Lazu Ș. et al

PHYTOCOENOTIC DIVERSITY OF GRASSLANDS WITH Chrysopogon gryllus (L.) Trin. IN EURASIA

Ștefan LAZU*, Ghenadie Petru TITICĂ*, Ludmila TALMACI*,**, Ana GUȚU*

*”Alexandru Ciubotaru” National Botanical Garden (Institute), MD 2002, Republic of Moldova, Chisinau, 18 Pădurii str. **Corresponding author e-mail: [email protected]

Abstract Chrysopogon gryllus (L.) Trin. is a species of savannoid grasses. In the Republic of Moldova, it is dominant in the xero-mesophytic grasslands of the southern steppe area (Bugeac), occurs on slopes with steppe vegetation, glades and forest edges and in the central area – on the side of forest roads, glades, as well as in xerophilous grasslands. The position of this species in the phytocoenotic classification becomes clearer if we examine its presence depending on the biogeographic regions. The analysis of the synthetic lists drawn up by different authors from the Republic of Moldova, Romania and Ukraine, countries whose territories with savannoid grasslands are part of the continental and steppe biogeographic regions, has helped us identify the species that is faithful to these communities - Thymus pannonicus All., which occurs in stable phytocoenoses of the ass. Thymo (pannonici) - Chrysopogonetum grylli Doniţă et. all. 1992. In the Mediterranean, Pannonian and alpine biogeographic regions, we have shown the dependence of these grasslands on the altitude of the site. At high altitudes (the alpine zone), in areas with slightly acid soils, there are communities of the ass. Agrostio (capilaris) - Chrysopogonetum gryllis Kojić 1959 and ass. Bromo (erectus) - Chrysopogonetum gryllis Horvat 1934. The greatest phytocoenotic diversity of grasslands with Chrysopogon gryllus (L.) Trin. is in the Mediterranean biogeographic region. Here, at the lowest altitudes, Chrysopogon gryllus (L.) Trin. occurs in discrete communities with Carex humilis, Schoenus nigricans, Teucrium capitatum, Bromus fibrosus, B. erectus, Globularia elongata, Nerium oleander, and in areas with a little higher altitudes - Asphodelus ramosus. In the biogeographic regions of the Black Sea and the Anatolian plateau, it does not grow in grasslands as dominant species, but persists as a component of the grassy vegetation in sparse forests of Quercus coccifera, Arbutus andrachne, Fantanezis phillirea, Olea sylvestris, Arbutis unedonis, Pinus brutia, Juniperus excelsa etc. In the steppes of Anatolia (550-650 m altitude), Chrysopogon gryllus is a component of the grasslands with Stipa arabica. The syntaxonomy of the grasslands with Chrysopogon gryllus (L.) Trin. requires in- depth analysis based on the ecology and the habitat (heliophilous and thermophilous) of these phytocenoses, as well as the environment with Bothriochloa ischaemum (L.) Keng., both of which are part of the savannoid Andropogoneta. The necessity arises to open phytocenotic classes of the dry grasslands Chrysopogono-Bothriochloetea n. n. with the meridional-latitudinal distribution to the west ord. Chrysopogonetalia n. n., and to the east ord. Bothriochloetalia n. n..

Keywords: syntaxonomy, dry grassland, biogeographic zone.

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INTRODUCTION

Chrysopogon gryllus (L.) (Sburlino et al., 2013) and Trin. (common name - Goldbeard encompasses various biogeographic Grass) is a species of savannoid regions - continental, steppe, grasses, which occurs in dry (xeric) Pannonian, Black Sea, Anatolian, grasslands and has a quite large alpine, it is mostly found in the range (from the south of France to Mediterranean region at various the west of the Himalayas (India), altitudes. Thailand and Australia). It is Chrysopogon gryllus is a remarkable that in the western areas paleobotanical species with of its range, the grasslands are Mediterranean-Southwest Asian represented by a single species of range, which, on the territory of the the genus Chrysopogon - Ch. Republic of Moldova, appeared gryllus, and in Asia (western after the withdrawal of Sarmatian Himalayas) apart from waters in the Middle Miocene sub- Chrysopogon gryllus, Chrysopogon epoch (20-5 million years ago), montanus, Ch. aciculatus, Ch. growing in forest steppes with sub- fulvus, Ch. serrulatus also occur. In Mediterranean floristic elements Australia, the genus Chrysopogon is (Lavrenko, 1940; Gheideman, represented by 10 species, 1989). including: Chrysopogon gryllus (L.) In the Republic of Moldova, Trin. (Blake, 1944; Ganatsas et al., the savannoid grasslands with 2004; Joshi, et al., 1964; Chrysopogon gryllus occur in the Neamsuvan et al., 2009; Qureshi et Bugeac steppe, as well as in the al., 1964; Sundriyal, 1995; Singh glades of subarid forests of Quercus Vikaspal et al., 2015), in Northwest pubescens with Cotinus coggygria Africa, Dobignard et al., 2010, in the steppe biogeographic region mentioned the presence of C. as well as in glades of the gryllus, C. aucheri (Boies.) Stapf., mesophytic forests with Quercus C. plumulosus Hochst. robur, Q. petraea and Fagus Chrysopogon gryllus is a silvatica (nature reserves ”Codrii” geophyte, grows about 50-180 cm and ”Plaiul fagului”) - the tall, is dominant mainly in sunny, continental biogeographic region. In xerophilous, xero-mesophilous, the Republic of Moldova, subthermal grasslands, on Chrysopogon gryllus is a plant chernozem soil and ”brown earth” species considered rare, because on various substrate (sand, loess, these grasslands are intensely limestone, river gravel, marl, scree grazed, but the current climate etc.), it forms stable, discrete conditions, as well as the impact of communities with a specific habitat. the global warming favour an It is mostly dependent on expansion within the areas with temperature, which ranges in the favourable soil and climate limit of the isotherm 7.5-10.5 °C conditions, in protected areas.

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occurring in productive grasslands, used as pastures, often degraded, but with different grass species, which does not clarify the situation, neither from the point of view of savannoid grasslands nor in terms of evolution, including the impact of global warming (figure 1, 2 and 3).

Figure 1. Chrysopogon gryllus (L.) Trin. on a slope with steppe vegetation (Iordanovca v., Basarabeasca d.).

Having a large range and an obvious tendency toward expansion, we decided to analyse the floristic and phytocoenotic composition of the grasslands with C. gryllus on its large natural range. From west to east, we notice that the altitude, at which these grasslands occur, ranges from 80-100 m in the west to 2000-3000 m in the east (Himalaya Figure 2. Chrysopogon gryllus (L.) Mountains in western India). In the Trin. in a grassland with steppe Mediterranean Basin (North Africa, vegetation (Moscovei v., Cahul d.). Anatolian plateau, Balkan Mountains, Eastern Alps), it is a species with compact natural range,

MATERIAL AND METHOD

In the study on the Southern Moldavian Plain, grasslands with Chrysopogon Nikolaeva, 1950-1955 - the glades gryllus in the Republic of Moldova, with steppe vegetation in the forests we used the results of with downy oak, Vitko, 1980-1981 - phytosociological studies carried the reserve with medicinal plants out in different periods of the 20th from the villages Minjir, Poruceni, century - Săvulescu, 1927; Selişte, Leu (Nisporeni), the edge Zahariadi, 1932-1934 - the hills and the glades of the mesophytic with steppe vegetation from the forests Kononov, 1956, as well as

Romanian Journal of Grassland and Forage Crops (2019) 20 57 Lazu Ș. et al herbarium specimens collected in 2015; Titica, 2017, 2018; the 21st century - from forest edges Lebedenco village, Cahul district, and grasslands in the forests with Titica, 2017; Iordanovca v., downy oak (Hânceşti) - Vorniceni, Basarabeasca dist. - Titica, 2018, as Bucovăţ, Codreanca (Străşeni), well as phytocenotic studies on Sadova (Călăraşi) (Şabanova, 2012, them, analysed by Şabanova, 2012. 2014); the Nature Reserve “Ciumai” It is obvious that these grasslands Postolache, 2017; slopes with with Chrysopogon gryllus, in the steppe vegetation (Moscovei v., Republic of Moldova, tend to Cahul distr.) Miron et al. 2017; in expand. the Bugeac Plain - Ghendov et al.

Figure 3. Phytocoenosis with Chrysopogon gryllus (L.) Trin. (Lebedenco v., Cahul d.)

