Research Paper |
Corresponding author: Jürgen Dengler ( dr.juergen.dengler@gmail.com ) Academic editor: Victor Chepinoga
© 2024 Denys Vynokurov, Alla Aleksanyan, Thomas Becker, Idoia Biurrun, Dariia Borovyk, George Fayvush, Itziar García-Mijangos, Martin Magnes, Salza Palpurina, Ute Becker, Asun Berastegi, Beata Cykowska-Marzencka, Iwona Dembicz, Dieter Frank, Andreas Hilpold, Philipp Kirschner, Helmut Mayrhofer, Marine Oganesian, Iuliia Vasheniak, Jürgen Dengler.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Vynokurov D, Aleksanyan A, Becker T, Biurrun I, Borovyk D, Fayvush G, García-Mijangos I, Magnes M, Palpurina S, Becker U, Berastegi A, Cykowska-Marzencka B, Dembicz I, Frank D, Hilpold A, Kirschner P, Mayrhofer H, Oganesian M, Vasheniak I, Dengler J (2024) Dry grasslands and thorn-cushion communities of Armenia: a first syntaxonomic classification. Vegetation Classification and Survey 5: 39-73. https://doi.org/10.3897/VCS.119253
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Aim: To provide the first syntaxonomic, plot-based classification of the dry grasslands and thorn-cushion communities in Armenia. Study area: Armenia. Methods: We sampled 111 vegetation plots (10 m2) and recorded environmental and structural parameters. We collected additional 487 relevés from surrounding countries for a broad-scale comparison. We used modified TWINSPAN to derive a syntaxonomic classification system, whose units were then compared among each other regarding species composition, structure, site conditions and distribution. Results: The classification of Armenian vegetation plots resulted in a 12-cluster solution. Unsupervised classification of the broad-scale dataset yielded five main groups, which were used for the high-level syntaxonomic assignments of the Armenian data. We assigned about half of the plots of the Armenian dataset to the Festuco-Brometea, while the remaining represented a potential new class, preliminarily called “Ziziphora tenuior-Stipa arabica grasslands”. Most of the syntaxa below class level are new to science, therefore we provide formal descriptions of three orders (Plantagini atratae-Bromopsietalia variegatae, Onobrychido transcaucasicae-Stipetalia pulcherrimae, Cousinio brachypterae-Stipetalia arabicae), four alliances (Acantholimono caryophyllacei-Stipion holosericeae, Artemision fragrantis, Onobrychido michauxii-Stipion capillatae, Onobrychido transcaucasicae-Stipion pulcherrimae) and six associations. We found significant differences in the topographic, climatic and soil characteristics, and structural parameters, species life forms and distribution range types between the grassland types at different syntaxonomic levels. The mean species richness was 47.3 (vascular plants: 46.8, bryophytes: 0.4, lichens: 0.1). Conclusions: We found remarkable differences of the Armenian dry grasslands from the previously known units and described most of the higher syntaxa and all the associations as new to science. Our study provides arguments for a potential new class of Ziziphora tenuior-Stipa arabica grasslands separate both from the Euro-Siberian Festuco-Brometea and the Anatolian Astragalo-Brometea. Finally, we found plot scale richness of vascular plants clearly above the Palaearctic average of dry grasslands and that of non-vascular plants clearly below, which calls for further biodiversity analyses.
Taxonomic reference:
Abbreviations: EDGG = Eurasian Dry Grassland Group; DCA = detrended correspondence analysis; ICPN = International Code of Phytosociological Nomenclature (
Armenia, classification, dry grassland, Festuco-Brometea, Irano-Turanian, mountain steppe, Onobrychido transcaucasicae-Stipetalia pulcherrimae, Plantagini atratae-Bromopsietalia variegatae, species richness, syntaxonomy, thorn-cushion community, Cousinio brachypterae-Stipetalia arabicae
Armenia is a land-lock country located in the southern part of the Lesser Caucasus, belonging to two global biodiversity hotspots: the Caucasian and the Irano-Anatolian (
Grasslands and shrublands in the Middle East and the Caucasus areas are of great interest as they host a high biodiversity of species and habitats (
Due to the abovementioned importance of the typology of habitats and vegetation, there is a growing international consensus on the need for coherent vegetation classification systems based on the analysis of vegetation-plot data (
Up to date, there is no formalised plot-based classification system for Armenia, which was a Soviet Socialist Republic until 1991. As the Braun-Blanquet approach to vegetation classification (
Although the phytosociological approach has not been applied to the survey of Armenian grasslands, those of neighbouring countries of the Southern Caucasus have been at least fragmentarily studied in Transcaucasia (Azerbaijan:
Many of the data used for these phytosociological surveys are stored in vegetation-plot databases. In the last decades small regional and/or personal databases have been compiled in large vegetation plots databases. The European Vegetation Archive (EVA,
It is therefore both an opportunity and a challenge to record vegetation plots and apply the Braun-Blanquet classification approach to Armenia – as its vegetation, to our knowledge, has never been studied according to this approach. Therefore, the Eurasian Dry Grassland Group (EDGG; www.edgg.org) conducted an international research expedition (called “Field Workshop”) in Armenia to collect standardised, high-quality vegetation-plot data from dry grasslands and thorn-cushion communities throughout the country. In principle the EDGG Field Workshops aim at collecting such data for regional studies on biodiversity patterns (
In this paper, we used the plot data sampled during the Field Workshop to provide the first syntaxonomic classification scheme of the dry grasslands and thorn-cushion communities of the country, using numerical methods of unsupervised classification and determination of diagnostic species. Specifically, we asked:
Armenia is a South Caucasian republic, bordering Georgia, Azerbaijan, Turkey, and Iran. It is a landlocked country with a total area of 29,740 km2, at about 145 km from the Black Sea and 175 km from the Caspian Sea. It lies between 38°50' and 41°18' northern latitude and between 43°27' and 46°37' eastern longitude, and measures 400 km along its main axis (north-west to south-east). Armenia is generally a mountainous country, having its lowest point at 375 m a.s.l. and culminating at 4,095 m a.s.l. in the Aragats, with an average elevation of 1,850 m a.s.l.
The location of Armenia at the intersection of two phytogeographical subkingdoms (Boreal and Ancient Mediterranean), together with the diversity in climatic conditions and the active geological processes, have resulted in the formation of diverse ecosystems and high biodiversity with a high level of endemism (
A wide range of climatic zones are distinguished within Armenia, which experiences large climatic contrasts because of its intricate terrain and the big climatic gradients (
From the orographical and physico-geographical points of view, Armenia forms the northern edge of the system of folded-block mountains of the Armenian Highland. Unlike the Greater Caucasus, Armenia and the Lesser Caucasus are not a single, distinct watershed ridge. It is a system of coulisse-spaced ridges that merge with the mountain formations of the inner parts of the Armenian Highland and adjacent high areas (
In our study we tried to cover as much of the country’s dry grassland diversity as possible within 11 days, with a focus on the northwestern and central parts (Figure
So far, the syntaxonomy of grassland or thorn-cushion vegetation of Armenia hasn’t been developed yet. The only existing vegetation typologies are based on the dominance approach. The first overview of the Armenian vegetation types was performed by
Afterwards, the classification of natural grasslands of Armenia was done by
In the Transcaucasus region, the first work with a description of syntaxa following the Braun-Blanquet approach was done by M. Guinochet in Azerbaijan and Georgia.
For the Northern Caucasus,
In Turkey, several high-level syntaxonomic units have been established for dry grasslands and thorn-cushion vegetation.
In another bordering region, Iran, xerophilous grassland and scrub communities were first delineated by
In general, the class Astragalo-Brometea is the most widely used name to unite the dry grasslands and thorn-cushion vegetation in the western part of the Irano-Turanian region. However, its conceptual boundaries have undergone significant changes over time, both geographically and physiognomically. Many researchers now extend its scope to include tragacanth vegetation not only from the subalpine belt but also from the lower elevations, as well as chamaephyte-dominated phryganoid vegetation, non-tragacanth dry grasslands at lower elevations, saline steppes, and gypsophilous rocky grasslands (
We sampled 111 plots of 10 m2- size (Suppl. materials
Other environmental and structural parameters that were recorded in situ following the EDGG sampling methodology (
Soil was collected as mixed samples from the uppermost 15 cm of the soil in five random points inside each plot. After air drying and sieving to the fine-soil fraction, the following parameters were measured in the lab: pH (in H2O), electrical conductivity (μS cm-1), organic C content (%), humus content (%), N content (%), and C/N ratio. Southing was calculated from aspect as -cos (aspect).
The nomenclature of vascular plants was standardised to
To be able to identify the high-level syntaxonomic units, we digitised from literature and used for comparison relevés from the bordering regions, focusing on the original diagnosis of the high-level units of similar vegetation types: 230 plots from Anatolia (Turkey) and 51 plots from Northern Iran. Among them, the original diagnosis of the class Astragalo-Brometea
Climatic data were extracted from the CHELSA Climate database (
Maps were created using QGIS software (
In order to assess the distribution ranges of species, we analysed their distribution maps (according to
In addition, we classified all the species into one of the Raunkiær plant life forms: therophytes, geophytes, hemicryptophytes, chamaephytes, and phanerophytes. The data are also available in the Suppl. material
Unsupervised classification for both the West Asian and Armenian dataset was done in JUICE 7.0 (
DCA-Ordination was performed with Canoco 5 (
Differences in variables between syntaxa were tested by univariate ANOVA using SPSS 22 (IBM, Armonk, NY, US). We tested whether the assumptions of ANOVA (normal distribution, equal variance) were sufficiently met by visually inspecting the frequency distribution of the residuals and by testing for homogeneity of variance according to Levene (
We selected a TWINSPAN resolution where the terminal clusters were floristically still well-characterised and not too small. These clusters were then assigned to the rank of association. Alliance, order and class levels were assigned to higher cut levels of the dendrogram, with the double aim to have floristically well differentiated and ecologically and chorologically interpretable units on a comparable level as these hierarchies have in
After defining the hierarchical units, we carefully checked the syntaxonomic literature of the neighbouring countries to determine whether syntaxa with this content already existed. If this was the case, we took the established name. If not, we formally described new syntaxa according to the ICPN (
Modified TWINSPAN resulted into 16 clusters with five main groups of clusters: A (clusters 1–3), B (4–6), C (7), D (8–10) and E (11–16) (Figure
The group A (clusters 1–3) completely consisted of the relevés originally assigned to the class Astragalo-Brometea, including the type order Astragalo-Brometalia
The second group B (clusters 4–6) comprised plots from the high-mountain steppe vegetation. Cluster 4 consisted mainly of relevés from the northern slope of the Alborz Mountains in Iran, assigned to the association Alchemilletum plicatissimae Klein et Lacoste 1994. Plots from the Northern Caucasus region assigned to the alliance Artemisio chamaemelifoliae-Bromopsion variegatae Vynokurov in
Thorn-cushion communities from Eastern Anatolia are combined in the group C (cluster 7). They were originally assigned to the order Festuco oreophilae-Veronicetalia orientalis
Clusters 8–10 formed group D. It consisted exclusively of plots from Armenia. Among them, cluster 8 contained the most xeric communities sampled in the driest parts of Armenia, followed by cluster 10. Cluster 9 was transitional between the groups D and E.
The group E (clusters 11–16) was formed by plots containing the more ‘typical’ Festuco-Brometea species, mostly from the region of the Northern Caucasus. Cluster 11 was comprised mainly of rocky grasslands belonging to the order Asphodelino tauricae-Euphorbietalia petrophilae Vynokurov in
Results of the Modified TWINSPAN classification for the broad-scale comparison involving plots from the bordering countries (n = 598). The width of the bars is proportional to the number of included plots. The main groups (letters) and terminal clusters (numbers) are described in the text. Blue colour indicates clusters that predominantly or completely consisted of Armenian plots.
DCA of the West Asian dataset (DCA with supplementary variables, eigenvalues/gradient lengths/cumulative explained variation of axis 1: 0.6313/5,37/4.02, axis 2: 0.4207/4.94/6.69). Vectors (environmental variables): BIO1: annual mean temperature; BIO7: temperature annual range; BIO12: annual precipitation; elevation: elevation (m a.s.l.); N species: vascular plant richness.
