12 ACTA BOT. CROAT. 81 (1), 2022 Acta Bot. Croat. 81 (1), 12–22, 2022 CODEN: ABCRA 25 DOI: 10.37427/botcro-2022-027 ISSN 0365-0588 eISSN 1847-8476 An Asphodelus ramosus dominated plant community in Montenegro: fringe or grassland? Milica Stanišić-Vujačić¹*, Danijela Stešević¹, Sead Hadžiablahović², Danka Caković¹, Urban Šilc3 1 University of Montenegro, Faculty of Natural Sciences and Mathematics, Džordža Vašingtona bb, 81000 Podgorica, Montenegro 2 Environmental Protection Agency of Montenegro, IV Proleterske 19, 81000 Podgorica, Montenegro 3 ZRC SAZU, Institute of Biology, Novi trg 2, 1000 Ljubljana, Slovenia Abstract – Our phytosociological study in Montenegro (Ćemovsko polje) deals with the syntaxonomy of arid grass- lands in the Adriatic region and, in particular, different interpretations of plant communities dominated by Asphodelus ramosus. The main aims of this study were to contribute to knowledge of the composition of dry grasslands dominated by Asphodelus ramosus in Montenegro and to compare instances of Asphodelus ramosus dominated vegetation along the Adriatic. Our vegetation dataset included 82 phytosociological relevés: 17 from our recent field work and 72 relevés of South European Asphodelus ramosus communities. Ordination analysis (NMDS) was used for comparison of Asphodelus ramosus dominated communities in the Adriatic region. The Asphodelus ramosus community from Montenegro was classified into Bromo erecti-Chrysopogonetum grylli. The analysis revealed two distinct vegetation groups: grassland communities of the vegetation class Festuco-Brometea from Montenegro, Croatia and Albania, and edge vegetation of the new class Charybdido pancratii-Asphodeletea ramosi from Italy. Comparison with similar vege- tation types shows high similarity with associations on the eastern Adriatic coast, where they are treated as grassland communities belonging to the alliance Chrysopogono grylli-Koelerion splendentis, order Scorzoneretalia villosae, class Festuco-Brometea. Keywords: Asphodelus ramosus, Bromo erecti-Chrysopogonetum grylli, dry grasslands, Festuco-Brometea, Montenegro Introduction Secondary dry grassland communities in the Mediter- ranean have a zoo-anthropogenic origin; they have devel- oped over centuries or even millennia of traditional land use, featuring practices such as mowing, grazing, temporary abandonment of arable fields, and/or other disturbance re- gimes (Apostolova et al. 2014, Valkó et al. 2018). Mediter- ranean and sub-Mediterranean dry grasslands are consid- ered to be among the floristically richest vegetation types (Apostolova et al. 2014) and, at the same time, a very impor- tant habitat for endangered, rare and endemic species, so they are included in the list of Habitats of European Interest (Council Directive 92/43/EEC on the conservation of natu- ral habitats and of wild fauna and flora, 1992), as well as the European Red List of Habitats (Janssen et al. 2016). Com- pared to other parts of Europe, dry grasslands of the Balkan peninsula are still well preserved (Apostolova et al. 2014). The Balkan peninsula is well known for its rich flora and well-preserved vegetation, because it was a glacial ref- uge for animal and plant species (Griffiths et al. 2004). Its biodiversity is considered to be among the highest in Eu- rope (Apostolova et al. 2014). The peninsula is character- ized by the presence of a broad spectrum of dry grasslands (e.g., Horvat et al. 1974, Jovanović et al. 1986, Apostolova et al. 2014, Terzi 2015, Aćić et al. 2015, Matevski et al. 2018). In some Balkan countries (Slovenia, Serbia, Bulgaria, Cro- atia, North Macedonia), dry grassland vegetation has been intensively researched (Apostolova et al. 2014, Matevski et al. 2018). In contrast to these countries, research into this type of vegetation in Montenegro has considerable discon- tinuity, and comprenhensive studies, especially in the sub- Mediterranean and Mediterranean region, are rare (Pulević and Bulić 2012). * Corresponding author e-mail: milicas@ucg.ac.me ASPHODELUS RAMOSUS DOMINATED PLANT COMMUNITY ACTA BOT. CROAT. 