The phytocenotic assessment Chrysopogon gryllus was carried of grasslands with Chrysopogon out according to the floristic and gryllus in Eurasia is not ecological principles, delimiting the homogeneous. In the European area, group of associations. Currently, in the phytocenotic classification is the communities with Chrysopogon presented according to the principle gryllus as a dominant species, of the phytosociological school of according to the phytocoenological Braun-Blanquet, 1933, 1964, that is classification (Braun-Blanquet, to say, the constancy or frequency 1964) accepted by most experts in of the species is used for definition phytocoenology, there are various in the syntaxonomic nomenclature frequent species of grasses that of the association. In the Republic reflect the local ecological of Moldova, the phytocenotic conditions, specific to the evaluated classification of dry grasslands with region. The dominant species

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Chrysopogon gryllus is not both biogeographic regions distributed evenly everywhere, but (continental and steppe). To sometimes its range intersects with elucidate the specific traits of the range of Bothriochloa locations and of the faithful species ischaemum, which has expanded in to Chrysopogon gryllus, which the northern hemisphere due to occurs in various biogeographic global warming and aridization regions, we analysed the floristic (Lazu, 2014; Lazu et al., 2016; composition and the abundance in 2017; 2018). the phytocenoses of the grasslands In the Republic of Moldova, presented by authors in synthetic the grasslands with Chrysopogon lists: Ukraine - Dubyna D.V. gryllus were studied from the point (1996); Romania - Puşcaru, of view of phytocoenology by Soroceanu et al., 1963; Borza, 1963; Şabanova, 2012; 2014, who Csuros and Nidermaier, 1966; identified the group of associations - Dihoru, 1970; Ivan et al., 1993; Chrysopogon gryllus, with Festuca Doniţă et al., 2005; Oprea and valesiaca, on the basis of 17 relevés Sârbu, 2005; Drăgulescu and from the glades of forests with Schumacher, 2006; Ștefan et al., downy oak. 2007; Chifu et al., 2014; Italy - The author distinguished the Meyer, 1975; Nimis and Fonda, second group of associations of 1997; Sburlino et al., 2013; Serbia - Chrysopogon gryllus with Jovanovici et al., 2017; Djodjovici Bothriochloa ischaemum, on the et al., 2012, Djodjovici et al., 2013; basis of the analysis of 19 relevés, Aćić et al., 2015; Croatia - on slopes with steppe vegetation, in Bernhardt et al, 2006; Ilijanici et al., areas with continental climate. 1972; Jasprica et al., 2007; Albania These groups of associations with - Fanelli et al., 2015; Greece - Pirini Chrysopogon gryllus cannot be et al., 2014; Bulgaria - Tachev et analysed in more detail because the al., 2010; Pedaşenco et al., 2013; synthetic tables have not been Macedonia - Matevski et al., 2015 presented. By analysing the papers (table 1 and figure 4). on phytocoenology, published in the The phytocoenotic studies Republic of Moldova (Şabanova, on the grasslands with Chrysopogon 2012, 2014), concerning the gryllus, carried out until now, in the communities with Chrysopogon Republic of Moldova, are gryllus, as well as the papers written insufficient. We determined the by authors of the above-mentioned position of this species in the phytosociological studies, carried phytocoenotic classification by out in the evaluated locations, we analysing synthetic tables based on have found that the faithful species the wide natural range presented by of such grasslands is Thymus numerous authors, who made marschalianus Wild. (T. pannonicus phytosociological assessments in All. According to Ciocârlan 1990, various biogeographic regions. 1992), and this species occurs in Analysing, in general, the synthetic

Romanian Journal of Grassland and Forage Crops (2019) 20 59 Lazu Ș. et al lists with Chrysopogon gryllus to representative – the characteristic which we had access, we saw the species to the habitat or the faithful need to learn more about the plant species, which are constant chorology and the capacity of according to the concept Braun- coexistence and socialization of the Blanquet, 1964. The results dominant species (Chrysopogon obtained can be seen in the table 1 gryllus (L.) Trin.) with the local and figure 4.

Figure 4. Distribution of phytocoenoses with dry grasslands with Chrysopogon gryllus (L.) Trin., in the biogeographic regions of Europe.

RESULTS AND DISCUSSION

Chrysopogon gryllus (L.) vegetation. This relict grass species Trin., syn. Andropogon gryllus L., has been mentioned (Nykolaeva, Pollinia gryllus Spr., in the 1963; Şabanova, 2012) in Republic of Moldova, is a species of monographs and scientific articles tussock grass, which occurs in as an herbaceous species that grows grasslands in the southern areas of up to 1.5 m tall, xeromesophyte and the country, particularly, in the with high fidelity to the habitat of glades of the forests of Quercus savannoid grasslands that are pubescens with Cotinus coggygria located on the northern border of the and in grasslands with steppe natural range and which passes

Romanian Journal of Grassland and Forage Crops (2019) 20 60 Lazu Ș. et al through the central area of the Brachypodium pinnatum, Republic of Moldova, bordering Calamagrostis epigejos, Dactylis with the Central “Codri” area. glomerata, Elytrigia repens, Şabanova, 2012, 2014, Elytrigia intermedia, Melica ciliata, described the associative character Poa angustifolia, Bromus inermis. of this species, in the forests of A lower abundance is characteristic downy oak, with the group of ass. of the species of Poaceae with dense Chrysopogon gryllus - Festuca caespitose roots: Koeleria cristata, valesiaca from the steppe Stipa capillata, Stipa pinnata, Stipa biogeographic region and the group tirsa, as well as those with spongy of the ass. Chrysopogon gryllus - caespitose roots, such as: Bothriochloa ischaemum (the Bothriochloa ischaemum, Phleum continental biogeographic region). phleoides, steppe shrubs: The author, on the basis of 17 Chamaecitisos austriacus, Thymus relevés (in the table 1 and figure 4 N marschallianus, Caragana mollis, 1), has detected the presence of the semi-ephemeroids - Adonis vernalis, group of associations Chrysopogon Doronicum hungaricum, Lathyrus gryllus - Festuca valesiaca with 123 pannonicus, Pulsatilla grandis, species, 103 of which are diverse. Pulsatilla nigricans, Viola ambigua, The following species are the most ephemeroids - Crocus reticulatus, characteristic for these associations Gagea minima, Muscari neglectum, and are present in all the ephemerals - Myosotis micrantha, descriptions: Thymus marchalianus, Veronica verna. Constant species - Galium campanulatum, Festuca Thymus marschalianus, Festuca valesiaca, Koeleria cristata, Phleum valesiaca, Koeleria cristata, phleoides, Elytrigia repens, Asperula cynanhica, Galium Elytrigia intermedia, Achillea campanulatum, Galium verum, pannonica, Securigera varia, Eryngium campestre, Hypericum Dianthus membranaceus, Medicago perforatum, Inula hirta, Medicago romanica, Teucrium chamaedrys, romanica, Potentilla impolita, Adonis vernalis, Crocus reticulatus, Crocus reticulatus. Arabidopsis thaliana, Myosotis In these phytocoenoses with micrantha, Veronica verna. The Chrysopogon gryllus as dominant following species are more species, the species with ordinary: Bothriochloa ischaemum, characteristics of steppe plants Melica ciliata, Euphorbia stepposa, constitute 69.9 % and the Galium verum, Helichrysum mesophytes - 23.6 %. According to arenarium, Hypericum perforatum, ecological criteria, xeromesophytes Inula hirta, Linum austriacum, make up 43.95 % and Jurnea mollisima, Otites moldavica, mesoxerophytes - 22.8 %. Stachys officinalis, Lathyrus The author considers this pannonicus, Lathyrus pallescens. association secondary, evolving Among Poaceae, the species with from those with Bothriochloa rhizomes are more abundant: ischaemum.

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Postolache Gh. (2017) Muntenia, at an altitude between mentioned the presence of 150 and 450 m. The average annual Chrysopogon gryllus in the temperature, in this area, is 10.0- grasslands of the “Ciumai” Nature 10.5 °C. Precipitation - 500- Reserve, as a plant that had been 650 mm. Relief - plateaus and included in the Red Book of the slightly inclined slopes. Soil - dark Republic of Moldova (table 1, brown chernozem on loess or figure 4, N 2). limestone. The sources mentioned Miron, 2017, mentioned the presence of 22 associations with Chrysopogon gryllus as vulnerable Chrysopogon gryllus as a taxon plant, occurring in the steppe with high degree of fidelity to the grassland, (283 ha) Moscovei habitat, including species with a village, Cahul district, where restricted range around these Thymus marschalianus was also biogeographic zones (steppe and present (table 1, figure 4, N 3). continental), but the species faithful Ghendov et al., 2015, Titica, to all the sites where Chrysopogon 2017, 2018, mentioned the presence gryllus occurs is Thymus of Chrysopogon gryllus in the pannonicus (Doniţă and Popescu, Bugeac Plain, accompanied 2005). everywhere by Thymus Dihoru, 1970, mentioned marschalianus, with varying the ass. Chrysopogonetum grylli degrees of abundance, but faithful to dobrogicum, which occurred on the these communities (table 1, figure 4, Babadag Plateau, whose plant N 4, 5, 6). communities were found on sunny, Dubyna et al., 1996, on the southwest-facing slopes and in marine soils in Jebriansk (Odessa glades of forests with Quercus district), detected the presence of pubescens. Dauco (gutati) - Chrysopogonetum The average coverage was grylli Popescu and Doltu, 1980 with 90 % (50-100 %) and the 28 species, including the following stratification consisted of 4 levels. diagnostic species: Chrysopogon The first layer, of 20 cm in height, gryllus (V), Plantago lanceolata included Teucrium polium, (V), Cynanchum acutum (III), Inula Linosyris villosa, Alyssum hirsutum salicina (V), Dianthus bessarabicus etc., the second layer, of 30 cm - (III), Apera maritima (III), Linum Festuca valesiaca; the third layer, austriacum (III). Thymus of 60 cm - Stipa joannis, Stachys marschalianus (I) also occurred in transsilvanica, Centaurea this area (table 1, figure 4, N 7). jurneifolia, Orlaya grandiflora, In steppe and continental Allium rotundum etc. and the fourth grasslands of Romania, - 100 cm - Chrysopogon gryllus, Chrysopogon gryllus occurs in Agropyron intermedium, Verbascum phytocoenoses found in the south of lanaticum, Salvia nutans. Festuca Transylvania, Banat, Dobrogea, valesiaca, Bothriochloa ischaemum, Central Moldavian Plateau and Orlaya grandiflora and Elytrigia