In our 111 plots of 10 m2, we recorded a total of 739 vascular plant, 40 bryophyte and 13 lichen taxa (subspecies, species, aggregates and sections, further as ‘species’). The species richness per plot ranged from 21 to 85, with a mean of 47.3. On average there were 46.8 vascular plant, 0.4 bryophyte and 0.1 lichen species per plot. The most frequent vascular plant was Galium verum (in 72% of all plots), followed by Thymus kotschyanus (59%), Teucrium capitatum (58%), Poa bulbosa (55%), Dactylis glomerata (54%), Scutellaria orientalis aggr. (53%), Koeleria macrantha (51%), Stachys recta (50%) and Potentilla recta aggr. (50%). The most frequent bryophytes were Syntrichia ruralis (27%), Ptychostomum imbricatulum (19%) and Syntrichia montana (14%). Lichens were absent in most plots, with the most frequent one (Cladonia foliacea) reaching just 4%.
The most meaningful modified TWINSPAN classification of the plots from Armenia resulted in the 12-cluster solution (Figure
Results of the Modified TWINSPAN classification of the Armenian plots (n = 111). The terminal clusters were interpreted as associations or, if represented by too few plots, as informal communities at association rank. The first cluster (X) consisted only of one plot of scree vegetation. Codes at the tips of the other clusters correspond to Table
Ordination of the plots with the assignment to one of these clusters revealed that the first axis of the DCA graph (Figure
DCA ordination of the 111 Armenian plots with assignment to the 12 distinguished units at association level (DCA with supplementary variables, eigenvalues/gradient lengths/cumulative explained variation of axis 1: 0.5825/5.12/5.76, axis 2: 0.3160/3.64/8.89). Triangles indicate members of the class Ziziphora tenuior-Stipa arabica grasslands, circles members of the class Festuco-Brometea. Vectors: BIO1: annual mean temperature; BIO12: annual precipitation; BIO17: precipitation of driest quarter; CAU: cover of Caucasian species in %; chamaephytes: cover of chamaephytes in %; cover herb layer: cover of the herb layer in %; cover litter: cover of the litter; elevation: elevation in m a.s.l.; EUR: cover of European species in %; height herb layer: height of the herb layer; humus: soil humus content in %; IT: cover of Irano-Turanian species in %; MED: cover of Mediterranean species in %; pH: pH values of the plot soil samples; soil depth: mean soil depth of plot; southerness: -cos (aspect); therophyte: cover of therophytes in %.
Resulting from our analyses of the Armenian data and the comparison with the syntaxa of neighbouring territories, we propose the following syntaxonomic scheme for the dry grassland and thorn-cushion communities of Armenia, including a single plot with an unclear assignment (Table
Syntaxonomic scheme for the dry grasslands and thorn-cushion communities of Armenia based on the 111 plots analysed in this paper.
Unclear class (scree communities) |
Euphorbia orientalis-Melilotus officinalis community |
Potential class 1 Ziziphora tenuior-Stipa arabica grasslands |
Order 1.1 Cousinio brachypterae-Stipetalia arabicae Vynokurov et al. 2024 |
Alliance 1.1.1 Onobrychido michauxii-Stipion capillatae Vynokurov et al. 2024 |
1.1.1.1 Stachys lavandulifolia-Astracantha condensata community |
1.1.1.2 Marrubio parviflorae-Stipetum capillatae Vynokurov et al. 2024 |
Alliance 1.1.2 Artemision fragrantis Vynokurov et al. 2024 |
1.1.2.1 Noaeo mucronatae-Artemisietum fragrantis Vynokurov et al. 2024 |
Alliance 1.1.3 Acantholimono caryophyllacei-Stipion holosericeae Vynokurov et al. 2024 |
1.1.3.1 Acantholimono caryophyllacei-Stipetum holosericeae Vynokurov et al. 2024 |
1.1.3.2 Stachys inflata-Acantholimon vedicum community |
Class 2. Festuco-Brometea Br.-Bl. et Tx. ex Soó 1947 |
Order 2.1 Plantagini atratae-Bromopsietalia variegatae Vynokurov et al. 2024 |
Alliance 2.1.1 Artemisio chamaemelifoliae-Bromopsion variegatae Vynokurov in |
2.1.1.1 Ranunculo caucasici-Bromopsietum variegatae Vynokurov et al. 2024 |
2.1.1.2 Tragopogon reticulatus-Astracantha aurea community |
Order 2.2 Onobrychido transcaucasicae-Stipetalia pulcherrimae Vynokurov et al. 2024 |
Alliance 2.2.1. Onobrychido transcaucasicae-Stipion pulcherrimae Vynokurov et al. 2024 |
2.2.1.1 Trisetum flavescens-Stachys macrostachys community |
2.2.1.2 Onobrychis transcaucasica-Vicia canescens subsp. variegata community |
2.2.1.3 Globulario trichosanthae-Stipetum pulcherrimae Vynokurov et al. 2024 |
2.2.1.4 Seslerio phleoidis-Onobrychidetum cornutae Vynokurov et al. 2024 |
The proposed classification of the Armenian dry grassland and thorn-cushion communities is shown in the synoptic table (abbreviated version: Table
Euphorbia orientalis-Melilotus officinalis scree community (Figure
One cluster in our analysis consisted of only a single relevé of scree vegetation. For this instance, we assume that a corresponding vegetation type needs to be described in the future in the rank of an order or even a class when enough relevant data is available. The aforementioned relevé was sampled in the Vayots Dzor Province, near Hermon (39.8812°N, 45.43254°E), 1,739 m a.s.l., aspect 135°, inclination 46°, 2 July 2019, total vegetation cover: 50%:
Alyssum alyssoides: 0.5, Arenaria serpyllifolia aggr.: 0.1, Asperula arvensis: 0.2, Buglossoides arvensis: 0.1, Bupleurum commutatum: 0.01, Caucalis platycarpos: 0.1, Cerastium ruderale: 3, Chaerophyllum bulbosum: 0.5, Cleome ornithopodioides: 0.01, Convolvulus arvensis: 0.1, Coronilla coronata: 2, Crepis pulchra: 2, Euphorbia orientalis: 15, Galium spurium: 1, Galium tenuissimum: 0.3, Holosteum marginatum: 0.1, Lactuca viminea: 0.5, Lamium amplexicaule: 0.01, Medicago rigidula: 0.1, Melica taurica: 1, Melilotus officinalis: 5, Michauxia laevigata: 0.5, Nepeta trautvetteri: 0.3, Noccaea perfoliata: 0.01, Prangos ferulacea: 1, Reichardia dichotoma: 0.3, Salvia verticillata: 30, Sanguisorba minor: 2, Saponaria orientalis: 0.2, Secale vavilovii: 2, Stachys recta: 0.5, Valerianella uncinata: 0.1, Vicia sativa: 0.3, Zosima absinthiifolia: 4.
Potential class 1: Ziziphora tenuior-Stipa arabica grasslands – Western Asian dry grasslands and xeric thorn cushion communities
Diagnostic species: Achillea arabica, Aegilops cylindrica, Agropyron cristatum, Allium pseudoflavum, Alyssum turkestanicum, Androsace maxima, Anisantha tectorum, Arabis auriculata aggr., Arenaria serpyllifolia aggr., Artemisia fragrans, Asperula arvensis, Bromus danthoniae, B. japonicus, B. squarrosus, Centaurea aggregata, Chardinia orientalis, Crepis sancta, Crupina vulgaris, Dianthus orientalis, Helianthemum ledifolium, Helichrysum plicatum, Holosteum marginatum, H. umbellatum, Marrubium parviflorum, Meniocus linifolius, Minuartia hamata, Noaea mucronata, Poa bulbosa, Sideritis montana, Stachys inflata, S. lavandulifolia, Stipa arabica, S. capillata, S. holosericea, Taeniatherum caput-medusae subsp. crinitum, Tanacetum aureum, Teucrium capitatum, Thymelaea passerina, Xeranthemum squarrosum, Ziziphora tenuior.
Ecology and distribution. Communities of the potential new class occur in the lower elevations in dry conditions and include semi-desert vegetation, xeric thorn-cushion communities, and xeric grasslands. Within Armenia, it is represented by one order and three alliances.
Order 1.1: Cousinio brachypterae-Stipetalia arabicae – Western Asian dry grasslands and xeric thorn cushion communities
Diagnostic species: Identical with those of the monotypic class.
Ecology and distribution. Communities of this order are distributed in Transcaucasia and possibly even broader within Western Asia. We expect them to occur throughout Western Asia in dry conditions (semi-deserts, dry steppe-like communities, low-elevation thorn-cushion communities). Also, according to our analysis, communities of this order may occur even in higher elevations on rocky substrates.
Alliance 1.1.1: Onobrychido michauxii-Stipion capillatae – Transcaucasian rocky dry grasslands
Diagnostic species: Astracantha condensata, Onobrychis michauxii, Salvia aethiopis, Stachys lavandulifolia, Teucrium capitatum, Veronica multifida (mainly negatively differentiated central alliance).
Ecology and distribution. Communities of this alliance are distributed in higher elevations than those of the other two alliances included in this order. This alliance is a transitional unit between this order and the order Onobrychido transcaucasicae-Stipetalia pulcherrimae, comprising Transcaucasian mountain steppes (see below).
1.1.1.1: Stachys lavandulifolia-Astracantha condensata community (Figure
Diagnostic species: Asperula arvensis, Astracantha condensata, Centaurea phrygia subsp. abbreviata, Crepis ciliata, Euphorbia orientalis, Gypsophila elegans, Herniaria hirsuta, Leptunis trichodes, Melica taurica, Nepeta racemosa, Onobrychis michauxii, Onosma setosa, Salvia aethiopis, Sempervivum transcaucasicum, Stachys lavandulifolia, Tanacetum aureum, Teucrium orientale, Tragopogon dubius, Viola occulta, Zosima absinthiifolia.
Structure, ecology and distribution. We sampled this vegetation type in the Gegharkunik (vicinity of the town of Sevan, Shorja) and Vayots Dzor (Hermon, vicinity of Gnishik and Khachik) provinces. These communities were located at the most south-facing rocky slopes with shallow soil and low humus content. The herb layer was sparse and with a high representation of Irano-Turanian species, e.g. Astracantha condensata, A. microcephala, Stachys lavandulifolia, Teucrium orientale.
1.1.1.2: Marrubio parviflorae-Stipetum capillatae (Figure
Diagnostic species: Allium cardiostemon, Centaurea ovina aggr., Euphorbia condylocarpa, Marrubium parviflorum, Stipa capillata.
Structure, ecology and distribution: These communities were sampled on slopes with shallow rocky substrates in Gegharkunik (Ardanish), Lori (near Shirakamut) and Vayots Dzor (vicinity of Gnishik and Khachik) provinces. The association differed by a higher herb layer cover compared to the previous community and the highest participation of hemicryptophytes among all associations of the class. The dominant species were Festuca valesiaca aggr., Onobrychis cornuta and Teucrium capitatum.
Alliance 1.1.2: Artemision fragrantis – Transcaucasian wormwood semi-deserts
Diagnostic species: Agropyron cristatum, Allium pseudoflavum, Alyssum turkestanicum, Androsace albana, Arenaria serpyllifolia aggr., Artemisia fragrans, Astragalus hyalolepis, Bromopsis riparia, Ceratocephala falcata, Cousinia brachyptera, Crupina vulgaris, Cuscuta araratica, Consolida hispanica, Didymodon tophaceus (B), Meniocus linifolius, Minuartia hamata, Noaea mucronata, Peganum harmala, Polygala hohenackeriana, Sclerocaryopsis spinocarpos, Syntrichia caninervis (B).
Ecology and distribution: Artemisia fragrans semi-deserts in Armenia are distributed in the lowest elevations in the country. We did not sample other semi-desert types, but we can expect that Armenian loamy and sandy semi-deserts will also be probably included in this unit. In our dataset, this alliance is represented by a single association.
1.1.2.1: Noaeo mucronatae-Artemisietum fragrantis (Figure
Diagnostic species: identical with those of the monotypic alliance.