81 (1), 2022 13 In our study, we focused on research into Asphodelus ramosus-dominated vegetation. Asphodelus ramosus is a species native to the Mediterranean and Macaronesian re- gions: southern Europe, northern Africa, the Middle East, Mediterranean islands and Canary Islands. It can be found in forest edges and grasslands, generally on basic soils of a certain depth, forming very dense populations in grazed areas, from the coast up to 1000 m, occasionally reaching 2150 m in the mountain systems of North Africa (Diaz- Lifante and Valdés 1996). It is unpalatable for grazing animals. Research into plant communities dominated by Asphode- lus spp. (and Asphodelus ramosus in particular) has become very intensive in recent years, especially in the western and central Mediterranean (Allegrezza et al. 2015, Biondi et al. 2016, 2017). In the Balkan peninsula, similar vegetation has been described from Albania (Fanelli et al. 2015) and Croatia (Horvatić 1934, 1939, 1963, Šegulja 1969, 1970, Hećimović 1984, Jasprica and Ruščić 2013, Jasprica et al. 2016). Today, there are different opinions about the syntaxo- nomical classification of Asphodelus ramosus dominated vegetation in Europe and its position in the landscape (as grasslands or fringe (saum) communities). In the eastern Adriatic, these communities have been classified within the grassland class Festuco-Brometea, while in the central and western Adriatic, they have been classified as heliophilous edge vegetation of Trifolio-Geranietea sanguinei (Biondi et al. 2014, Allegrezza et al. 2015) or recently into a new class Charybdido pancratii-Asphodeletea ramosi Biondi et al. 2016 (Biondi et al. 2016, 2017). The aim of this study was to (i) contribute to knowledge of the composition of dry grasslands dominated by Asphodelus ramosus in Montenegro and (ii) compare examples of Asphodelus ramosus dominated vegetation along the Adriatic and its syntaxonomical classification and classifi- cation into Natura 2000 habitat types. Materials and methods Study area Ćemovsko polje is a part of Zetska ravnica plain in Montenegro, between the rivers Morača, Cijevna and Ribnica. It covers 165 km2 and extends from Podgorica, the capital of Montenegro, to Skadar Lake. The altitude of the investigated area ranges from 12 to 30 m a.s.l. During the Pleistocene period, moraine material eroded from the mountains was carried along the rivers Morača and Cijevna into the area of Ćemovsko polje (Radojičić 2015). The dom- inant types of soil are eutric cambisol and rendzina, which are formed on fluvio-glacial deposits, and consequently are often shallow and skeletoid (Burić et al. 2017). The area of Ćemovsko polje has a Mediterranean climate with hot sum- mers – Csa (Burić and Micev 2008). A grassland ecosystem is dominant in this area. The studied Asphodelus ramosus – dominated vegetation occurs in pastures used for grazing by cattle and sheep. A significant area of Ćemovsko polje is occupied by vineyards and plantations of peach and other kinds of fruit (Radojičić 2015). The area investigated has also recently been heavily impacted by urbanization (Burić et al. 2017). Although many floristic studies have been performed in the area of Ćemovsko polje (Černjavski et al. 1949, Hadžiablahović 2010, 2018, Stešević et al. 2014), the vegeta- tion has remained poorly studied (Černjavski et al. 