Romanian Journal of Grassland and Forage Crops (2019) 20 62 Lazu Ș. et al intermedia were species with grylli Doniţa et al., 1992 is particularly high abundance- mentioned in the table 1 and figure dominance. Diagnostic species of 4, N 9. the association of xerophilous Doniţă N., Popescu et al., plants, occurring in Dobrogea, 2005, identified the habitat of Daco- according to the author’s opinion, Balkan grasslands of Chrysopogon were the following: Trigonella gryllus and Festuca rupicola, which gladiata, Conringia austriaca, are among the Mediterranean Lactuca viminea, Onobrychis grasses of the plant association gracilis. From the synthetic list Thymo pannonici – made up on the basis of 21 relevés Chrysopogonetum grylli Doniţă et from Dobrogea, Dihoru, 1970, al., 1992 and occur on the Central distinguished phytocenoses with Moldavian Plateau (Muntenia, Chrysopogon gryllus in the ass. Moldova), Babadag Plateau Chryso pogonetum grylli (Dobrogea) and Transylvanian dobrogicum a. n., with a range of Plateau. The altitude, in this area is local species, but the species about 150-450 m, the average Thymus marschalianus (Th. temperature is 8-10.5 °C and the pannonicus) (II) also occurred here amount of precipitation - 550- and was close to the ecology of the 650 mm. The relief consists of dominant species Chrysopogon plateaus and slopes with slight gryllus, and this association inclination. Soils - dark brown included the basic components of chernozem on loess or limestone. the regional association Thymo Edifying and characteristic species - (pannonici)-Chrysopogonetum Chrysopogon gryllus, Thymus grylli Doniţa et al., 1992 (table 1, pannonicus. Other important figure 4, N 8). species: Thesium linophyllon, Ivan et al., 1993, described the Senecio jacovea, Asperula Chrysopogoneta that occured in cynanchica, Stachys recta, Romania, which included the Artemisia campestris, Scabiosa following companion specific ochroleuca, Adonis vernalis, floristic elements: Festuca rupicola, Trifolim montanum, Hypericum Potentilla recta, Convolvulus elegans, H. perforatum, Trifolium cantabrica, Falcaria vulgaris, campestre, Coronilla varia. These Cleistogenes serotina, Dorycnium Daco-Balkan grasslands differ from herbaceum, Vinca herbacea, those found in Banat and western Astraga- lus dasyanthus, as well as Oltenia in the presence of numerous the characteristic species: Pontic elements and the absence of Chrysopogon gryllus and Thymus Illyrian species (table 1, figure 4, N pannonicus. The authors mentioned 10). the lack of Illyrian species and the Șabanova, 2012, mentioned presence of numerous Pontic that Chrysopogon gryllus, in the elements. The association Thymo Republic of Moldova occurred at (pannonici) - Chrysopogonetum the north-eastern border of its range,

Romanian Journal of Grassland and Forage Crops (2019) 20 63 Lazu Ș. et al in Bucovăţ, Grebleşti, Codreanca of grass communities with and Negrești, Străşeni district, and Chrysopogon gryllus as dominant Sadova, Călăraşi district. Based on species, in Oltenia, southern Banat the analysis of 19 descriptions, the and to a lesser degree, in author mentioned the presence of Transylvania. Such plant communities of the group of communities occur less commonly associations Chrysopogon gryllus - in Moldova, Muntenia and Bothriochloa ischaemum, which Dobrogea, where the climate is consisted of 121 species, 104 of colder and drier. which were diverse. The coverage They occur on plateaus and was 80-100 % on 100 m2, and 48 gentle slopes in hill forests with species were identified in this area: common oak, sessile oak and beech. Chrysopogon gryllus (3), The floristic composition is poor, Bothriochloa ischaemum (3), and dominant species are Festuca valesiaca (2), Thymus Chrysopogon gryllus with marschalianus, Koeleria cristata, Bothriochloa ischaemum, Festuca Asperula cynanchica, Galium valesiaca, Koeleria gracilis etc. campanulatum, Galium verum, In Oltenia and Banat, Aira Eryngium campestre, Hypericum elegans, Sieglingia decumbens, perforatum, Inula hirta, Medicago Vulpia myurus and Haynaldia romanica, Potentilla impolita, villosa also occur. The author has Potentilla recta, Securigera varia, mentioned some species of the Stachys officinalis, Teucrium family Fabaceae, which are quite chamaedrys, Crocus reticulatus, uncommon in the studied areas – Stipa capillata, Stipa pennata, Medicago falcata, Dorycnium Bromopsis inermis, Dactylis herbaceum, Trifolium arvense, glomerata, Elytrigia intermedia, T.striatum, T. campestre, Lotus Elytrigia repens, Melica ciliata, Poa corniculatus, Onobrychis arenaria angustifolia, Adonis vernalis, etc., but the following species are Doronicum hungaricum, Lathyrus more frequent: Thymus pannonicus, Pulsatilla grandis, marschallianus, Plantago Pulsatilla nigricans, Viola ambigua, lanceolata, Eryngium campestre, Gagea pusilla, Muscari neglectum, Achillea collina, Euphorbia Caragana mollis, Chamaecytisus cyparissias, Filipendula hexapetala, austriacus, Achillea pannonica, Asperula cynanchica. There are Veronica barrelieri. The species different types of soil in the studied faithful to this community is area – levigated chernozem, Thymus marschalianus, which is rendzina, podzol, marshy soil, part of the association Thymo alluvia, and the determining factor (pannonici) - Chrysopogonetum for the ass. Thymo (pannonici) – grylli Doniţa et al., 1992 (table 1, Chrysopogonetum grylli Doniţa et figure 4, N 11). al. 1992 is the moisture in the soil Puşcaru-Soroceanu et al., and air (table 1, figure 4, N 12). 1963, characterized the distribution

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Table 1 The syntaxonomy of grasslands with Chrysopogon gryllus (L.) Trin., in natural habitats, according to the biogeographic regions

Association or Diagnostic species with the Name of the proposed Bibliographic floristic list of the frequency indicated by the Site association No. source author author

relevés

the map

ic region ic

Number of Number

according to to according (m) Altitude

Biogeograph Glades in forests with Ass. Chrysopogon Thymus marschallianus Willd. Ass. Thymo (pannonici) Şabanova, 1. Quercus pubescens 150-250 17 gryllus – Festuca (T.pannonicus All.) according to – Chrysopogonetum 2014 Willd. (R. Moldova) valesiaca V.Ciocârlan, 1988 (V) grylli Doniţa et al., 1992 Postolache, “Ciumai” Nature 2. 45-93 - Floristic list Thymus marschallianus Willd. - ” - ” - 2017 Reserve (R. Moldova) Slope with steppe Miron et al., vegetation 3. 75-100 - Floristic list Thymus marschallianus Willd. - ” - ” - Steppe 2017 Moscovei v., Cahul distr. (R. Moldova) Plain with steppe Titica Gh., vegetation Lebedenco 4. 90 - Floristic list Thymus marschallianus Willd. - ” - ” - 2017 v., Cahul distr. (R. Moldova) Ghendov et Bugeac Plain, (R. 5. - - Floristic list Thymus marschallianus Willd. - ” - ” - al., 2015 Moldova) Plain with steppe vegetation, Ass. Thymo (pannonici) 6. Titica, 2018 Iordanovca v., 250 - Floristic list Thymus marschallianus Willd – Chrysopogonetum Basarabeasca distr. grylli Doniţa et al., 1992 (R. Moldova)

Steppe Steppe Ass. Dauco guttati Marine sands, Dubyna et al., – Chrysopogonetum 7. Jebriansk (Odessa - 11 Thymus marschallianus Willd - ” - ” - 1996 grylli Popescu, district, Ukraine) Sanda, Doltu, 1980

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Ass. Chrysopogon Babadag Plateau Thymus marschallianus Willd 8. Dihoru, 1970 200-350 21 grylli dobrogicum - ” - ” - (Romania) (II) Dihoru, 1970 Ass. Tymo (pannonici) – Ivan et al., Southern Moldova 9. 300-400 53 Chrysopogonetum Thymus pannonicus All. (II) - ” - ” - 1993 (Romania) grylli Doniţă N. et al., 1992 Ass. Tymo Muntenia, Moldova, (pannonici) – Doniţă et al., Dobrogea, 10. 150-450 - Chrysopogonetum Thymus pannonicus All. (V) - ” - ” - 2005 Transylvania grylli Doniţă N. et (Romania) al., 1992 Slopes with steppe Ass. Chrysopogon