Structure, ecology and distribution: This association is typical for the Aragatsotn province (vicinity of Dashtadem and Tatool). The sampled plots were distributed at the lowest elevations with the highest mean annual temperature and lowest mean annual precipitation compared to the other studied associations. The communities were dominated by Artemisia fragrans, Poa bulbosa, and Taeniatherum caput-medusae subsp. crinitum. The herb layer is relatively sparse and with a high representation of therophytes (e.g. Alyssum turkestanicum, Bromus squarrosus, Ceratocephala falcata, Crupina vulgaris, Sclerocaryopsis spinocarpos) and characteristic chamaephytes (Artemisia fragrans, Noaea mucronata).
Alliance 1.1.3: Acantholimono caryophyllacei-Stipion holosericeae – Transcaucasian dry grasslands and xeric thorn-cushion communities
Diagnostic species: Acantholimon vedicum, Aegilops cylindrica, Aethionema carneum, Arabis auriculata aggr., Bromus japonicus, Bunium microcarpum, Chardinia orientalis, Crepis sancta, Crupina vulgaris, Ephedra procera, Galium verticillatum, Gaudiniopsis macra, Helianthemum ledifolium, Noccaea perfoliata, Papaver minus, Roemeria hybrida, Stachys inflata, Stipa arabica, S. holosericea, Taeniatherum caput-medusae subsp. crinitum, Petrorhagia cretica, Ziziphora tenuior.
Ecology and distribution: This unit comprises vegetation traditionally known as ‘highland xerophytic vegetation’. It includes dry grasslands and xeric tragacanth communities distributed above the semi-desert belt and below the mountain steppe altitudinal belt. We distinguish one association and one community within this alliance.
1.1.3.1: Acantholimono caryophyllacei-Stipetum holosericeae (Figure
Diagnostic species: Acantholimon caryophyllaceum, Aegilops triuncialis, Carduus hamulosus, Crepis sancta, Geranium lucidum, Lomelosia rotata, Noccaea annua, Stipa zalesskii subsp. pontica, Torilis arvensis, Xeranthemum squarrosum.
Structure, ecology and distribution: We sampled this association mainly in the Vayots Dzor province (Hermon, vicinities of Areni, Gnishik and Khachik), and also in one locality in Aragatsotn province (near Tatool). The communities were distributed on shallow soils, but with higher humus content and lower gravel cover compared to the other associations of this class. Acantholimon caryophyllaceum, Stipa holosericea, and Taeniatherum caput-medusae subsp. crinitum are the dominant species in this association. Among other species, Irano-Turanian elements often occur, such as Achillea arabica, Eryngium billardierei, Hypericum scabrum, and Thymus kotschyanus.
1.1.3.2: Stachys inflata-Acantholimon vedicum community (Figure
Diagnostic species: Acantholimon vedicum, Aegilops biuncialis, A. cylindrica, Aethionema carneum, Androsace maxima, Arabis auriculata aggr., Artemisia fragrans, Aspicilia hispida (L), Astragalus ornithopodioides, Bunium microcarpum, Callipeltis cucullaria, Camelina laxa, Chardinia orientalis, Cousinia daralaghezica, Crossidium squamiferum (B), Crucianella exasperata, Crupina vulgaris, Cuscuta pedicellata, Ephedra procera, Galium verticillatum, Gaudiniopsis macra, Helianthemum ledifolium, Holosteum marginatum, Lactuca tuberosa, Lamium amplexicaule, Onobrychis atropatana, Papaver minus, Petrorhagia cretica, Stachys inflata, Stipa arabica, Tanacetum aureum, Trinia glauca, Valerianella coronata, Ziziphora tenuior.
Structure, ecology and distribution: We sampled this vegetation type in Ararat (vicinity of Tigranashen) and Vayots Dzor (vicinity of Gnishik) provinces. Communities were distributed at lower elevations with high mean annual temperature and low mean annual precipitation. The substrate differed by the most alkaline soil reaction (mean pH: 8). The herb layer was sparse and with a high representation of Irano-Turanian and Mediterranean therophyte species (Aegilops spp., Crupina vulgaris, Petrorhagia cretica, Ziziphora tenuior), while the cover of hemicryptophytes was the lowest among all studied communities. These communities did not have clear dominants, but Chardinia orientalis, Stachys inflata, Stipa arabica, and S. sareptana subsp. anisotricha occurred with higher cover than the other species. The species richness of vascular plants, bryophytes and lichens was higher compared to the other associations of the class.
Class 2: Festuco-Brometea – Mesoxeric and xeric basiphilous grasslands of temperate Europe and adjacent regions
Diagnostic species: Abietinella abietina (B), Achillea millefolium aggr., Artemisia absinthium, Bupleurum falcatum aggr., Campanula glomerata aggr., C. rapunculoides, C. stevenii, Cirsium leucocephalum, Dactylis glomerata, Galium verum, Koeleria albovii, Leontodon hispidus, Linum nervosum, L. tenuifolium, Lotus corniculatus, Onobrychis transcaucasica, Origanum vulgare, Phleum phleoides, Pimpinella saxifraga aggr., Plantago atrata, Poa pratensis aggr., Polygala anatolica, Polygonum cognatum, Potentilla recta aggr., Scabiosa bipinnata, Securigera varia, Stachys macrostachys, Taraxacum sect. Taraxacum, Thalictrum minus, Trifolium alpestre, T. ambiguum, T. trichocephalum, Trisetum flavescens.
Ecology and distribution: Within Armenia, this class comprises meso-xeric grasslands and mountain steppes at higher elevations. We distinguish two orders representing different altitudinal belts.
Order 2.1: Plantagini atratae-Bromopsietalia variegatae – High-mountain meso-xeric grasslands of the Caucasus
Diagnostic species: Achillea millefolium aggr., Ajuga orientalis, Alchemilla sericea, Arenaria blepharophylla aggr., A. gypsophiloides, Aster alpinus, Brachypodium pinnatum, Bromopsis variegata, Campanula collina, C. stevenii, Cirsium leucocephalum, Festuca ovina aggr., Schedonorus pratensis, Festuca rubra aggr., Filipendula vulgaris, Gagea glacialis, Galium cordatum, Gentiana septemfida, Huynhia pulchra, Koeleria albovii, Lathyrus digitatus, Lomelosia caucasica, Lotus corniculatus, Luzula multiflora, Medicago papillosa, Muscari armeniacum, Myosotis alpestris, Ornithogalum sigmoideum, Papaver orientale, Pedicularis condensata, Phleum alpinum, Pilosella officinarum aggr., Pimpinella saxifraga aggr., Plantago atrata, Poa pratensis aggr., Pohlia nutans (B), Polygonum cognatum, Potentilla argentea, Psephellus xanthocephalus, Pseudoleskella tectorum (B), Pulsatilla albana, Ranunculus caucasicus, Rumex acetosella, Senecio pseudo-orientalis, Stachys macrantha, Stipa tirsa, Taraxacum sect. Taraxacum, Thymus collinus, Tortula acaulon (B), Trifolium ambiguum, T. spadiceum, T. trichocephalum, Verbascum speciosum, Veronica denudata, V. gentianoides.
Ecology and distribution: Communities of this order occupy the highest sampled elevations in Armenia: upper subalpine and lower alpine belts. They form a particular unit recognized in the dominant approach typology: mountain meadow steppes. Beyond this elevation belt, they are replaced by the alpine grasslands which possibly belong to the class Juncetea trifidi Hadač in Klika et Hadač 1944 (Festucetalia woronowii
Alliance 2.1.1: Artemisio chamaemelifoliae-Bromopsion variegatae – Caucasian subalpine and lower-alpine meso-xeric grasslands
Diagnostic species: Achillea millefolium aggr., Alchemilla sericea, Arenaria gypsophiloides, Bromopsis variegata, Campanula collina, C. stevenii, Festuca ovina aggr., F. rubra aggr., Huynhia pulchra, Koeleria albovii, Lathyrus digitatus, Lomelosia caucasica, Lotus corniculatus, Myosotis alpestris, Plantago atrata, Polygonum cognatum, Potentilla argentea, Ranunculus caucasicus, Taraxacum sect. Taraxacum, Trifolium ambiguum, T. trichocephalum, Veronica denudata, V. gentianoides.
Ecology and distribution: This unit was described from the Main Range of the North Caucasus (
2.1.1.1: Ranunculo caucasici-Bromopsietum variegatae (Figure
Diagnostic species: Alchemilla sericea, Arenaria blepharophylla aggr., Artemisia chamaemelifolia, Aster alpinus, Avenula pubescens, Brachypodium pinnatum, Bromopsis variegata, Campanula collina, C. stevenii, Carex caryophyllea, Cirsium leucocephalum, Schedonorus pratensis, Galium cordatum, Huynhia pulchra, Lomelosia caucasica, Luzula multiflora, Medicago papillosa, Muscari armeniacum, Myosotis alpestris, Pedicularis condensata, Phascum cuspidatum (B), Phleum alpinum, Pilosella officinarum aggr., Pimpinella saxifraga aggr., Plantago atrata, Polygonum cognatum, Psephellus xanthocephalus, Pulsatilla albana, Ranunculus caucasicus, Rumex acetosella, Stachys macrantha, Stipa tirsa, Tragopogon graminifolius, Trifolium trichocephalum, Veronica gentianoides.
Structure, ecology and distribution: We sampled this vegetation type at the steep north-facing slopes at elevations around 2,100 m, mainly in the Shirak province (vicinities of Amasia and Zorakert), and also in Gegharkunik province (Ardanish). The localities were characterised by a high mean annual precipitation (around 700–900 mm). The soil reaction was slightly acidic (mean pH: 6.5). The association differed by high species richness and the highest participation of Caucasian species, e.g. Dianthus cretaceus, Lomelosia caucasica, Stachys macrantha, and Trifolium trichocephalum, among all studied communities. Graminoids were dominant, particularly Brachypodium pinnatum, Bromopsis variegata, Carex humilis, and Phleum alpinum.
2.1.1.2: Tragopogon reticulatus-Astracantha aurea community (Figure
Diagnostic species: Arenaria dianthoides, A. gypsophiloides, Astracantha aurea, Campanula stevenii, Elytrigia repens, Gagea glacialis, Koeleria albovii, Lathyrus digitatus, Papaver orientale, Plantago atrata, Senecio pseudo-orientalis, Tragopogon reticulatus, Trifolium ambiguum, T. spadiceum, Trisetum flavescens, Verbascum speciosum.
Structure, ecology and distribution: We sampled this community mainly in the Gegharkunik province (Selim pass), and also in the Aragatsotn province (near the fortress of Amberd). Most of the localities were situated at 2,300–2,400 m a.s.l. and represented the highest elevations among all studied sites. The soil reaction was slightly acidic. These communities are characterised by low species richness with a high participation of Transcaucasian and Caucasian species, e.g. Arenaria dianthoides, Astracantha aurea, Koeleria albovii. Festuca ovina aggr. and Plantago atrata were the dominant species.
Order 2.2: Onobrychido transcaucasicae-Stipetalia pulcherrimae – Transcaucasian mountain steppes
Diagnostic species: Artemisia absinthium, Bupleurum falcatum aggr., Campanula glomerata aggr., C. rapunculoides, Cerinthe minor, Dactylis glomerata, Euphrasia pectinata, Galium verum, Globularia trichosantha, Helictochloa armeniaca, Hypericum perforatum, Klasea radiata, Linum nervosum, L. tenuifolium, Lotus corniculatus, Nepeta nuda, Onobrychis transcaucasica, Origanum vulgare, Phlomis tuberosa, Polygala anatolica, Rosa spinosissima, Salvia verticillata, Scabiosa bipinnata, Securigera varia, Stachys macrostachys, S. recta, Stipa pennata, S. pulcherrima, Teucrium chamaedrys, Thalictrum minus, Vicia canescens subsp. variegata, Viola ambigua.
Ecology and distribution: Mountain steppes in the Transcaucasus form a distinct altitudinal belt above highland xerophyte vegetation (Cousinio brachypterae-Stipetalia arabicae) and below mountain meadow steppes (Plantagini atratae-Bromopsietalia variegatae). In our dataset, the order is represented by one alliance.