1949, Hadžiablahović 2018). Potential natural vegetation of the wider area of Podgorica and Lake Skadar is Apulian-south- east Adriatic meso-supra-Mediterranean Quercus trojana forests with Pistacia species (Bohn et al. 2000-2003) or precisely Macedonian oak forest ‘Quercetum trojanae montenegrinum Blečić et Lakušić 1975’ (recte: Quercetum trojanae Em 1958). Illyrian sub-Mediterranean rocky grasslands on shallow calcareous soils of the alliance Chrysopogono grylli-Koelerion splendentis Horvatić 1973 prevail in the area of Ćemovsko polje (Hadžiablahović 2010). Sampling and data analysis From April to June 2019, we sampled Asphodelus ramosus-dominated plant communities on Ćemovsko polje according to the Braun-Blanquet method (Braun-Blanquet 1964). We made 17 relevés and the size of plots was 25 m2. The minimum distance between plots was 100 m. The min- imum coverage value of Asphodelus ramosus for it to be con- sidered dominant was 25%. Each plot was visited in April and again in June. Cover values of species and total vegeta- tion cover are based on the summer aspect (Tab. 1, Appen- dix 1). GPS coordinates were recorded for each plot (Appen- dix 2). All relevés made during fieldwork and relevés from the literature were entered into the Turboveg (Hennekens and Schaminée 2001) database. Relevés from the literature were used for comparison with vegetation data from our fieldwork (Appendix 3). We obtained characteristic species of the association Bromo erecti-Chrysopogonetum grylli according to Horvatić (1963). Diagnostic taxa of alliance and order were assigned according to Terzi (2015), while the diagnostic taxa of class- es were determined according to Terzi (2015) for the class Festuco-Brometea, Mucina et al. (2014) for other classes and Biondi et al. (2016) for the class Charybdido pancratii- Asphodeletea ramosi. Non-metric multidimensional scaling (NMDS, Kruskal 1964) was used to examine the overall variation in the spe- cies composition in the whole relevé dataset. Hellinger transformation of percentage cover values (5=87.5 %, 4=62.5 %, 3= 37.5 %, 2=12.5 %, 1=2.5 %, +=0.5 %, r=0.1 %) was used and Bray-Curtis as a measure of dissimilarity. NMDS was performed using the R package ‘vegan’ (Oksanen et al. 2017). For ecological interpretation of vegetation patterns, ecological indicator values (Pignatti et al. 2005) were pas- sively projected onto the NMDS graph. Weighted by species cover, mean indicator values were calculated for each relevé using JUICE software (Tichý 2002). STANIŠIĆ-VUJAČIĆ M., STEŠEVIĆ D., HADŽIABLAHOVIĆ S., CAKOVIĆ D., ŠILC U. 14 ACTA BOT. CROAT. 81 (1), 2022 The nomenclature of taxa is in accordance with Euro+Med (2006) and the nomenclature of higher syntaxa according to Mucina et al. (2014). The taxonomy of Asphodelus ramosus species was some- times ambiguous. Previously, the species Asphodelus microcarpus Viv. was accepted as valid by many authors but, after a taxonomic revision of the genus Asphodelus in the western Mediterranean (Diaz-Lifante and Valdés 1996), it has been considered a synonym of Asphodelus ramosus L. (Euro+Med 2006). Results The Asphodelus ramosus plant community from Ćemovsko polje (Montenegro) is represented by 17 relevés in the phytosociological table (Tab. 1). The dominant species of the association are Asphodelus ramosus and Chrysopogon gryllus, while the most frequent species are Asphodelus ramosus, Sanguisorba minor, Poa bulbosa, Teucrium capitatum, Sideritis romana subsp. purpurea, and Anacamptis papilionacea. Asphodelus ramosus is also considered to be the only characteristic species of the asso- ciation. The community develops in two clear phenological aspects. In the spring aspect (Fig. 2A), the dominant species is