Continental Şabanova, vegetation (Străşeni, gryllus – Thymus marschallianus Willd. 11. 250-350 19 - ” - ” - 2012 Călăraşi, Sadova) (R. Bothriochloa (V) Moldova) ischaemum Puşcaru- Ass. Thymo (pannonici) Ass. Chrysopogon Thymus marschallianus Willd. 12. Sorocianu, Banat (Romania) 200-300 10 – Chrysopogonetum gryllus (IV) 1963 grylli Doniţa et al., 1992

Ass. Chrysopogon Csuros and gryllus Transylvanian plateau 13. Niedermaier, 400-500 - transilvanicum Thymus pannonicus All. (IV) - ” - ” - (Romania) 1966 Csuros, Nedem., 1966

Continental Continental Drăgulescu Ass. Chrysopogono- Southern and caricetum humilis 14. Transylvanian plateau 430-525 12 Thymus pannonicus All. (V) - ” - ” - Schumacher, Zolyomi (1950) (Romania) 2006 1958

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Ass. Thymo “Glodeni Hayfields” pannonici - Oprea and 15. Nature Reserve, - 2 Chrysopogonetum Thymus pannonicus All. (+) - ” - ” - Sârbu, 2005 Vaslui (Romania) grylli Doniţă et al., 1992 Ass. Medicagini (minimae) – Festucetum Ass. Thymo (pannonici) Chifu et al., Transylvania, Banat, 16. 140-380 34 valesiace Vagner Thymus pannonicus All. (I) – Chrysopogonetum 2014 Crişana (Romania) 1941 subass. grylli Doniţa et al., 1992 Chrysopogonetum grylli n.n. Ass. Carici humilis - Chrysopogonetum Ass. Carici humulus - Low 17. Meyer, 1975 North-East Italy grylli Carex humilis Leyss. Vest (2) Chrysopogonetum grylli altitudes - fumanietosum Kojić i 1959 - galietosum Ass. Schoeno Ass. Schaeno (nigricans) (nigricans) -Chrysopogonetum grylli

Continental Chrysopogonetum n. n. The highland Nimis and grylli Schoenus nigricans L. and Ass. Chamaecytiso 18. “Friulian Plain” (NE 50-300 55 Fonda, 1997 Ass. Chamaecytiso Chamaecytiso purpureus (purpureus)- Italy) (purpureus) – Chrysopogonetum grylli Chrysopogonetum n. n. grylli Ass. Teucrio capitati – Ass. Teucrio capitati – Sburlino et al., The north shore of the 19. Sand dunes 18 Chrysopogonetum Teucrium capitatum Chrysopogonetum grylli 2013 Adriatic Sea (Italy) grylli Sburlino et Sburlino et al., 2008 al., 2008

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Ass. Bromo Ass. Bromo (erectus) - Bernhardt et Krk and Cres islands (erectus) 20. - 17 Bromus erectus Huds. (V) Chrysopogonetum grylli al., 2006 (Croatia) Chrysopogonetum Horva 1934 grylli H – ic 60 Ass. Globulario Ass. Globulario

Ilijanic et al., (elongata) - Globularia elongata Hagetschw. (elongata) – 21. West Croatia 200-220 10 1972 Chrysopogonetum (V) Chrysopogonetum grylli grylli Ilijanic et al., ex Terzi Ass. Chrysopogono Ass. Nerieto (oleandri) - Jasprica et al., The shore of the grylli – Nerietum 22. 84-97 5 Nerium oleander L. (V) Chrysopogonetum grylli 2007 Adriatic Sea (Croatia) oleandri Jasprica et Mediterranean Mediterranean Jasprica et al. 2007 al. 2007 Ass. Asphodelo Ass. Asphodelo (ramosus) Fanelli et al., The bank of Buna (ramosus) - 23. 277-484 - Chrysopogonetum Asphodelus ramosus 2015 River (Albania) Chrysopogonetum grylli grylli Horvatić Horvatić (1936) 1957 (1936) 1957 Ass. Bromo Mt. Studina Ass. Bromo (fibrosi) – Jovanovic et High (fibrosi) – Bromus fibrosus Hack. 24. Serpentino, Central 13 Chrysopogonetum grylli al., 2017 altitudes Chrysopogonetum (B. riparius Rehm) Balkans (Serbia) Tatic 1969 grylli Tatic 1969 Ass. Danthonio (alpina) Alpine Alpine Ass. Danthonio – Ivan et al., Oltenia, Transylvania Danthonia alpina Vest. Rchb. – Chrysopogonetum 25. 600 21 Chrysopogonetum 1993 (Romania) (III) grylli Kojić 1959 grylli Kojić 1959 Boscaiu (1970) 1972 Ass. Chrysopogon Ass.Teucrio (capitati)- Pirini et al., Northern-central gryllus – 26. 575-720 26 Teucrium capitatum (69) Chrysopogonetum grylli 2014 mountains of Greece Bothriochloa Sburlino et al., 2008 ischaemum

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Ass. Agrostio North-western Agrostis capillaris L. (A. tenuis Agrostio (capillari) – Pedaşenco et (capillari) – 27. mountains of 970-1400 40 Sibth.) according to V. Chrysopogonetum gryllis al., 2013 Chrysopogonetum Bulgaria Ciocârlan 1988 (V) Kojić 1959 grylli Kojić 1959 Ass. Djordjevic et Mt. Maljen (West 28 510 Chrysopogonetum Agrostis capillaris L. (V) - ” - ” - al., 2012 Serbia) grylli Gajić 1954 Ass. Bromo (erectus) - Tashev et al., Stara Planina Com. Chrysopogon 29. 755 - Bromus erectus Huds. Chrysopogonetum grylli 2010 (Bulgaria) gryllus Horvatić 1934 Ass. Petrarhagio Ass. Petrarhagio (haynaldianae) - (haynaldianae) - Matevski et South-western areas 704 30. 12 Chrysopogonetum Petrarhagia haynaldianae (+) Chrysopogonetum grylli al., 2015 of Macedonia (540-1300) grylli Motevski et Motevski et al., 2015, al., 2015 Kojić 1959 Ass. Chrysopogonetum Northern-central (grylli) serbicum Agrostio (capillari) – Acic et al., areas of the Balkan Agrostis capillaris Leyss 31. 600-1400 - Ass. Chrysopogonetum grylli 2015 Peninsula (Serbia, (cl. 8-9) Chysopogonetum Kojić 1959 Kosovo) pannonicum Stjep.- vest Pannonian Ass. Ass. Carici humulus - Djurdjevic et South Banat, Delibato Chrysopogonetum 32. - - Carex humilis Leyss. Vest (2) Chrysopogonetum grylli al., 2013 sands (Serbia) pannonicum Stjep.- Kojić 1959 vest

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Csuros and Niedermaier, ecological subtypes represented by 1966, presented a monographic xerophilous, xero-mesophilous, study on the distribution of the meso-xerophilous and even species Chrysopogon gryllus, which mesophilous plant communities. was considered by the authors as a However, they are reflected in the Paleo-Mediterranean-Southwest types of habitats by the presence of Asian plant. The ecology of this plant species indicating habitat dominant plant implies that the pH conditions. In Romania, there are should be slightly acid-neutral to over 22 phytosociological units with slightly alkaline, and the analysis of Chrysopogon gryllus, and all of the soil, on which this species them include the species Thymus grows, shows a decrease in the soil pannonicus and match the habitat pH at the upper level (between 6.7 and the ass. Thymo (pannonici) - and 7.73 at 2-10 cm), as compared Chrysopogonetum grylli Doniţa et with the lower one (20-30 cm), by al. 1992 (table 1, figure 4, N 14). 1.3-1.73 units - the ecological limit Oprea and Sârbu, 2005, at which the soil pH ranges from studying the flora and vegetation of slightly acid to alkaline, which the “Glodeni Hayfields” Nature shows that the limit of the range of Reserve, Vaslui (Romania), these phytocoenoses is determined discovered the presence of by the acid pH of the soil and not by phytocoenoses of the ass. Thymeto heat or moisture - ass. Thymo pannonici - Chrysopogonetum grylli (pannonici) - Chrysopogonetum Doniţa et al., 1992 all. Festucion grylli Doniţa et al. 1992 (table 1, valesiacae Klika 1931, ord. figure 4, N 13). Festucetalia valesiacae Br.-Bl. et Drăgulescu and Tx. ex Br.-Bl. 1949, Cl. Festuco- Schumacher, 2006, have analysed Brometea Br.-Bl. et Tx. în Br.-Bl. the grasslands of southern 1949. It is noteworthy that the Transylvania (430-525 m altitude), abundance of Festuca valesiaca attributed by them to the ass. makes up 1-2 units, which is by 3-4 Chrysopogono-Caricetum humilis units lower than the abundance of Zolyomi (1950) 1958, where there Chrysopogon gryllus (table 1, figure are such differential species as 4, N 15). Dorycnium herbaceum, Thymus Chifu et al., 2014, pannonicus, Salvia transsilvanica, mentioned the communities with Dianthus carthusianorum, Carex Chrysopogon gryllus as sub- humilis, Jurnea mollis ssp. association included in the ass. transilvanica and Astragalus Medicagini minimae - Festucetum austriacus. The authors note that the valesiacae Wagner 1941. The sub- Chrysopogoneta, in Romania, are association was characterised as affected by the habitat conditions thermophilic and included the specific to the given area, which are differential species Chrysopogon characterized by the presence of the gryllus (IV) and Dasypyrum