Alliance 2.2.1: Onobrychido transcaucasicae-Stipion pulcherrimae – Transcaucasian mountain steppes
Diagnostic species: Campanula rapunculoides, Cerinthe minor, Dactylis glomerata, Linum nervosum, L. tenuifolium, Onobrychis transcaucasica, Origanum vulgare, Scabiosa bipinnata, Securigera varia, Stachys macrostachys, S. recta.
Ecology and distribution: Despite its high floristic heterogeneity, we unite all mountain steppes into one alliance. We distinguish two informal communities and two associations.
2.2.1.1: Trisetum flavescens-Stachys macrostachys community (Figure
Diagnostic species: Arenaria graminea, Artemisia absinthium, Chaerophyllum roseum, Gagea caroli-kochii, Silene cephalantha, Stachys macrostachys, Trisetum flavescens, Verbascum cheiranthifolium.
Structure, ecology and distribution: We sampled this vegetation type at the elevations 1,950–2,300 m a.s.l. in Aragatsotn (near Amberd fortress), Gegharkunik (Selim pass, Shorja) and Shirak (vicinity of Amasia) provinces. These communities develop on soils with high humus content. The herb layer is relatively dense with dominance of grasses (Elytrigia intermedia aggr., Phleum nodosum, Poa pratensis aggr., Trisetum flavescens) and legumes (Securigera varia, Trifolium alpestre, Vicia tenuifolia subsp. variabilis, V. canescens subsp. variegata).
2.2.1.2: Onobrychis transcaucasica-Vicia canescens subsp. variegata community (Figure
Diagnostic species: Arabis hirsuta, Campanula bononiensis, Campyliadelphus chrysophyllus (B), Chaerophyllum aureum, Daphne oleoides subsp. kurdica, Helictochloa armeniaca, Klasea radiata, Lathyrus latifolius, Linum nervosum, Nepeta nuda, Onobrychis transcaucasica, Origanum vulgare, Phlomis tuberosa, Primula veris subsp. macrocalyx, Rhinanthus subulatus, Salvia nemorosa, Securigera varia, Seseli libanotis, Stachys macrostachys, Stipa zalesskii subsp. canescens, Thalictrum minus, Valeriana officinalis aggr., Vicia canescens subsp. variegata, Viola ambigua.
Structure, ecology and distribution: We recorded relevés of this community only in the Vayots Dzor province, at north-facing slopes (inclination 10–40°) in the vicinity of Gnishik and between Khachik and Areni. Communities differed by closed herb layer and high litter cover. The species composition is characterised by a high participation of forbs with European distribution, such as Campanula bononiensis, Klasea radiata, and Securigera varia. The dominant species is Vicia canescens subsp. variegata.
2.2.1.3: Globulario trichosanthae-Stipetum pulcherrimae (Figure
Diagnostic species: central association (no diagnostic species)
Structure, ecology and distribution: We sampled this vegetation type at elevations around 1,900–2,200 m a.s.l. in the provinces of Gegharkunik (Ardanish and vicinity of the town of Sevan), Shirak (Jajur pass) and Vayots Dzor (vicinities of Gnishik, Khachik and Areni). In the species composition, prevailing groups of species were European (Potentilla recta aggr., Stachys recta, Stipa pulcherrima) and Irano-Turanian (Eryngium billardierei, Onobrychis michauxii, Thymus kotschyanus, Ziziphora clinopodioides), followed by Caucasian endemics (Astragalus cancellatus, Centaurea pseudoscabiosa, Scabiosa bipinnata). The dominant species of the association was Stipa pulcherrima.
2.2.1.4: Seslerio phleoidis-Onobrychidetum cornutae (Figure
Diagnostic species: Abietinella abietina (B), Adonis volgensis, Androsace chamaejasme, Asperula affinis, Asphodeline taurica, Briza media, Campanula rapunculoides, C. sibirica, Coronilla coronata, Euphorbia esula aggr., Euphrasia pectinata, Fritillaria caucasica, Homalothecium lutescens (B), Hypnum cupressiforme (B), Leucanthemum vulgare, Linum tenuifolium, Pimpinella saxifraga aggr., Pinus sylvestris, Pontechium maculatum, Psephellus karabaghensis, Sesleria phleoides, Spiraea crenata, Thalictrum foetidum, Viola alba.
Structure, ecology and distribution: We sampled this association at elevations 1,940–2,070 m a.s.l. in Gegharkunik (Ardanish, Shorja, vicinity of the town of Sevan) and Shirak (Jajur pass) provinces. The association differed by the highest mean total species richness and richness of bryophytes across all studied communities. The species composition is represented by a high participation of European species of grasses and forbs, such as Briza media, Campanula rapunculoides, Galium verum, Pimpinella saxifraga aggr., and Stachys recta. Carex humilis, Onobrychis cornuta and Teucrium chamaedrys are the dominant species.
Distribution of five described vegetation alliances in Armenia based on sampled vegetation plots (n = 110). 1.1.1 – Onobrychido michauxii-Stipion capillatae, 1.1.2 – Artemision fragrantis, 1.1.3 – Acantholimono caryophyllacei–Stipion holosericeae, 2.1.1 – Artemisio chamaemelifoliae–Bromopsion variegatae, 2.2.1 – Onobrychido transcaucasicae-Stipion pulcherrimae. Basemap: SRTM elevation model for Armenia (obtained from Earth Resources Observation and Science (
Abbreviated constancy table of the classes, orders, alliances and associations/communities distinguished among the Armenian dry grasslands and thorn-cushion communities. Dark grey indicates diagnostic species at the respective level, while this colour is copied to all subordinate levels. Light grey refers to transgressive diagnostic species at the higher level (only given if the sum of the phi values of the syntaxa at the lower level without the target syntaxon is > 0). Frames indicate differential species at the association level. Diagnostic species are sorted by decreasing phi values in their target unit, while companion species are sorted by decreasing total constancy. While all diagnostic species at the class level are shown, at the order, alliance and association level only the first seven plus all with a phi-value > 0.5 were kept. Additionally, all species with a total constancy of ≥ 20% were also kept. Species that did not meet any of these criteria are not shown here. The complete version of this table, including also the precise phi values and the individual relevés is available as Suppl. material
Syntaxon | All | 1 | 2 | 1.1 | 2.1 | 2.2 | 1.1.1 | 1.1.2 | 1.1.3 | 2.1.1 | 2.2.1 | 1.1.1.1 | 1.1.1.2 | 1.1.2.1 | 1.1.3.1 | 1.1.3.2 | 2.1.1.1 | 2.1.1.2 | 2.2.1.1 | 2.2.1.2 | 2.2.1.3 | 2.2.1.4 | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Number of plots | 110 | 47 | 63 | 47 | 21 | 42 | 17 | 13 | 17 | 21 | 42 | 7 | 10 | 13 | 10 | 7 | 13 | 8 | 8 | 7 | 16 | 11 | |
Mean species richness in 10 m2 (all) | 47.4 | 45.0 | 49.2 | 45.0 | 50.1 | 48.8 | 43.5 | 45.0 | 46.6 | 50.1 | 48.8 | 42.4 | 44.2 | 45.0 | 45.3 | 48.4 | 56.8 | 39.3 | 49.1 | 52.3 | 42.3 | 55.8 | |
Mean species richness in 10 m2 (vascular plants) | 46.9 | 44.6 | 48.7 | 44.6 | 49.3 | 48.3 | 43.2 | 44.9 | 45.9 | 49.3 | 48.3 | 41.9 | 44.1 | 44.9 | 44.6 | 47.7 | 55.6 | 39.1 | 48.9 | 51.4 | 41.9 | 55.3 | |
Mean species richness in 10 m2 (byophytes) | 0.4 | 0.2 | 0.5 | 0.2 | 0.6 | 0.4 | 0.3 | 0.1 | 0.3 | 0.6 | 0.4 | 0.6 | 0.1 | 0.1 | 0.3 | 0.3 | 0.9 | 0.0 | 0.3 | 0.7 | 0.3 | 0.5 | |
Mean species richness in 10 m2 (lichens) | 0.1 | 0.1 | 0.1 | 0.1 | 0.2 | 0.0 | 0.0 | 0.0 | 0.4 | 0.2 | 0.0 | 0.0 | 0.0 | 0.0 | 0.4 | 0.4 | 0.2 | 0.1 | 0.0 | 0.1 | 0.1 | 0.0 | |
Cl. 1 | |||||||||||||||||||||||
Xeranthemum squarrosum | 25 | 57 | . | 57 | . | . | 41 | 77 | 59 | . | . | 29 | 50 | 77 | 80 | 29 | . | . | . | . | . | . | |
Teucrium capitatum | 58 | 87 | 37 | 87 | . | 55 | 100 | 77 | 82 | . | 55 | 100 | 100 | 77 | 90 | 71 | . | . | . | 14 | 94 | 64 | |
Taeniatherum caput-medusae subsp. crinitum | 22 | 51 | . | 51 | . | . | 24 | 69 | 65 | . | . | 14 | 30 | 69 | 70 | 57 | . | . | . | . | . | . | |
Holosteum umbellatum | 24 | 51 | 3 | 51 | . | 5 | 41 | 54 | 59 | . | 5 | 14 | 60 | 54 | 50 | 71 | . | . | . | . | . | 18 | |
Crupina vulgaris | 18 | 40 | 2 | 40 | . | 2 | 6 | 69 | 53 | . | 2 | 14 | . | 69 | 40 | 71 | . | . | . | . | 6 | . | |
Thymelaea passerina | 15 | 34 | 2 | 34 | . | 2 | 35 | 38 | 29 | . | 2 | 14 | 50 | 38 | 20 | 43 | . | . | . | . | 6 | . | |
Tanacetum aureum | 23 | 40 | 10 | 40 | . | 14 | 47 | 31 | 41 | . | 14 | 86 | 20 | 31 | 10 | 86 | . | . | . | 14 | 6 | 36 | |
Helichrysum plicatum | 24 | 45 | 8 | 45 | 5 | 10 | 29 | 69 | 41 | 5 | 10 | 14 | 40 | 69 | 20 | 71 | . | 13 | 13 | 14 | 13 | . | |
Marrubium parviflorum | 15 | 32 | 2 | 32 | . | 2 | 41 | 23 | 29 | . | 2 | . | 70 | 23 | 50 | . | . | . | . | . | 6 | . | |
Bromus danthoniae | 10 | 23 | . | 23 | . | . | 18 | 15 | 35 | . | . | 14 | 20 | 15 | 30 | 43 | . | . | . | . | . | . | |
Bromus squarrosus | 15 | 30 | 3 | 30 | . | 5 | 12 | 38 | 41 | . | 5 | 14 | 10 | 38 | 50 | 29 | . | . | . | . | 6 | 9 | |
Sideritis montana | 22 | 40 | 8 | 40 | . | 12 | 53 | 69 | 6 | . | 12 | 71 | 40 | 69 | 10 | . | . | . | . | 14 | 13 | 18 | |
Poa bulbosa | 55 | 77 | 40 | 77 | 33 | 43 | 41 | 100 | 94 | 33 | 43 | . | 70 | 100 | 100 | 86 | 15 | 63 | 75 | . | 69 | 9 | |
Achillea arabica | 20 | 38 | 6 | 38 | 5 | 7 | 29 | 54 | 35 | 5 | 7 | . | 50 | 54 | 60 | . | . | 13 | 25 | . | 6 | . | |
Anisantha tectorum | 6 | 15 | . | 15 | . | . | 12 | . | 29 | . | . | 14 | 10 | . | 30 | 29 | . | . | . | . | . | . | |
Centaurea aggregata | 9 | 19 | 2 | 19 | . | 2 | 18 | 15 | 24 | . | 2 | 29 | 10 | 15 | 10 | 43 | . | . | 13 | . | . | . | |
Dianthus orientalis | 6 | 15 | . | 15 | . | . | 12 | 23 | 12 | . | . | 29 | . | 23 | . | 29 | . | . | . | . | . | . | |
All. 1.1.1 | |||||||||||||||||||||||
Veronica multifida | 23 | 26 | 21 | 26 | 19 | 21 | 65 | 8 | . | 19 | 21 | 57 | 70 | 8 | . | . | 31 | . | 13 | 29 | 25 | 18 | |
Assoc. 1.1.1.1 | |||||||||||||||||||||||
Astracantha condensata | 11 | 17 | 6 | 17 | . | 10 | 41 | 8 | . | . | 10 | 86 | 10 | 8 | . | . | . | . | . | . | 13 | 18 | |
Stachys lavandulifolia | 8 | 17 | 2 | 17 | . | 2 | 41 | . | 6 | . | 2 | 71 | 20 | . | 10 | . | . | . | . | . | 6 | . | |
Gypsophila elegans | 5 | 9 | 3 | 9 | . | 5 | 24 | . | . | . | 5 | 57 | . | . | . | . | . | . | . | 14 | . | 9 | |
Onobrychis michauxii | 11 | 17 | 6 | 17 | . | 10 | 41 | 8 | . | . | 10 | 71 | 20 | 8 | . | . | . | . | 13 | . | 19 | . | |
Viola occulta | 2 | 4 | . | 4 | . | . | 12 | . | . | . | . | 29 | . | . | . | . | . | . | . | . | . | . | |
Herniaria hirsuta | 2 | 4 | . | 4 | . | . | 12 | . | . | . | . | 29 | . | . | . | . | . | . | . | . | . | . | |
Crepis ciliata | 2 | 4 | . | 4 | . | . | 12 | . | . | . | . | 29 | . | . | . | . | . | . | . | . | . | . | |
Onosma setosa | 2 | 4 | . | 4 | . | . | 12 | . | . | . | . | 29 | . | . | . | . | . | . | . | . | . | . | |
Centaurea phrygia subsp. abbreviata | 2 | 4 | . | 4 | . | . | 12 | . | . | . | . | 29 | . | . | . | . | . | . | . | . | . | . | |
Nepeta racemosa | 8 | 13 | 5 | 13 | . | 7 | 29 | . | 6 | . | 7 | 57 | 10 | . | 10 | . | . | . | . | 14 | . | 18 | |
Asperula arvensis | 31 | 53 | 14 | 53 | . | 21 | 65 | 46 | 47 | . | 21 | 100 | 40 | 46 | 60 | 29 | . | . | 13 | . | 44 | 9 | |
Assoc. 1.1.1.2 | |||||||||||||||||||||||
Stipa capillata | 23 | 40 | 10 | 40 | . | 14 | 47 | 46 | 29 | . | 14 | . | 80 | 46 | 50 | . | . | . | . | . | 31 | 9 | |
Allium cardiostemon | 2 | 4 | . | 4 | . | . | 12 | . | . | . | . | . | 20 | . | . | . | . | . | . | . | . | . | |
Euphorbia condylocarpa | 7 | 9 | 6 | 9 | . | 10 | 24 | . | . | . | 10 | . | 40 | . | . | . | . | . | 13 | . | 6 | 18 | |
All. 1.1.2 | |||||||||||||||||||||||
Noaea mucronata | 10 | 23 | . | 23 | . | . | . | 77 | 6 | . | . | . | . | 77 | . | 14 | . | . | . | . | . | . | |
Cousinia brachyptera | 8 | 19 | . | 19 | . | . | 6 | 62 | . | . | . | . | 10 | 62 | . | . | . | . | . | . | . | . | |
Astragalus hyalolepis | 13 | 26 | 3 | 26 | 5 | 2 | 12 | 77 | . | 5 | 2 | . | 20 | 77 | . | . | 8 | . | . | . | 6 | . | |
Allium pseudoflavum | 16 | 36 | 2 | 36 | . | 2 | 12 | 85 | 24 | . | 2 | 29 | . | 85 | 20 | 29 | . | . | . | . | 6 | . | |
Minuartia hamata | 24 | 51 | 3 | 51 | . | 5 | 29 | 100 | 35 | . | 5 | 29 | 30 | 100 | 20 | 57 | . | . | 13 | . | 6 | . | |
B | Syntrichia caninervis | 6 | 15 | . | 15 | . | . | . | 46 | 6 | . | . | . | . | 46 | . | 14 | . | . | . | . | . | . |
Androsace albana | 4 | 9 | . | 9 | . | . | . | 31 | . | . | . | . | . | 31 | . | . | . | . | . | . | . | . | |
Polygala hohenackeriana | 4 | 9 | . | 9 | . | . | . | 31 | . | . | . | . | . | 31 | . | . | . | . | . | . | . | . | |
Meniocus linifolius | 16 | 38 | . | 38 | . | . | 24 | 77 | 24 | . | . | 14 | 30 | 77 | . | 57 | . | . | . | . | . | . | |
Alyssum turkestanicum | 9 | 21 | . | 21 | . | . | 12 | 54 | 6 | . | . | 29 | . | 54 | 10 | . | . | . | . | . | . | . | |
Arenaria serpyllifolia aggr. | 32 | 57 | 13 | 57 | 10 | 14 | 59 | 100 | 24 | 10 | 14 | 86 | 40 | 100 | 30 | 14 | 15 | . | 38 | . | 13 | 9 | |
All. 1.1.3 | |||||||||||||||||||||||
Chardinia orientalis | 11 | 26 | . | 26 | . | . | 12 | 15 | 47 | . | . | . | 20 | 15 | 40 | 57 | . | . | . | . | . | . | |
Crepis sancta | 15 | 34 | 2 | 34 | . | 2 | 12 | 31 | 59 | . | 2 | . | 20 | 31 | 70 | 43 | . | . | 13 | . | . | . | |
Stipa holosericea | 11 | 26 | . | 26 | . | . | 18 | 8 | 47 | . | . | . | 30 | 8 | 50 | 43 | . | . | . | . | . | . | |
Aegilops cylindrica | 7 | 17 | . | 17 | . | . | . | 15 | 35 | . | . | . | . | 15 | 30 | 43 | . | . | . | . | . | . | |
Roemeria hybrida | 4 | 9 | . | 9 | . | . | . | . | 24 | . | . | . | . | . | 20 | 29 | . | . | . | . | . | . | |
Noccaea perfoliata | 26 | 38 | 17 | 38 | 14 | 19 | 35 | . | 71 | 14 | 19 | 29 | 40 | . | 60 | 86 | 23 | . | . | 57 | 6 | 27 | |
Bromus japonicus | 15 | 30 | 5 | 30 | . | 7 | 24 | 15 | 47 | . | 7 | 14 | 30 | 15 | 40 | 57 | . | . | 13 | . | 6 | 9 | |
Assoc. 1.1.3.1 | |||||||||||||||||||||||
Torilis arvensis | 3 | 6 | . | 6 | . | . | . | . | 18 | . | . | . | . | . | 30 | . | . | . | . | . | . | . | |
Acantholimon caryophyllaceum | 5 | 9 | 2 | 9 | . | 2 | . | . | 24 | . | 2 | . | . | . | 40 | . | . | . | . | 14 | . | . | |
Lomelosia rotata | 7 | 15 | 2 | 15 | . | 2 | . | 23 | 24 | . | 2 | . | . | 23 | 40 | . | . | . | . | . | 6 | . | |
Carduus hamulosus | 2 | 4 | . | 4 | . | . | . | . | 12 | . | . | . | . | . | 20 | . | . | . | . | . | . | . | |
Stipa zalesskii subsp. pontica | 2 | 4 | . | 4 | . | . | . | . | 12 | . | . | . | . | . | 20 | . | . | . | . | . | . | . | |
Geranium lucidum | 2 | 4 | . | 4 | . | . | . | . | 12 | . | . | . | . | . | 20 | . | . | . | . | . | . | . | |
Aegilops triuncialis | 6 | 15 | . | 15 | . | . | 12 | . | 29 | . | . | . | 20 | . | 40 | 14 | . | . | . | . | . | . | |
Assoc. 1.1.3.2 | |||||||||||||||||||||||
Acantholimon vedicum | 5 | 11 | . | 11 | . | . | . | . | 29 | . | . | . | . | . | . | 71 | . | . | . | . | . | . | |
Stachys inflata | 11 | 21 | 3 | 21 | . | 5 | 6 | . | 53 | . | 5 | 14 | . | . | 20 | 100 | . | . | 25 | . | . | . | |
Bunium microcarpum | 8 | 13 | 5 | 13 | . | 7 | . | . | 35 | . | 7 | . | . | . | . | 86 | . | . | 25 | . | . | 9 | |
Petrorhagia cretica | 4 | 9 | . | 9 | . | . | . | . | 24 | . | . | . | . | . | . | 57 | . | . | . | . | . | . | |
Helianthemum ledifolium | 11 | 26 | . | 26 | . | . | 6 | 23 | 47 | . | . | . | 10 | 23 | 20 | 86 | . | . | . | . | . | . | |
Gaudiniopsis macra | 5 | 11 | . | 11 | . | . | . | 8 | 24 | . | . | . | . | 8 | . | 57 | . | . | . | . | . | . | |
Ephedra procera | 3 | 6 | . | 6 | . | . | . | . | 18 | . | . | . | . | . | . | 43 | . | . | . | . | . | . | |
Papaver minus | 3 | 6 | . | 6 | . | . | . | . | 18 | . | . | . | . | . | . | 43 | . | . | . | . | . | . | |
Aethionema carneum | 3 | 6 | . | 6 | . | . | . | . | 18 | . | . | . | . | . | . | 43 | . | . | . | . | . | . | |
Arabis auriculata aggr. | 9 | 19 | 2 | 19 | . | 2 | 6 | 15 | 35 | . | 2 | . | 10 | 15 | 10 | 71 | . | . | . | . | . | 9 | |
Lamium amplexicaule | 4 | 6 | 2 | 6 | 5 | . | . | . | 18 | 5 | . | . | . | . | . | 43 | 8 | . | . | . | . | . | |
Camelina laxa | 4 | 9 | . | 9 | . | . | 6 | . | 18 | . | . | . | 10 | . | . | 43 | . | . | . | . | . | . | |
Galium verticillatum | 7 | 13 | 3 | 13 | . | 5 | 6 | . | 29 | . | 5 | . | 10 | . | 10 | 57 | . | . | . | 14 | 6 | . | |
Ziziphora tenuior | 13 | 30 | . | 30 | . | . | 6 | 38 | 47 | . | . | 14 | . | 38 | 30 | 71 | . | . | . | . | . | . | |
Aegilops biuncialis | 2 | 4 | . | 4 | . | . | . | . | 12 | . | . | . | . | . | . | 29 | . | . | . | . | . | . | |
Cuscuta pedicellata | 2 | 4 | . | 4 | . | . | . | . | 12 | . | . | . | . | . | . | 29 | . | . | . | . | . | . | |
Valerianella coronata | 2 | 4 | . | 4 | . | . | . | . | 12 | . | . | . | . | . | . | 29 | . | . | . | . | . | . | |
Onobrychis atropatana | 2 | 4 | . | 4 | . | . | . | . | 12 | . | . | . | . | . | . | 29 | . | . | . | . | . | . | |