Romanian Journal of Grassland and Forage Crops (2019) 20 70 Lazu Ș. et al villosum (I). The author delimited South Eurasian - 10.5 %; 6. North this sub-association after analysing Eurasian - 12.5 %. In the given area, 34 relevés focused on areas located by cluster analysis, the presence of at 140-380 m altitude and, at the two associations was determined - same time, it reflected a more Schoeno (nigricans) - mesophilic character of the Chrysopogonetum grylli n. n. and vegetation within the described ass. Chamaecytiso (purpureus) association, and Thymus pannonicus Chrysopogonetum grylli n. n. (table Aqll., in the synthetic table, has the 1, figure 4, N 18). frequency - 1. We consider the Sburlino, 2013, studied the presence of the phytocoenoses of phytocoenotic diversity on the north the ass. Thymo (pannonici) - coast of the Adriatic Sea – the grass Chrysopogonetum grylli Doniţa et communities on the slopes of sand al. 1992 (table 1, figure 4, N 16). dunes, as well as those occurring in Meyer, 1975, in his doctoral the more humid areas found thesis, analysed the dry grasslands between them. The authors with Chrysopogon gryllus in north- identified the presence of the ass. eastern Italy, where he identified the Teucrio capitati – ass. Carici humulus - Chrysopogonetum grylli Sburlino et Chrysopogonetum grylli, subass. al., 2008, communities of Fumanetosum with the variants - oligotrophic grasses of the northern Diplachne serotina and tipicum, as forest steppe, found between the well as subass. galietosum with the isotherms 7.5-10.5 °C (table 1, variants tipicum, Leontodon figure 4, N 19). teneciflorus, Vinca minor, al. In the continental Bromion and Orno-Ostrion. The biogeographic area, at altitudes of author noted that such communities up to 1090 m, in the glades of the are also found in Yugoslavia, Carpathian forests of Crivilui-Buzău Hungary and Romania, at low (Romania), Ştefan et al., 2007, altitudes (table 1, figure 4, N 17). mentioned the ass. Taraxaco Nemis et al., 1997, carried (serotini) - Bothriochloetum out the phytogeographic analysis of Burduja et al., 1956, where the steppe-like vegetation on the Chrysopogon gryllus occurred, but hills of the north-eastern Italy with had low abundance. the altitude 50-300 m, where he Imbrea et al., 2008, identified 6 phytogeographic groups mentioned that, on the on the rocky with features relevant to the Balkan- areas of “Cheile Globului” Nature Illyrian and the east alpine regions. Reserve (Mehedinţi) in the south- 1. 34 % include endemic, sub- west of Romania, in forest endemic, Illyrian-Balkan and NW phytocoenoses of the ass. Carpino sub-Mediterranean; 2. South- orientalis - Quercetum cerris Klika European - sub-Mediterranean - 1938, Chrysopogon gryllus (I) and 18.8%; 3. Larger European range - Thymus pannonicus All. (III) 16.6 %; 4. Eastern Pontic - 7.6 %; 5. occurred at altitudes of 100-200 m.

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Lyubenova et al., 2011, serpentines, located between Mt. mentioned the presence of Vujan and Ilijak hill (Serbia), Chrysopogon gryllus on the dominated by Stipa novakii. Danubian Plain (Bulgaria) with Chrysopogon gryllus occurs in the Cotino-Quercetum cerridis. Altitude areas with the ass. Stipetum novakii - 300-1000 m. a. n. (327-360 m altitude, pH 5.5-8), Sopotleva and A postolova where the climate is humid and 2014, determined the presence of warm (t = 10.8 °C). These areas are Chrysopogon gryllus in the surrounded by forests with Quercus transition zone between two cerris, Q. pubescens, Cotinus biogeographic regions - Euro- coggygria etc. Asiatic steppe and Mediterranean In the continental and steppe sclerophytic vegetation. The bioregions, Chrysopogon gryllus, at average altitude in this area is 111- low altitudes, grows in grass 356 m. Here, the species communities, as a dominant species, Chrysopogon gryllus is part of the but in the zone of thermophilic ass. Chrysopogonetum grylli alpine forests, it is a species with (Bulgaria). low abundance, since it prefers Feoli et al., 1980, analysed areas with more light, such as the abandoned grasslands of the glades or sparse forest vegetation. Karst region (Italy), with Quercus Bernhardt et al., 2006, in pubescens, Q. petraea, Fraxinus their study on the xerophytic ornus, Ostrya carpinifolia, grasslands on the NE Adriatic Juniperus communis and Cotinus islands Cres and Krk (Croatia), coggygria, where the populations of mentioned the presence of plant Chrysopogon gryllus were communities of the ass. Bromo recovering. The authors found that (erectus) – Chrysopogonetum grylli some mesophytes had disappeared H-ic 60, on neutral to alkaline soil, from that area and were replaced by in the humid habitat of the all. xerophytes. Scorzonerion villosae H-ic 49; ord. Lonati et al.,2010, studied Scorzonero – Chrysopogonetalia the syntaxonomy and synecology of grylli H-ic 60; Cl. Festuco- the communities with Erica cinerea Brometea (table 1, figure 4, N 20). L. in the Alps (north-western Italy). Ilijanic et al., 1972, At 480-670 m altitude, the edaphic described the grasslands containing group Chrysopogon gryllus with Chrysopogon gryllus, in the western characteristic grasses of cl. Festuco- areas of Croatia, where there are dry Brometea occurs and is surrounded grasslands of the Cl. Festuco- by Mediterranean forests (Quercus Brometea ass. Globulario pubescens, Q.petraea, Castanea (elongata) - Chrysopogonetum sativa, Fraxinus ornus, Erica grylli, with a floristic composition cinerea). of sub-Mediterranean type. The Kabas et al., 2013, described given region, with grassy vegetation the grasslands found on rocky dominated by Chrysopogon gryllus,

Romanian Journal of Grassland and Forage Crops (2019) 20 72 Lazu Ș. et al is found at an altitude of 200-220 m. (1936) 1958, all. Chrysopogono- The soil is brown at the uppermost Saturejion Horvat & Horvatić in horizon, but becomes red or lightly Horvatić 1934, ord. Scorzonero - yellow at deeper profiles, at about Chrysopogonetalia Horvatić & 40 cm (table 1, figure 4, N 21). Horvat in Horvatić 1963, cl. Jasprica et al., 2007, on the Festuco-Brometea Br.-Bl. et Tuxen Adriatic coast, indicated the ex Klika et Horvatić 1963 (table 1, presence of grass communities of figure 4, N 23). the ass. Chrysopogono grylli – Koci et al., 2017, studied the Nirietum oleandri Jasprica et al., flora and vegetation of the 2007, all. Rubo ulmifoli – Nerion southwest part of Vlora Bay oleandri, ord. Tamaricetalia, cl. (Albania), which was represented Nerio-Tamaricetea (Braun-Blanquet mainly by three families being et Bolos 1957). These communities respectively Poaceae - 28 species, are characterized as resistant to long Asteraceae - 16 species and periods of drought and include Fabaceae - 13 species, the components of the ord. Quercetalia Mediterranean elements ilicis and Scorzonero- predominated - 49 %. On cliffs, Chrysopogonetalia. According to there are forests of Quercus the authors, they occur all along the pubescens, Q. ilex with the Mediterranean coastline (table 1, herbaceous layer consisting mostly figure 4, N 22). of Chrysopogon gryllus, Dactilis Fanelli et al., 2015, carried glomerata and Asparagus out a research on the vegetation of acutifolius. The woody layer of the Buna River-Velipoja Protected forests, in the studied area, is Landscape (Albania). On carbonate dominated by Cupressus substrate, there are perennial grasses sempervirens (90- 95 %). specific to the Mediterranean Zupančič et al., 1987, region. The author indicated the analyzed the relief of Slovenia from presence of communities of the ass. phytogeographic point of view and Asphodelo (ramosus) – mentioned the presence of Chrysopogonetum grylli Horvatić grasslands with Chrysopogon (1936) 1957, described in Dalmatia gryllus in the pre-alpine zone, where by Horvatić in 1958 and 1963, and sub-Mediterranean elements then – in Albania (at 83-152 m and predominated (Sesleria autumnalis, 277-484 m altitude), as well as in Ruscus aculeatus, Chrysopogon the sparse forests of Quercus gryllus, Luzula nivea) in Ostrya - trojana. There are no Fagetum forests. Chrysopogoneta in areas with Pipenbaher et al., 2008, Mediterranean climate, where the studied the sub-Mediterranean grasslands consist mainly of Stipa Illyrian grass vegetation of bromoides. The authors indicated (Slovenia) in two totally different the association Asphodelo - sites - with limestone and flysch Chrysopogonetum grylli Horvatić substrates. In this area, they