Cousinia daralaghezica | 2 | 4 | . | 4 | . | . | . | . | 12 | . | . | . | . | . | . | 29 | . | . | . | . | . | . | |
Astragalus ornithopodioides | 2 | 4 | . | 4 | . | . | . | . | 12 | . | . | . | . | . | . | 29 | . | . | . | . | . | . | |
Crucianella exasperata | 2 | 4 | . | 4 | . | . | . | . | 12 | . | . | . | . | . | . | 29 | . | . | . | . | . | . | |
Stipa arabica | 14 | 30 | 2 | 30 | . | 2 | 18 | 31 | 41 | . | 2 | 29 | 10 | 31 | 20 | 71 | . | . | . | . | 6 | . | |
Cl. 2 | |||||||||||||||||||||||
Lotus corniculatus | 39 | 2 | 67 | 2 | 90 | 55 | 6 | . | . | 90 | 55 | . | 10 | . | . | . | 92 | 88 | 88 | 57 | 31 | 64 | |
Achillea millefolium aggr. | 47 | 13 | 73 | 13 | 95 | 62 | 29 | . | 6 | 95 | 62 | 43 | 20 | . | 10 | . | 92 | 100 | 75 | 57 | 63 | 55 | |
Dactylis glomerata | 55 | 26 | 76 | 26 | 57 | 86 | 47 | . | 24 | 57 | 86 | 43 | 50 | . | 40 | . | 54 | 63 | 100 | 86 | 88 | 73 | |
Poa pratensis aggr. | 33 | 11 | 49 | 11 | 71 | 38 | 18 | 15 | . | 71 | 38 | . | 30 | 15 | . | . | 69 | 75 | 50 | 86 | 13 | 36 | |
Scabiosa bipinnata | 34 | 6 | 54 | 6 | 38 | 62 | 12 | 8 | . | 38 | 62 | 29 | . | 8 | . | . | 54 | 13 | 63 | 43 | 63 | 73 | |
Campanula glomerata aggr. | 18 | . | 32 | . | 33 | 31 | . | . | . | 33 | 31 | . | . | . | . | . | 15 | 63 | 50 | 71 | 13 | 18 | |
Potentilla recta aggr. | 51 | 28 | 68 | 28 | 71 | 67 | 41 | 15 | 24 | 71 | 67 | 29 | 50 | 15 | 40 | . | 69 | 75 | 88 | 43 | 81 | 45 | |
Galium verum | 73 | 53 | 87 | 53 | 81 | 90 | 47 | 62 | 53 | 81 | 90 | 71 | 30 | 62 | 60 | 43 | 77 | 88 | 88 | 100 | 81 | 100 | |
Phleum phleoides | 25 | 6 | 40 | 6 | 52 | 33 | 6 | . | 12 | 52 | 33 | 14 | . | . | 20 | . | 54 | 50 | 25 | 71 | 31 | 18 | |
Pimpinella saxifraga aggr. | 17 | . | 30 | . | 48 | 21 | . | . | . | 48 | 21 | . | . | . | . | . | 69 | 13 | 13 | . | 6 | 64 | |
Polygala anatolica | 18 | 2 | 30 | 2 | 29 | 31 | . | 8 | . | 29 | 31 | . | . | 8 | . | . | 46 | . | . | 57 | 31 | 36 | |
Leontodon hispidus | 19 | 4 | 30 | 4 | 48 | 21 | . | 8 | 6 | 48 | 21 | . | . | 8 | 10 | . | 62 | 25 | 13 | 57 | 13 | 18 | |
Bupleurum falcatum aggr. | 13 | . | 22 | . | 19 | 24 | . | . | . | 19 | 24 | . | . | . | . | . | 31 | . | . | 43 | 19 | 36 | |
Trifolium alpestre | 12 | . | 21 | . | 29 | 17 | . | . | . | 29 | 17 | . | . | . | . | . | 38 | 13 | 50 | . | . | 27 | |
Trisetum flavescens | 13 | 2 | 21 | 2 | 29 | 17 | . | . | 6 | 29 | 17 | . | . | . | 10 | . | 8 | 63 | 63 | 14 | . | 9 | |
Ord. 2.1 | |||||||||||||||||||||||
Trifolium ambiguum | 13 | . | 22 | . | 62 | 2 | . | . | . | 62 | 2 | . | . | . | . | . | 46 | 88 | 13 | . | . | . | |
Campanula stevenii | 22 | . | 38 | . | 81 | 17 | . | . | . | 81 | 17 | . | . | . | . | . | 77 | 88 | 50 | . | 6 | 18 | |
Plantago atrata | 27 | 2 | 46 | 2 | 86 | 26 | . | . | 6 | 86 | 26 | . | . | . | 10 | . | 85 | 88 | 13 | 29 | 25 | 36 | |
Trifolium trichocephalum | 11 | . | 19 | . | 52 | 2 | . | . | . | 52 | 2 | . | . | . | . | . | 62 | 38 | . | . | . | 9 | |
Bromopsis variegata | 23 | 11 | 32 | 11 | 76 | 10 | 29 | . | . | 76 | 10 | 57 | 10 | . | . | . | 92 | 50 | . | 14 | . | 27 | |
Myosotis alpestris | 8 | . | 14 | . | 38 | 2 | . | . | . | 38 | 2 | . | . | . | . | . | 46 | 25 | . | . | 6 | . | |
Koeleria albovii | 12 | . | 21 | . | 43 | 10 | . | . | . | 43 | 10 | . | . | . | . | . | 38 | 50 | . | . | 25 | . | |
[…] | |||||||||||||||||||||||
Veronica denudata | 21 | 9 | 30 | 9 | 57 | 17 | 12 | 8 | 6 | 57 | 17 | . | 20 | 8 | 10 | . | 54 | 63 | 25 | 14 | 13 | 18 | |
[…] | |||||||||||||||||||||||
Festuca ovina aggr. | 20 | 11 | 27 | 11 | 52 | 14 | . | 31 | 6 | 52 | 14 | . | . | 31 | 10 | . | 54 | 50 | 13 | . | 19 | 18 | |
[…] | |||||||||||||||||||||||
Taraxacum sect. Taraxacum | 23 | 6 | 35 | 6 | 62 | 21 | 12 | . | 6 | 62 | 21 | 14 | 10 | . | 10 | . | 69 | 50 | 25 | 43 | 6 | 27 | |
Assoc. 2.1.1.1 | |||||||||||||||||||||||
Ranunculus caucasicus | 6 | . | 11 | . | 33 | . | . | . | . | 33 | . | . | . | . | . | . | 54 | . | . | . | . | . | |
Huynhia pulchra | 5 | . | 10 | . | 29 | . | . | . | . | 29 | . | . | . | . | . | . | 46 | . | . | . | . | . | |
Lomelosia caucasica | 5 | . | 10 | . | 29 | . | . | . | . | 29 | . | . | . | . | . | . | 46 | . | . | . | . | . | |
Alchemilla sericea | 7 | . | 13 | . | 33 | 2 | . | . | . | 33 | 2 | . | . | . | . | . | 54 | . | . | . | . | 9 | |
Campanula collina | 5 | . | 8 | . | 24 | . | . | . | . | 24 | . | . | . | . | . | . | 38 | . | . | . | . | . | |
Polygonum cognatum | 13 | . | 22 | . | 48 | 10 | . | . | . | 48 | 10 | . | . | . | . | . | 69 | 13 | 25 | . | 6 | 9 | |
Phleum alpinum | 6 | . | 11 | . | 29 | 2 | . | . | . | 29 | 2 | . | . | . | . | . | 46 | . | 13 | . | . | . | |
Cirsium leucocephalum | 15 | . | 27 | . | 52 | 14 | . | . | . | 52 | 14 | . | . | . | . | . | 77 | 13 | 13 | 43 | . | 18 | |
Schedonorus pratensis | 7 | . | 13 | . | 29 | 5 | . | . | . | 29 | 5 | . | . | . | . | . | 46 | . | . | . | . | 18 | |
Stipa tirsa | 8 | . | 14 | . | 29 | 7 | . | . | . | 29 | 7 | . | . | . | . | . | 46 | . | . | . | 19 | . | |
B | Tortula acaulon | 4 | . | 6 | . | 19 | . | . | . | . | 19 | . | . | . | . | . | . | 31 | . | . | . | . | . |
Pedicularis condensata | 4 | . | 6 | . | 19 | . | . | . | . | 19 | . | . | . | . | . | . | 31 | . | . | . | . | . | |
Psephellus xanthocephalus | 4 | . | 6 | . | 19 | . | . | . | . | 19 | . | . | . | . | . | . | 31 | . | . | . | . | . | |
Arenaria blepharophylla aggr. | 5 | . | 10 | . | 24 | 2 | . | . | . | 24 | 2 | . | . | . | . | . | 38 | . | . | . | . | 9 | |
Rumex acetosella | 5 | . | 10 | . | 24 | 2 | . | . | . | 24 | 2 | . | . | . | . | . | 38 | . | 13 | . | . | . | |
Assoc. 2.1.1.2 | |||||||||||||||||||||||
Tragopogon reticulatus | 11 | 4 | 16 | 4 | 24 | 12 | 6 | 8 | . | 24 | 12 | . | 10 | 8 | . | . | . | 63 | 13 | . | . | 36 | |
Senecio pseudo-orientalis | 4 | . | 6 | . | 14 | 2 | . | . | . | 14 | 2 | . | . | . | . | . | . | 38 | 13 | . | . | . | |
Astracantha aurea | 6 | 2 | 10 | 2 | 19 | 5 | 6 | . | . | 19 | 5 | . | 10 | . | . | . | . | 50 | 13 | 14 | . | . | |
Lathyrus digitatus | 5 | . | 8 | . | 19 | 2 | . | . | . | 19 | 2 | . | . | . | . | . | 8 | 38 | . | . | 6 | . | |
Verbascum speciosum | 2 | . | 3 | . | 10 | . | . | . | . | 10 | . | . | . | . | . | . | . | 25 | . | . | . | . | |
Trifolium spadiceum | 2 | . | 3 | . | 10 | . | . | . | . | 10 | . | . | . | . | . | . | . | 25 | . | . | . | . | |
Gagea glacialis | 2 | . | 3 | . | 10 | . | . | . | . | 10 | . | . | . | . | . | . | . | 25 | . | . | . | . | |
Ord. 2.2 | |||||||||||||||||||||||
Stachys macrostachys | 9 | . | 16 | . | . | 24 | . | . | . | . | 24 | . | . | . | . | . | . | . | 75 | 57 | . | . | |
Stachys recta | 50 | 32 | 63 | 32 | 33 | 79 | 65 | 8 | 18 | 33 | 79 | 57 | 70 | 8 | 30 | . | 54 | . | 38 | 100 | 88 | 82 | |
Securigera varia | 35 | 15 | 49 | 15 | 38 | 55 | 24 | . | 18 | 38 | 55 | 29 | 20 | . | 30 | . | 54 | 13 | 75 | 100 | 44 | 27 | |
Cerinthe minor | 16 | 6 | 24 | 6 | . | 36 | 12 | . | 6 | . | 36 | . | 20 | . | 10 | . | . | . | . | 57 | 44 | 36 | |
Teucrium chamaedrys | 16 | 4 | 25 | 4 | 5 | 36 | 12 | . | . | 5 | 36 | 14 | 10 | . | . | . | 8 | . | 13 | 43 | 38 | 45 | |
Salvia verticillata | 27 | 15 | 37 | 15 | 10 | 50 | 24 | . | 18 | 10 | 50 | 43 | 10 | . | 30 | . | 15 | . | 38 | 43 | 50 | 64 | |
Stipa pulcherrima | 24 | 15 | 30 | 15 | . | 45 | 6 | 8 | 29 | . | 45 | . | 10 | 8 | 20 | 43 | . | . | 13 | 57 | 56 | 45 | |
Assoc. 2.2.1.1 | |||||||||||||||||||||||
Artemisia absinthium | 12 | . | 21 | . | 19 | 21 | . | . | . | 19 | 21 | . | . | . | . | . | 23 | 13 | 75 | 14 | . | 18 | |
Verbascum cheiranthifolium | 2 | . | 3 | . | . | 5 | . | . | . | . | 5 | . | . | . | . | . | . | . | 25 | . | . | . | |
Arenaria graminea | 2 | . | 3 | . | . | 5 | . | . | . | . | 5 | . | . | . | . | . | . | . | 25 | . | . | . | |
Chaerophyllum roseum | 2 | . | 3 | . | . | 5 | . | . | . | . | 5 | . | . | . | . | . | . | . | 25 | . | . | . | |
Assoc. 2.2.1.2 | |||||||||||||||||||||||
Klasea radiata | 8 | . | 14 | . | 5 | 19 | . | . | . | 5 | 19 | . | . | . | . | . | 8 | . | . | 71 | 19 | . | |
Linum nervosum | 17 | 6 | 25 | 6 | 5 | 36 | . | 15 | 6 | 5 | 36 | . | . | 15 | 10 | . | 8 | . | . | 100 | 19 | 45 | |