Romanian Journal of Grassland and Forage Crops (2019) 20 73 Lazu Ș. et al identified the ass. Danthonia planted with Pinus halepensis and (alpinae) - Scorzonetum villosae. P. pinea (200-300 m altitude). The Chrysopogon gryllus is a soil, at these sites, is shallow, component of the grass vegetation eroded, covered mostly by forest in the sites with lighter substrate and vegetation - Quercus coccifera and has a higher frequency. The soil pH, Q. pubescens, the herbaceous layer in such areas, is neutral to slightly is dominated by Chrysopogon acid; the average temperature is 8- gryllus. 12 °C. In the Mediterranean Čarni, 2003, mentioned that, biogeographic region, Chrysopogon on the Adriatic coastline, in the gryllus is present as a dominant north-west of Croatia (264-504 m species in grassy phytocenoses, altitude), at the edge of forests of forming communities with diverse Quercus pubescens with Cotinus local species of xeric grasslands. coggygria, there were grass Maestre and Cortina, 2002, communities with Chrysopogon mentioned the spatial distribution of gryllus of the ass. Knautio illyricae the components of the – Melampyretum carstinise as. Mediterranean semi-arid steppe nova. vegetation with Stipa tenacissima L. Karmiris et al., 2010, studied - as dominant species and the diet of animals living in Brachipodium retusum (Pers.) P. Mediterranean shrublands. At 360- Beaw. - co-dominant species, which 520 m altitude, in central represent 73 % of the total perennial Macedonia, Greece, nearly 50 % of vegetation cover. The steppes of the the area is covered by scattered south-eastern Spain do not include kermes oak (Quercus coccifera) such grass species as Chrysopogon stands, and the herbaceous layer is gryllus and Bothriochloa dominated by Chrysopogon gryllus ischaemum (altitude - 460 m, and Dichanthium ischaemum. Such temperature - 15.8 °C, precipitation species as Festuca valesiaca, - 388 mm), because they are found Cynodon dactylon, Bromus at the south-western boundary of the tectorum, Anthoxanthum odoratum, xeromesophytic grasslands. Dactylis glomerata also occur in Jovanovici Sl. et al. 2017 this area. The Fabaceae family is (Serbia) demonstrated the represented by Trifolium stellatum, distribution of the grass Medicago polymorpha, Trifolium communities with Chrysopogon arvense, T. campestre, Vicia cracea. gryllus on limestone serpentines in Other species – Malva sylvestris, the Balkan Mountains, at high Cichorium intybus, Sanguisorba altitudes, where they identified minor, Capsella bursa-pastoris, grassland vegetation of the ass. Leontodon hispidus, Taraxacum Bromo (fibrosi) – Chrysopogonetum officinale, Sonchus oleraceus. grylli Tatici 1969 (table 1, figure 4, Ganatsas et al., 2004, N 24). described some areas, in Greece,

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Ivan et al., 1993, based on subjected to the cluster analysis, relevés sampled in the area of the four plant communities were Transylvanian Plateau (Romania), at distinguished in the area, namely the altitude of 600 m, emphasized Artemisia campestris-Dasypyrum the specific floral composition of villosum, Chrysopogon gryllus- grasslands with Chrysopogon Bothriochloa ischaemum, Satureja gryllus, accompanied by Danthonia montana-Artemisia alba and Stipa alpina (III) - an oligotrophic capillata-Koeleria macrantha. xeromezophytic grass species, These communities were classified which grows on slightly acid soil. in cl. Festuco-Brometea, ord. Other species that occur in this area: Astragalo-potentillietalia. The Festuca valesiaca (IV), Asperula authors consider these communities cynanchica (II), Achillea setaceae steppe-like or secondary grasslands, (II), Trifolium montanum (IV), with some species interpreted as Teucrium montanum (III), Eryngium relicts of the Tertiary period, and campestre (III), Teucrium the secondary grasslands as a result chamaedrys (III), Filipendula of degradation of forests with vulgaris (III), Salvia pratensis (IV), Quercus trojana. Steppe species Euphorbia cyparissias (III), occur in open oak woodlands or Sanguisorba minor (II), Dianthus shibljak-type shrublands. carthusianorum (IV), Trifolium The authors also mention campestris (III), Prunella laciniata that low-altitude dry grasslands of (III), Thymus globarescens (II), north-central Greece are located Anthylis vulneraria (II), Phleum near the south-eastern European montanum (II) etc. The zonal margins of the vegetation class component of these Festuco-Brometea. The authors Chrysopogoneta is missing – concluded that the steppe-like Thymus pannonicus. These grasslands around the Lakes of communities of savanoid grasses Vegoritida and Petron are clearly occur at the northern border of this secondary steppe grasslands but it is biogeographic region and contact rather likely that populations of with the dry steppe grasslands of cl. steppe species have occurred Festuco-Brometea. The authors continuously in deciduous accepted to include such plant woodlands and shrublands, in communities in the ass. Dantonia landscapes that “were probably (alpina) - Chrysopogonetum grylli never densely wooded throughout Kojić 1959; Boșcaiu (1970) 1972 the Holocene”. Chrysopogon gryllus (table 1, figure 4, N 25). and Bothriochloa ischaemum are Pirini et al., 2014, presented present in all clusters mentioned by a study on the steppe-like grassland the authors, and this fact suggests a vegetation in the hills around the close connection of these species lakes of Vegoritida and Petron, in with the Balkan (alpine and north-central Greece. Based on 245 Mediterranean) habitat. After relevés of vascular plant species, analyzing the synthetic table

Romanian Journal of Grassland and Forage Crops (2019) 20 75 Lazu Ș. et al presented by the authors, we Djordjevic et al., 2012, highlighted the dominance of the described communities of the ass. species Thymus sibthorpii in the Chrysopogonetum grylli grass communities of the ass. Gajić 1954; all. Chrysopogono - Chrysopogon gryllus - Bothriochloa Danthonion calycinae Kojić i 1959; ischaemum Sburlino et al., 2008. ord. Festucetalia valesiaceae Br.- The authors conclude that the dry Bl. et Tx. 1943, cl. Festuco- grasslands of Greece are secondary Brometea Br.-Bl. et Tx. 1943, found steppes, formed because of at 510-490 m altitude. The intensive grazing, in the sparse frequency of Agrostis capillaris - forests of Quercus trojana as well (V) (3 relevés), ass. Agrostis as in the Mediterranean forests with (capillaris) - Chrysopogon grylli Q.coccifera, which has led to the Kojić, 1959 (table 1, figure 4, N creation of shrublands of the 28). shibljak type (wooded deciduous Tashev et al., 2010, pastureland of the subcontinental contributed to the study of habitat parts of the Balkans and the Black diversity in the Western Stara Sea area). Under postglacial and Planina Mountains (Bulgaria), current climatic as well as land use where they identified the biotope of (grazing) conditions in our area, and sub-Pannonian steppe with depending on the degree of their Chrysopogon gryllus and demand for light, steppe species Bothriochloa ischaemum. One of may have occurred either in open the most common species for these oak woodlands or in various kinds steppes is Bromus erectus with ass. of shibljak-type degradation stages Bromo (erectus) - (table 1, figure 4, N 26). Chrysopogonetum grylli Horvatić Pedashenko et al., 2013, 1934 (table 1, figure 4, N 29). carried out a research on the Matevski et al., 2015, ecological diversity and studied the syntaxonomy of the syntaxonomy of the dry grasslands rocky grasslands on carbonate of NW Bulgarian Mountains. They bedrocks in the west and southwest identified, at high altitudes, on of the Republic of Macedonia, limestone substrate with alkaline where they have identified the ass. pH, plant communities of the ass. Petrorhagio haynaldianae - Agrostio (cappilari) - Chrysopogonetum grylli n.n., in Chrysopogonetum grylli Kojić areas with the average temperature 1959; all. Chrysopogono grylli - 11.2 °C and altitude from 540 m to Danthonion calycinae Kojić 1959; 1300 m. Chrysopogon gryllus and ord. Brachipodietalia pinnati Bothriochloa ischaemum occur in Korneck 1974; cl. Festuco - arid grasslands all over the Balkans Brometea Br.-Bl. et Tx. et Klika et tTable 1, figure 4, N 30). Hadac 1944 (table 1, figure 4, N Velev, 2005, analyzed the 27). dynamics of grassland vegetation in the Lovech province (Bulgaria) over