Origanum vulgare | 13 | . | 22 | . | 10 | 29 | . | . | . | 10 | 29 | . | . | . | . | . | 15 | . | 13 | 86 | 6 | 36 | |
Onobrychis transcaucasica | 19 | 4 | 30 | 4 | 14 | 38 | 12 | . | . | 14 | 38 | . | 20 | . | . | . | 23 | . | 13 | 100 | 31 | 27 | |
Thalictrum minus | 9 | . | 16 | . | 14 | 17 | . | . | . | 14 | 17 | . | . | . | . | . | 15 | 13 | 25 | 71 | . | . | |
B | Campyliadelphus chrysophyllus | 4 | . | 6 | . | 5 | 7 | . | . | . | 5 | 7 | . | . | . | . | . | 8 | . | . | 43 | . | . |
Helictochloa armeniaca | 5 | . | 8 | . | . | 12 | . | . | . | . | 12 | . | . | . | . | . | . | . | . | 43 | 6 | 9 | |
Vicia canescens subsp. variegata | 18 | 6 | 27 | 6 | 14 | 33 | 6 | . | 12 | 14 | 33 | 14 | . | . | 20 | . | 23 | . | 25 | 86 | 13 | 36 | |
Rhinanthus subulatus | 2 | . | 3 | . | . | 5 | . | . | . | . | 5 | . | . | . | . | . | . | . | . | 29 | . | . | |
Lathyrus latifolius | 2 | . | 3 | . | . | 5 | . | . | . | . | 5 | . | . | . | . | . | . | . | . | 29 | . | . | |
Valeriana officinalis aggr. | 2 | . | 3 | . | . | 5 | . | . | . | . | 5 | . | . | . | . | . | . | . | . | 29 | . | . | |
Campanula bononiensis | 2 | . | 3 | . | . | 5 | . | . | . | . | 5 | . | . | . | . | . | . | . | . | 29 | . | . | |
Melampyrum sp. | 2 | . | 3 | . | . | 5 | . | . | . | . | 5 | . | . | . | . | . | . | . | . | 29 | . | . | |
Seseli libanotis | 8 | . | 14 | . | 14 | 14 | . | . | . | 14 | 14 | . | . | . | . | . | 15 | 13 | 13 | 57 | 6 | . | |
Assoc. 2.2.1.4 | |||||||||||||||||||||||
Linum tenuifolium | 15 | . | 25 | . | . | 38 | . | . | . | . | 38 | . | . | . | . | . | . | . | 13 | . | 44 | 73 | |
Coronilla coronata | 4 | . | 6 | . | . | 10 | . | . | . | . | 10 | . | . | . | . | . | . | . | . | . | . | 36 | |
Asphodeline taurica | 4 | . | 6 | . | . | 10 | . | . | . | . | 10 | . | . | . | . | . | . | . | . | . | . | 36 | |
Androsace chamaejasme | 4 | . | 6 | . | . | 10 | . | . | . | . | 10 | . | . | . | . | . | . | . | . | . | . | 36 | |
B | Hypnum cupressiforme | 13 | 4 | 19 | 4 | 14 | 21 | . | 8 | 6 | 14 | 21 | . | . | 8 | . | 14 | 15 | 13 | 13 | . | . | 73 |
Campanula sibirica | 9 | 6 | 11 | 6 | . | 17 | 18 | . | . | . | 17 | 43 | . | . | . | . | . | . | . | . | . | 64 | |
Sesleria phleoides | 6 | 2 | 10 | 2 | 5 | 12 | . | 8 | . | 5 | 12 | . | . | 8 | . | . | 8 | . | . | . | . | 45 | |
Asperula affinis | 5 | . | 8 | . | . | 12 | . | . | . | . | 12 | . | . | . | . | . | . | . | . | . | 6 | 36 | |
Campanula rapunculoides | 11 | . | 19 | . | 5 | 26 | . | . | . | 5 | 26 | . | . | . | . | . | 8 | . | 13 | 43 | . | 64 | |
Pontechium maculatum | 7 | . | 13 | . | 14 | 12 | . | . | . | 14 | 12 | . | . | . | . | . | 23 | . | . | . | . | 45 | |
Fritillaria caucasica | 3 | . | 5 | . | . | 7 | . | . | . | . | 7 | . | . | . | . | . | . | . | . | . | . | 27 | |
Differential species in more than one association | |||||||||||||||||||||||
Arenaria dianthoides | 5 | . | 8 | . | 14 | 5 | . | . | . | 14 | 5 | . | . | . | . | . | . | 38 | 25 | . | . | . | |
Gagea caroli-kochii | 5 | . | 8 | . | 10 | 7 | . | . | . | 10 | 7 | . | . | . | . | . | . | 25 | 38 | . | . | . | |
Silene cephalantha | 5 | . | 8 | . | 10 | 7 | . | . | . | 10 | 7 | . | . | . | . | . | . | 25 | 38 | . | . | . | |
Companion species | |||||||||||||||||||||||
Thymus kotschyanus | 60 | 66 | 56 | 66 | 48 | 60 | 59 | 69 | 71 | 48 | 60 | 43 | 70 | 69 | 60 | 86 | 31 | 75 | 50 | 71 | 56 | 64 | |
Scutellaria orientalis aggr. | 54 | 62 | 48 | 62 | 43 | 50 | 71 | 54 | 59 | 43 | 50 | 100 | 50 | 54 | 50 | 71 | 15 | 88 | 50 | 57 | 44 | 55 | |
Koeleria macrantha | 52 | 55 | 49 | 55 | 43 | 52 | 59 | 77 | 35 | 43 | 52 | 57 | 60 | 77 | 40 | 29 | 62 | 13 | 38 | 29 | 63 | 64 | |
Elytrigia intermedia aggr. | 47 | 49 | 46 | 49 | 19 | 60 | 65 | 46 | 35 | 19 | 60 | 71 | 60 | 46 | 50 | 14 | 8 | 38 | 88 | 57 | 44 | 64 | |
Eryngium billardierei | 38 | 43 | 35 | 43 | 5 | 50 | 24 | 46 | 59 | 5 | 50 | 14 | 30 | 46 | 60 | 57 | . | 13 | 50 | 43 | 69 | 27 | |
Convolvulus lineatus | 36 | 51 | 25 | 51 | . | 38 | 71 | 62 | 24 | . | 38 | 43 | 90 | 62 | 40 | . | . | . | 25 | . | 50 | 55 | |
Medicago sativa | 35 | 34 | 37 | 34 | 14 | 48 | 29 | 46 | 29 | 14 | 48 | . | 50 | 46 | 50 | . | 15 | 13 | 38 | 71 | 44 | 45 | |
Alyssum alyssoides | 35 | 43 | 29 | 43 | 19 | 33 | 41 | 38 | 47 | 19 | 33 | 43 | 40 | 38 | 40 | 57 | 23 | 13 | 50 | . | 56 | 9 | |
Festuca valesiaca aggr. | 35 | 23 | 43 | 23 | 48 | 40 | 41 | 8 | 18 | 48 | 40 | 14 | 60 | 8 | 20 | 14 | 54 | 38 | 75 | . | 38 | 45 | |
Euphorbia seguieriana | 32 | 40 | 25 | 40 | 10 | 33 | 24 | 62 | 41 | 10 | 33 | 43 | 10 | 62 | 50 | 29 | 8 | 13 | 13 | 29 | 56 | 18 | |
Centaurea ovina aggr. | 27 | 38 | 19 | 38 | . | 29 | 47 | 46 | 24 | . | 29 | . | 80 | 46 | 30 | 14 | . | . | . | 14 | 56 | 18 | |
B | Syntrichia ruralis | 27 | 21 | 32 | 21 | 24 | 36 | 6 | 31 | 29 | 24 | 36 | . | 10 | 31 | 50 | . | 23 | 25 | 38 | 57 | 19 | 45 |
Dianthus cretaceus | 26 | 21 | 30 | 21 | 52 | 19 | 29 | 31 | 6 | 52 | 19 | 14 | 40 | 31 | . | 14 | 54 | 50 | 13 | 29 | 6 | 36 | |
Medicago x varia | 26 | 26 | 27 | 26 | 29 | 26 | 41 | 23 | 12 | 29 | 26 | 43 | 40 | 23 | 20 | . | 8 | 63 | 13 | . | 44 | 27 | |
Ziziphora clinopodioides | 25 | 19 | 30 | 19 | 14 | 38 | 24 | . | 29 | 14 | 38 | . | 40 | . | 40 | 14 | . | 38 | 13 | 43 | 56 | 27 | |
Hypericum scabrum | 25 | 32 | 19 | 32 | . | 29 | 41 | . | 47 | . | 29 | 29 | 50 | . | 60 | 29 | . | . | 13 | . | 56 | 18 | |
Leontodon asperrimus | 25 | 13 | 33 | 13 | 33 | 33 | 29 | . | 6 | 33 | 33 | 14 | 40 | . | 10 | . | 23 | 50 | 25 | 14 | 38 | 45 | |
Thesium ramosum | 22 | 36 | 11 | 36 | 5 | 14 | 53 | 23 | 29 | 5 | 14 | 43 | 60 | 23 | 30 | 29 | 8 | . | 13 | 29 | 6 | 18 | |
Plantago lanceolata | 22 | 15 | 27 | 15 | 10 | 36 | 35 | . | 6 | 10 | 36 | 14 | 50 | . | 10 | . | 8 | 13 | 63 | 43 | 25 | 27 | |
Onobrychis cornuta | 21 | 19 | 22 | 19 | . | 33 | 29 | 8 | 18 | . | 33 | . | 50 | 8 | 20 | 14 | . | . | . | 29 | 44 | 45 | |
Falcaria vulgaris | 20 | 17 | 22 | 17 | 10 | 29 | 18 | 8 | 24 | 10 | 29 | 14 | 20 | 8 | 40 | . | 8 | 13 | 13 | 57 | 38 | 9 |
Dry grassland and thorn-cushion communities of Armenia that do not belong to the Festuco-Brometea. A. Euphorbia orientalis-Melilotus officinalis scree community near Hermon (Vayots Dzor Province); B. Stachys lavandulifolia-Astracantha condensata community (alliance 1.1.1); C. Marrubio parviflorae-Stipetum capillatae (alliance 1.1.1); D. Semi-deserts of the association Noaeo mucronatae-Artemisietum fragrantis (alliance 1.1.2); E. Highland xerophytic vegetation of the Acantholimono caryophyllacei-Stipetum holosericeae (alliance 1.1.3); F. Stachys inflata-Acantholimon vedicum community (alliance 1.1.3) (Photos: A, C, E: Jürgen Dengler; B: Thomas Becker; D: Denys Vynokurov; F: Dariia Borovyk).
Dry grasslands of Armenia classified within the Festuco-Brometea. A. Mountain meadow steppes in the Lake Arpi National Park with the association Ranunculo caucasici-Bromopsietum variegatae (alliance 2.1.1); B. Tragopogon reticulatus-Astracantha aurea community (alliance 2.1.1); C. Trisetum flavescens-Stachys macrostachys community (alliance 2.2.1); D. Onobrychis transcaucasica-Vicia canescens subsp. variegata community (alliance 2.2.1); E. Globulario trichosanthae-Stipetum pulcherrimae (alliance 2.2.1); F. mountain steppes near the Sevan Lake with the association Seslerio phleoidis-Onobrychidetum cornutae (alliance 2.2.1). (Photos: A: Philipp Kirschner; B, C: Dariia Borovyk; D, F: Jürgen Dengler; E: Denys Vynokurov).
Communities within the Festuco-Brometea predominantly thrived at higher elevations, especially those of Plantagini atratae-Bromopsietalia variegatae, reaching up to 2,400 m a.s.l. (Figure
In contrast, communities of the Ziziphora tenuior-Stipa arabica grasslands with its order Cousinio brachypterae-Stipetalia arabicae occurred at lower elevations, primarily below 2,000 m a.s.l. Among its three alliances, Artemision fragrantis, which comprises wormwood semi-deserts, thrived at the lowest altitudes, ranging from 1,300 to 1,600 m a.s.l. Additionally, the Stachys inflata-Acantholimon vedicum community, which belongs to the alliance Acantholimono caryophyllacei–Stipion holosericeae, also occupied comparably low elevations, at about 1,600 m a.s.l. Other units of this order generally occupy elevations not exceeding 2,000 m a.s.l.