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15 years. The average altitude of the Kosovo), which include the north- sites studied by the author is 1000- central Balkan Peninsula (south- 1200 m. In the areas covered mainly eastern Pannonian Plain). They are by forest vegetation with Fagus found along the Danube, Tisa, Sava, sylvatica and Quercus cerris, there Begej and Tamiš rivers. Pannonian are communities of herbaceous dry grasslands form xerophilous plants with Agrostis capillaries, grassy communities at various Pteridium aqulegium (L.) Kechn. altitudes (from low altitudes in the and Festuca valesiaca, which occur floodplain of rivers to mountain in large areas in the Bulgarian part areas of 600-1400 m) and on of the Balkan mountain range. The various types of soil: chernozem, phytocoenoses with Chrysopogon alkaline silicates, limestone with gryllus are secondary communities neutral pH, slightly acid sands, such in the forest vegetation. as loess and Pleistocene sediments Todorova et al., 2010, and deposits. The authors consider mentioned that, in the Bosnek that in the Central Balkans (Serbia, village (western Bulgaria), 20 % of Kosovo) traditional Pannonian dry the territory is covered by dry grasslands are communities of cl. grasslands, at 830-1500 m altitude, Festuco-Brometea, ord. Festucetalia on limestone substrate on karst vaginatae. By analysing the rocks. In this area, the authors have database made up of 1897 relevés identified phytocoenoses of the with 1323 species of vascular association Bromo moesiacae - plants, 11 clusters have been Stipetum epilosae a.n. all. Saturejon identified. Chrysopogon gryllus has montanae Horvat 1962, ord. been mentioned as dominant and Festucetalia Br.-Bl. et Tuxen 1943, constant species in the Cluster 2, on cl. Festuco-Brometea Br.-Bl. et the hills in the zone of Quercus Tuxen 1943. Chrysopogon gryllus frainetto and Q. cerris forest (table and Bothriochloa ischaemum occur 1, figure 4, N 31). in 5 of the 38 analyzed relevés. Djurdjević et al., 2013, In the alpine biogeographic demonstrated the allelopathic region, there are forest belts and effects of Chrysopogon gryllus L. in alpine grasslands with Chrysopogonetum Pannonicum Mediterranean and sub- Stjep.-Ves. steppe community in the Mediterranean vegetation, where region Deliblato Sands of Southern Chrysopogon gryllus occurs in Banat (Serbia), where they detected habitats with slightly acid soils and leaching of phenolic acid and alpine grasses - Agrostis capillaris, studied its impact on the Danthonia alpina, Bromus fibrosus, germination capacity of the seeds of Petrorhagia haynaldiana etc. Trifolium arvense and Poa pratense, Aćić et al., 2015, described which had low abundance and cover the ecology, biodiversity and in the ass. Carici humilis- classification of arid grasslands in Chrysopogonetum grylli Kojić the Central Balkans (Serbia, 1959. The abundance of Carex

Romanian Journal of Grassland and Forage Crops (2019) 20 77 Lazu Ș. et al humilis is (2) (table 1, figure 4, N present in the communities of 32). meadow grasses. Szollát et al., 2007, in their Hagyó, 2003, in an article on article on the vegetation of sandy the vegetation of marsh meadows of areas of Gödi láprét, Hungary, Zákányszék (Hungary), mentioned mentioned semi-natural steppes the presence of Chrysopogon dominated by Bromus erectus and gryllus in ass. Agrostio-Caricetum grasslands dominated by distantis and ass. Astragalo Chrysopogon gryllus, austriacae - Festucetum sulcatae, Brachypodium pinnatum and considering them transitional types Equisetum ramosissimum, with such of communities, from arid to humid abundant species as Stipa capillata, ones. Festuca rupicola, F. pseudovina, The analyzed meadows from Koeleria cristata, Carex the Pannonian Plain, containing caryophylla, Dactylis glomerata and Chrysopogon gryllus, commonly Centaurea sadleriana. In the occur in meadow habitats as studied grasslands found in the components of the ass. Agrostio Danubian Plain, of the Pannonian (alba) - Caricetum distans and biogeographic region, Chrysopogon Astragalo austriaceae - Festucetum gryllus has low abundance. sulcatae. The soil in this area is Dobolyi, 2002, studied the sandy. changes that occurred in the The Pannonian vegetation of grasslands found on biogeographic region comprises dolomite rocks in Csíki-Hegyek grasslands with Chrysopogon (Budaörs, Hungary), between 1996 gryllus, occurring at the border with and 2001. During that period, the the xeric steppes from cl. Festuco- proportion of species with sub- Brometea, thus, the analyzed lists Mediterranean, Pontic, Balkan and highlight the xerophytic character of Sarmatian elements increased, and the sites with Bromus erectus, as the distribution of phytosociological well as the expansion in the elements of Festucetalia valesiacae meadow habitats with Agrostis alba order, Festuco-Brometea class, such or Iris sibirica. as Chrysopogon gryllus, have The "Black Sea" experienced an increase in biogeographic region includes the abundance. Balkan and Anatolian coasts. In this Salamon-Albert and Lörincz, area, there are favourable conditions 2010, studied the vegetation of for the growth and expansion of the meadows, focusing on the ass. savannoid species Chrysopogon Iriditetum sibiricae, which gryllus, which is not dominant in contained plant communities that plant communities, but is part of the were specific to Hungary. The list grassland and forest phytocoenoses of components of this association occurring in the given region. also included Chrysopogon gryllus, Tzonev et al., 2005, studied which was very scattered, but still the psammophytic vegetation (on

Romanian Journal of Grassland and Forage Crops (2019) 20 78 Lazu Ș. et al dunes) of the Bulgarian Black Sea of light make it possible for this coast, where they detected, on species to grow in the forest areas. shifting dunes, the presence of grass The Anatolian biogeographic communities of the ass. Aurinio region is a plateau with uichtritziani - Artemisietum silvopastoral vegetation, which campestris and ass. Cakilo euxinae - includes various altitudes Salsolietum rutenicae, which favourable to forests, grasslands occured on mountain slopes in and forest steppes. Bulgaria and Turkey and differed Korkmaz et al., 2011, in the from those found on Dobrudzha study on the vegetation of the coast and “Golden Sands" resort in Kizilirmak valley (Turkey), the more savannoid floristic identified the presence of steppe composition. vegetation, and Chrysopogon Karaer et al., 2010, gryllus is present everywhere in the presented the results of a research forests of cl. Quercetea ilicis Br.-Bl. on the phytosociological and et de Bolos G. Vayreda 1950, where ecological structure of it has high frequency (V) in 10 Mediterranean enclaves along the relevés, at 300-400 m altitude. The stream valleys in inner parts of author mentioned 10 associations Black Sea region (Turkey). At the containing Chrysopogon gryllus: 1. altitude of 350-800 m, there are ass. Spyreaeo crenatae - Oleetum degraded forests of Quercus sylvestris Karaer et al. 2010. 2. Ass. coccifera, Pinus brutia, Arbutus Buxo sempervirentis - Arbutetum andrachne, A. unedonis and Olea unedonis Karaer et al. 2010, located sylvestris. They form forest at 300-400 m altitude, Chrysopogon communities with Chrysopogon gryllus (h = 70 cm) has a coverage gryllus, which occurs in the of 20 %, frequency - (I). 3. Ass. following associations: Siderito Scutellaria pinnatifidae - dichotomae - Quercetum cocciferae, Juniperetum excelsae Korkmaz., Cotino coggygriae - Arbutetum Engin, Kutbay et Galcin, altitude - andrachnes, Spiraeocrenatae - 350-600 m, height of Chrysopogon Oleetum sylvestris, Paliuro spinae- gryllus plants - 45-60 cm, coverage christi -Fontanesietum - 20-25 % and frequency - (II). Ass. philliraeoides, Buxo sempervirenti - Cotino coggygrie - Pinetum brutiae Arbutetum unedonis, Querco Korkmaz, Engin, Kutbay et Galciu, infectoriae - Pinetum brutiae. The the frequency of Chrysopogon grasses in these forest communities gryllus - (II), coverage - 20-25 %. h are 30-40 cm tall, their coverage is = 50-70 cm. In the Anatolian about 25-40 %, slope exposure - steppes, at the altitude of 550- southern or south-western. In these 650 m, Chrysopogon gryllus has associations, Chrysopogon gryllus is high frequency, together with Stipa in minority and its range includes arabica. It is part of the ass. Linario areas with slightly acid forest soils, carifoliae - Astragaletum but the temperature and the amount microcephali Korkmaz, Angin,