The altitudinal zonation reflected the climatic preferences of the communities. The Cousinio brachypterae-Stipetalia arabicae communities tended to prefer warmer and drier conditions, while the class Festuco-Brometea thrived in more mesic and cooler environments (Figure
Regarding soil characteristics, most communities preferred neutral or slightly alkaline soils, except for the order Plantagini atratae-Bromopsietalia variegatae, which occurred in slightly acidic soil conditions with a pH around 6.5 (Figure
In terms of skeleton content, communities of the Ziziphora tenuior-Stipa arabica grasslands generally tended to occur on sites with shallower soils and higher skeleton content (Figure
All communities of the class Festuco-Brometea tended to occur on sites with high humus content, with mean values within a narrow range of 7.5–8.1%. The tentative Ziziphora tenuior-Stipa arabica grasslands occurred on sites with significantly lower humus content, with mean values of 3.9–4.5% (Figure
Site characteristics of the grassland types at different syntaxonomic levels. Box plots (median, interquartile range, range and outliers) as well as arithmetic means (black points) are shown. For the codes of syntaxa, see Table
Soil characteristics of the grassland types at different syntaxonomic levels. Box plots (median, interquartile range, range and outliers) as well as arithmetic means (black points) are shown. For the codes of syntaxa, see Table
Herb layer cover and litter cover, proxies for ecosystem productivity, were notably higher in the communities belonging to the class Festuco-Brometea (Figure
Regarding the dominant life forms, we observed a strong differentiation between the two classes. Communities belonging to the class Festuco-Brometea had a significantly higher proportion of hemicryptophytes, which was highest in the case of the Ranunculo caucasici-Bromopsietum variegatae association (Figure
The proportion of species’ range types also showed a strong differentiation between the two classes (Figure
The presence of endemic species in the studied communities was also remarkable. While there were no significant differences detected between the two classes concerning narrow Transcaucasian endemics (Figure
Structure and dominant life forms of the grassland types at different syntaxonomic levels. Box plots (median, interquartile range, range and outliers) as well as arithmetic means (black points) are shown. For the codes of syntaxa, see Table
Proportions of range types of the grassland types at different syntaxonomic levels. Box plots (median, interquartile range, range and outliers) as well as arithmetic means (black points) are shown. For the codes of syntaxa, see Table
Total species richness in 10 m2 did not differ significantly among the higher syntaxa (Figure
Species richness in 10 m2-plots for different taxonomic groups compared at different syntaxonomic levels. Box plots (median, interquartile range, range and outliers) as well as arithmetic means (black points) are shown. For the codes of syntaxa, see Table
The type of the class Astragalo-Brometea, namely the order Astragalo-Brometalia, along with other orders traditionally associated with it (Drabo-Androsacetalia, Hyperico linarioidis-Thymetalia scorpilii, Onobrychido armenae-Thymetalia leucostomi), were grouped together as “group A” in the Twinspan analysis (Figure
Groups B and E comprised plots from meso-xeric, xeric, and rocky grasslands that can be categorised as belonging to the class Festuco-Brometea. Whereas plots within group E were previously assigned to order-level units (Asphodelino tauricae-Euphorbietalia petrophilae, Festucetalia valesiacae, Brachypodietalia pinnati), group B did not have any assignments to any syntaxonomic order. Considering the clear separation between group B and group E at the very basis of the dendrogram, and the fact that group B is, in contrast to group E, positioned above the Y-axis in the DCA ordination (Figure
Group D encompasses the driest communities sampled in Armenia, particularly cluster 8. The species present in this group are predominantly distributed in the Irano-Turanian region, such as Artemisia fragrans, Eryngium billardierei, Noaea mucronata, Stipa arabica, S. holosericea, and others. This species composition suggests that similar vegetation types may also exist in other regions of Western Asia. Since there is no suitable class-level unit available, we propose that in the future, a new class should be established. To do so a comprehensive comparison involving more data from the surrounding regions would be needed. For now, in this paper, we refer to this unit as “Ziziphora tenuior-Stipa arabica grasslands”, combining the dry grassland, semi-desert and xeric thorn-cushion vegetation of Western Asia.
To summarise, we can classify all the vegetation plots of the bigger dataset into three classes: Astragalo-Brometea (groups A and C), Festuco-Brometea (groups B and E), and a tentative new class, “Ziziphora tenuior-Stipa arabica grasslands”. This was well supported by the DCA ordination (Figure
We can identify two distinct vegetation classes in Armenia: Festuco-Brometea and a novel class meant to encompass drier grasslands and thorn-cushion communities found at lower elevations. This finding aligns well with the outcomes of the TWINSPAN analysis of the Armenian plots (Figure
Cluster X in the TWINSPAN dendrogram corresponds to scree vegetation that currently cannot be assigned to any existing vegetation class. It appears to be similar to the Thlaspietea rotundifolii Br.-Bl. 1948 from temperate Europe or Drypidetea spinosae Quézel 1964 from the Mediterranean. In the North Caucasus, a class of high-altitude scree vegetation on siliceous outcrops, Lamio tomentosi-Chaerophylletea humilis Belonovskaya et al. 2014, exists. However, the latter mainly consists of subnival belt vegetation with a completely different floristic composition. Therefore, we cannot currently assign the aforementioned Armenian scree community to any existing class and leave it unassigned.
The remaining clusters in the left part of the dendrogram (clusters 1.1.1.1–1.1.3.2 in Figure
Further examination of the drier part of Armenian plots (clusters 1.1.1.1–1.1.3.2 on the dendrogram, Figure
Relevés from clusters 2.1.1.1–2.1.1.2 were previously categorized under group B in the earlier section (broad-scale comparison), together with plots from the North Caucasus belonging to the alliance Artemisio chamaemelifoliae-Bromopsion variegatae. Given their separation from the other clusters at a high level (Figure
Clusters 2.2.1.1–2.2.1.4 corresponded to the so-called mountain steppes, following the classification of
With an average of 46.8 vascular plants in 10 m2, the dry grasslands of Armenia were significantly richer than the Palaearctic average of the three relevant ecological-physiognomic vegetation types (A.3 - Xeric grasslands and steppes; B.2 - Meso-xeric grasslands; D.3 - Garrigues and thorn-cushion communities) in the high-quality database GrassPlot (v.2.10; https://edgg.org/databases/GrasslandDiversityExplorer; see
Despite having compiled the available vegetation plot data, particularly the type relevés, of the relevant syntaxa described in the other countries of the Caucasus as well as Anatolia and Northern Iran, we found low correspondence of the Armenian dry grassland communities with these. It appears that only one of our five alliances had been described before, the Artemisio chamaemelifoliae-Brompsion variegatae from the Northern Caucasus, Russia (
Even at the class level we found that the more xeric dry grassland of the lower elevations in Armenia are floristically so profoundly different from either the Euro-Siberian Festuco-Brometea or the Anatolian-Iranian Astragalo-Brometea that they might be a class of their own. However, a formal description should wait for a plot-based broad-scale classification of all the dry grasslands in the Caucasus, Anatolia and Northern Iran, similar to the studies of Eastern and Central Europe by
Within Armenia, the next logical step would be to compile more plot data of dry grasslands with the same methodology to ensure that the system is complete and all the determined diagnostic species can be confirmed. Then the system could be translated into an electronic expert system that enables the automatic and unequivocal classification of new dry grassland plots (see the example by
Finally, our collected data of biodiversity, species composition and in situ environmental variables are also valuable for broad-scale analyses on biodiversity patterns and their drivers, global change projections and biogeographic analyses. For this purpose, we have already contributed them to the relevant international plot databases, namely EVA (
All original data from Armenia (species composition and header data as well as derived metrics of the plots) are provided in the Supplementary materials of this article.
A.A. and G.F. organised the 13th EDGG Field Workshop in Armenia, and together with A.B., A.H., D.F., D.B, D.V., I.B., I.G.-M., I.V., J.D., M.M., M.O., P.K., S.P., T.B. and U.B. collected the field data. A.A. and G.F. determined vascular plant specimens collected during the Field Workshop, I.D. critical Festuca species, D.B and D.V Stipa species, B.C.-M. bryophytes and H.M. lichens. A.B., D.B., D.V., I.B., I.V. and S.P. digitised the field forms and harmonised the data. D.V. performed the literature search and digitised vegetation plots from neighbouring countries. D.V. classified the relevés of the national and the supranational dataset and together with J.D. and developed the classification system for Armenia. J.D. determined the diagnostic species in the Armenian dataset and prepared the synoptic tables. T.B. assigned the life forms; D.V., T.B., and G.F. classified species distribution ranges. D.B. prepared the maps; T.B. performed the ANOVAs and prepared the boxplots and M.M. performed the ordinations. The manuscript was drafted by D.V. and J.D. with significant inputs by A.A., D.B., I.B., I.G.-M. and T.B. All authors checked, improved and approved the manuscript.
We are grateful to the Eurasian Dry Grassland Group (EDGG; https://edgg.org/) and the International Association for Vegetation Science (IAVS; https://www.iavs.org/) for providing travel grants that enabled the participation of some of the researchers. Further, we thank Aslan Ünal and Elena Belonovskaya for their participation in the field work as well as Jiří Danihelka and Marcin Nobis for their help in determining Stipa specimens. Finally, we thank Victor Chepinoga for the careful and fast editorial handling of the manuscript, Wolfgang Willner for advice on phytosociological nomenclature and Michael Glase for linguistic editing.
Formal descriptions of the new syntaxa according to the ICPN
For the diagnostic species we refer to the main text as well as Table
1.1 Cousinio brachypterae-Stipetalia arabicae ord. nov. hoc loco
Holotypus hoc loco: Artemision fragrantis Vynokurov et al. 2024 (this paper)
2.1 Plantagini atratae-Bromopsietalia variegatae ord nov. hoc loco
Holotypus hoc loco: Artemisio chamaemelifoliae-Bromopsion variegatae Vynokurov in
2.2 Onobrychido transcaucasicae-Stipetalia pulcherrimae ord. nov. hoc loco
Holotypus hoc loco: Onobrychido transcaucasicae-Stipion pulcherrimae Vynokurov et al. 2024 (this paper)
1.1.1 Onobrychido michauxii-Stipion capillatae all. nov. hoc loco
Holotypus hoc loco: Marrubio parviflorae-Stipetum capillatae Vynokurov et al. 2024 (this paper)
1.1.2 Artemision fragrantis all. nov. hoc loco
Holotypus hoc loco: Noaeo mucronatae-Artemisietum fragrantis Vynokurov et al. 2024 (this paper)
1.1.3 Acantholimono caryophyllacei-Stipion holosericeae all. nov. hoc loco
Holotypus hoc loco: Acantholimono caryophyllacei-Stipetum holosericeae Vynokurov et al. 2024 (this paper)
2.2.1 Onobrychido transcaucasicae-Stipion pulcherrimae all. nov. hoc loco
Holotypus hoc loco: Seslerio phleoidis-Onobrychidetum cornutae Vynokurov et al. 2024 (this paper)
1.1.1.2 Marrubio parviflorae-Stipetum capillatae ass. nov. hoc loco
Holotypus hoc loco: plot ID 81 in Suppl. material
1.1.2.1 Noaeo mucronatae-Artemisietum fragrantis ass. nov. hoc loco
Holotypus hoc loco: plot ID 65 in Suppl. material
1.1.3.1 Acantholimono caryophyllacei-Stipetum holosericeae ass. nov. hoc loco
Holotypus hoc loco: plot ID 71 in Suppl. material
2.1.1.1 Ranunculo caucasici-Bromopsietum variegatae ass. nov. hoc loco
Holotypus hoc loco: plot ID 18 in Suppl. material
2.2.1.3 Globulario trichosanthae-Stipetum pulcherrimae ass. nov. hoc loco
Holotypus hoc loco: plot ID 93 in Suppl. material
2.2.1.4 Seslerio phleoidis-Onobrychidetum cornutae ass. nov. hoc loco
Holotypus hoc loco: plot ID 13 in Suppl. material
Complete header data of the Armenian plots (*.xlsx)
Complete synoptic table of 110 classified Armenian plots with percent constancies and phi values for the syntaxa of all levels as well as the individual plots (*.xlsx)
Definition of additional species aggregates (*.pdf)
Value distribution of all recorded and analysed numerical environmental, structural and biodiversity variables (*.pdf)
Data sources of the West Asian and Caucasian dataset (*.pdf)
List of the vascular plants from the Armenian dataset with assignment of life forms and distribution ranges (*.pdf)
Synoptic table of the West Asian and Caucasian dataset with the results of the broad-scale comparison with five distinguished groups of clusters (A–E) (*.pdf)
Planned applications to the EuroVegChecklist Committee (EVCC) (*.pdf)