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Kytbay et Galcin. At the altitude of grasses - Chrysopogon montanus et 1250-1300 m, there are no al. (red soil, precipitation - 500- Chrysopogon gryllus plants in the 550 mm, altitude - 305-701 m, Anatolian steppes. It is obvious that depth of groundwater - 16-22 m). Chrysopogon gryllus occurs in areas Joshi et al., 1964, studied the with Mediterranean climate on grasses and sedges of certain areas alkaline soil on carbonate substrate. in Jhunjhunu district (Rajastan, Kaya et al., 2013, carried out India). The research was conducted a synecological and syntaxonomical in low-lying areas, sandy areas and research on the overgrazed steppe hilly areas with Chrysopogon vegetation on Arat Mountain gryllus, which was found only in (Turkey). Chrysopogon gryllus is hilly areas, where it formed present in the ass. Minuartio longitudinal belts over hilly slopes. formosae - Astragaletum Sundriyal, 1995, in the study diphtheritae, at 710-720 m altitude, on grassland forage production and with the frequency I, coverage - 70- management in the western 75 %, on slopes with northern Himalaya, mentioned the presence exposure and inclination of 15-20 °. of Chrysopogon gryllus as dominant It is part of the cl. Astragalo- species in the alpine zone (altitude > Brometea. Therophytes predominate 3000 m, t < 6-10 °C, rainfall < in these grasslands, as a result of 1000 mm; dry months - 5-6, winter overgrazing. snow - 6-7 months), together with Ünal S. et al. evaluated the Agrostis canina, Dactilis glomerata, vegetation of the Kayseri Province, Koeleria cristata, Poa pratensis, Central Anatolia (Turkey). The Phleum alpinum, Danthonia authors studied the steppe jacquemontii et al., climax species - vegetation at 1080-2264 m, where Rhododendron, Juniperus, the average temperature was Cotoneaster, Artemisia, Lonicera, 10.2 °C. The succession of grass Hippophae. The author mentioned species from these grasslands was the presence of Chrysopogon studied. The authors determined that gryllus in the montane zone Chrysopogon gryllus was in decline, (altitude = 2000-3000 m, t < 10 °C, but Festuca valesiaca, the dominant rainfall < 1000 mm, winter snow - species of xeric grasslands, was 3-4 months), climax species in expanding. forests - Betula utilis, Abies Qureshi et al., 1964, studied pindrow, Taxus baccata, Pinus the forest soils of the arid and semi- excelsa, Acer caesium. arid regions of India. They found Neamsuvan et al., 2009, that dry forests on red and yellow mentioned, for the first time, the soils predominated in the arid zone presence of Chrysopogon gryllus (Accacia senegal, Prosopis (L.) Trin. in a new location – spicigera et al.), the most common Vetiveria (Thailand), characterized species of shrubs were Tamarix by evergreen forest vegetation, at dioica, Zizyphus rotundifolia et al., the altitude of 1250-1500 m with

Romanian Journal of Grassland and Forage Crops (2019) 20 80 Lazu Ș. et al degraded grasslands, which are Rhododendron arboreum, often burned. Terminalia etc. Singh et al., 2015, studied Terzi, 2015, analyzed the the vegetation of degraded oak vegetation of the dry grasslands of forests (Quercus floribunda and Q. ord. Scorzonero-Chrysopogonetalia, leucotriphoros), at 1500-2350 m and modified its nomenclature, as altitude, in the Garhwal Mountains being strictly sub-Mediterranean, of (Himalaya). They mentioned that the order Scorzonerietalia villosae, Chrysopogon gryllus occurred in occurring in Albania, Bosnia and unaffected forests as well as in those Herzegovina, Bulgaria, Italy, affected by anthropogenic Kosovo, Montenegro, Slovenia and disturbance. Serbia. The vegetation of this order In the Central Himalaya, occurs in the western Balkans till Chrisopogon gryllus occurs in the the pre-alpine area. The author montane zone - 2000-3000 m mentions the species of the order as altitude (t = 15 °C; rainfall - 1000- meso-xerophilic, but included in cl. 2000 mm; dry months - 2-3), sub- Festuco-Brometea and considers montane – 1200-2000 m altitude (t that such communities are closer to = 15-20 °C; rainfall - 1000- the ecological requirements of 2500 mm; rainy days - 100-150; dry Scorzonera villosa, which occurs at months – 2-3) and sub-tropical zone lower altitude. - altitude < 1200 m (t > 20 °C; Romeo Di Pietro et al., rainfall - 1000 mm; dry months - 4- 2015, analyzed the nomenclature 5, sunny days < 100), together with and the syntaxonomy of dry Bothriochloa pertusa, Cynodon grasslands (steppes) in Europe. dactylon, Arundinella, Phragmites They mentioned the presence of etc., climax species - Quercus grassy communities of all. incana, Q. dilatata, Q. Chrysopogono-Danthonion in the semecarpifolia, Cedrus, Picea, Balkans and attributed the west Pinus, Terminalia etc. European grassland vegetation to In Eastern Himalaya, ord. Brometalia, cl. Festuco- Chrysopogon gryllus occurs in the Brometea. sub-montane zone - 1200-2000 m Redzic, 1999, carried out an altitude (t = 15-20 °C; rainfall - analytical study on the syntaxonomy 2000 mm; dry months - 3-4, humid of grasses of Cl. Festuco-Brometea climate) and sub-tropical - altitude < Br.-Bl. et Tx. 1943 from the 1200 m (t - 20 °C; rainfall Balkans and concluded that “they > 2000 mm; dry months <4), are significantly different from together with Agrostis, those in the western and central Bothriochloa, Festuca, Phalaris, Europe, and even from those in the Phleum, Phragmitis, Cynodon, eastern Europe, by the presence of Arundinella etc. with regional grass numerous species which belong to species, in forest vegetation - with the Balkan, Illyrian and the Quercus lamellosa, Magnolia, southeast Europe floristic element”.

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This phytocenotic class (Festuco- one - Festucetalia valesiacae Br.- Brometea) includes so many Bl. and Tx. 1943 - in xeric habitats. different orders – Scorzoneretalia Therefore, the savannoid villosae, Festucetalia valesiacae, grasslands Chrysopogono- Brometalia erecti and Astragalo- Bothriochloetea n.n. comprise the Potentilletalia, with numerous range of grasses in the phytocoenotic specifications, Mediterranean and sub- atypical to the class of grasslands in Mediterranean climate zone and the western and central Europe. This dry grasslands of the natural range situation emphasizes the need for of the dominant species further revision, according to Bothriochloa ischaemum and Redzic S., of the status of the class Chrysopogon gryllus, occurring in Festuco-Brometea, under mainly Europe and Asia, within the limits xeric conditions, encountered at of the isotherm 8-10.5 °C and different altitudes. The author also average amounts of precipitation of mentions that, by syntaxonomically 500-1000 mm, and all that would differentiating the xerophilous correspond to the habitat expressed vegetation from the by ecological formula - U1-2Tr1,R7- xeromesophilous vegetation, the 8T4,5H1 (Lazu, 2014). floristic composition is specified by This hypothesis requires the presence of hemicryptophytes further vegetation inventory, more and chamaephytes with more than in-depth studies on the current 35 % and of the Illyrian, Balkan and situation of savannoid grasslands south-east European floristic and more conclusive evidence in elements, and attributes the terms of paleobotany, plant grasslands with Chrysopogon geography and population genetics, gryllus to two orders of cl. Festuco in order to confirm the Brometea: the first one - syntaxonomic position of Scorzoneretalia villosa Horvatic communities with Chrysopogon 1975, with species of Mediterranean gryllus and Bothriochloa and sub-Mediterranean grasses in ischaemum. meso-xeric habitats, and the second

CONCLUSIONS

Chrysopogon gryllus, within Sea and Anatolia, it is an ordinary its wide range (continental, steppe, component of regional alpine, Mediterranean and phytocoenoses. Pannonian biogeographic regions), In the present grasslands forms discrete and stable with Chrysopogon gryllus, phytocoenoses, as dominant species occurring in the plains and glades and is accompanied by species of with steppe vegetation of the forests local grasses, and in the with Quercus pubescens and biogeographic regions of the Black Cotinus coggygria (steppe and

Romanian Journal of Grassland and Forage Crops (2019) 20 82 Lazu Ș. et al continental biogeographic regions), communities, and at low altitudes, it there are stable communities of the forms communities with other association Thymo (pannonici) - species of the genus Chrysopogon. Chrysopogonetum grylli Doniţă et The grasslands with al., 1992. Chrysopogon gryllus and In the Mediterranean Bothriochloa ischaemum spread biogeographic region, at an altitude over a large area, compactly of 480-670 m, within the range of accompanied by a group of their Mediterranean forests with Quercus faithful grasses, separating the petraea, Q. pubescens, Castanea xerophilous grasses of the cl. sativa, Fraxinus ornus, Erica Festuco-Brometea from savannah cinerea, with numerous shrubs - grasslands, which are more Rosa sp., Corylus avellana, Sorbus thermophilous and more humid, aria, Juniperus communis, Berberis reminiscent of the times when they vulgaris, Rosa canina etc. and were grazed by herds of herbivores grasses, such as Chrysopogon in the savannahs of the Tertiary era. gryllus and Bothriochloa The grassland communities ischaemum etc., there are isolated with Chrysopogon gryllus are more grasslands used as pastures, and in common on the European continent some sectors, the trees and shrubs (continental, steppe, Pannonian, have degraded to extinction. Mediterranean and alpine In the alpine biogeographic biogeographic regions), but they are region (500-1500 m altitude), there less common, or, at least, we have are dry grasslands with not found any bibliographic sources Chrysopogon gryllus as dominant mentioning their presence in North species, and in degraded forests, it Africa, Middle East and Southeast occurs as an ordinary component. Asia, and the habitat of these In the eastern part of its phytocenoses is included, in terms range, in the Himalaya Mountains, of climate, in the isotherm 7.5- Chrysopogon gryllus grows at 10.5 °C (Sburlino et al., 2013) and altitudes up to 3000 m and above, the precipitation limit > 500 mm where it forms monodominant (Sundriyal, 1995).

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