Asphodelus ramosus, while in summer (Fig. 2B), domi- nance is taken by the tall grass Chrysopogon gryllus. The spring aspect is also characterised by high frequency and coverage of Anemone hortensis, Poa bulbosa and Sanguisorba minor, and the summer one by Bupleurum veronense and Teucrium capitatum. In addition to the species characteristic for the class Festuco - -Brometea, there are several species of annual and epheme- ral grasslands of the classes Stipo-Trachynietea distachyae and Helianthemetea guttate, and seasonal perennial and ephemeroid pastures of the class Poetea bulbosae, all indi- cating the Mediterranean and grassland character of the studied plant community. The impact of grazing is evident from the many ruderal species: Avena barbata, Scandix pecten-veneris, Daucus guttatus, Scolymus hispanicus, Bromus squarrosus, Euphorbia exigua, Sonchus oleraceus (Tab. 1). The studied stands constitute an open grassland community, which develops on stony soil and is used for grazing (cattle and sheep). Total vegetation coverage is 60 – 80% in the summer aspect. Stones and pebbles have dimen- sions of up to 20 cm and their cover is 5 – 40%. If the stones are removed and these areas are used for mowing, the vegetation changes into dry grasslands of the alliance Vulpio- -Lotion (Hadžiablahović 2018). Using various numerical classifications, we tried to clas- sify the association Bromo erecti-Chrysopogonetum grylli into subassociations. This kind of vegetation is fairly uni- form in the study area, and there were no ecologically evi- dent/logical subunits. A comparison of the studied Asphodelus community from Montenegro with Asphodelus ramosus-dominated communities from around the Adriatic (Fig. 1) revealed two distinct vegetation groups (Tab. 2, Fig. 3). Communities from Albania, Croatia and Montenegro form one group, Fig. 2. Asphodelus ramosus dominated community on Ćemovsko polje, vicinity of Podgorica, Montenegro. A – spring aspect, B – sum- mer aspect. Fig. 1. Localities of data used of Asphodelus ramosus-dominated communities. See Tab. 2 for community abbreviations. AL – Al- bania, BA – Bosnia and Herzegovina, HR – Croatia, IT – Italy, ME – Montenegro, RS – Serbia, SI – Slovenia. ASPHODELUS RAMOSUS DOMINATED PLANT COMMUNITY ACTA BOT. CROAT. 81 (1), 2022 15 Table 1. Phytosociological table of the association Bromo erecti-Chrysopogonetum grylli in Ćemovsko polje (Montenegro). Relevé number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 % Plot size (m2) 25 Vegetation cover (%) 80 60 80 70 75 60 60 75 60 65 70 70 60 60 60 60 60 Bromo erecti-Chrysopogonetum grylli Asphodelus ramosus 4 3 4 3 4 3 3 3 3 4 3 3 3 3 4 3 4 17 Chrysopogono grylli-Koelerion splendentis Bupleurum veronense + + + . . + . + . . + . + + + . . 9 Carduus nutans subsp. micropterus . + . + . . . + . . . . . + . . . 4 Scorzoneretalia villosae Chrysopogon gryllus 1 2 2 3 2 2 1 2 . 1 3 2 2 2 2 1 . 15 Eryngium amethystinum + + 1 . 1 + 1 + + . 1 1 + 1 1 1 1 15 Koeleria lobata + + + . + 1 . . + . 1 . + 1 1 + 1 12 Seseli montanum subsp. tommasinii 1 . . . + . + . + . . 1 . . + 1 . 7 Satureja subspicata . . . + . . . . + 2 . . . 2 2 . + 6 Festuca stricta subsp. sulcata . . . + . + . 1 2 . . + . . . . . 5 Medicago prostrata + . . . . . . . . . 1 + . + . . . 4 Plantago holosteum . + + + . . . . . . . + . . . . . 4 Bunium alpinum subsp. montanum . . + . . . . . + + . . . . + . . 4 Scorzonera villosa . . 1 + . . . . + + . . . . . . . 4 Festuco-Brometea Sanguisorba minor 3 2 3 2 2 + 1 2 + 1 2 2 2 1 1 2 1 17 Poa bulbosa 1 1 1 2 1 2 2 1 + + 1 + 2 1 1 2 1 17 Teucrium capitatum + 1 1 + + 1 1 1 2 + 1 2 + 2 2 1 1 17 Hypericum perforatum + + + + + + . 1 + . + + + + + 1 + 15 Hippocrepis ciliata + . 1 . + 1 + . . 1 + + + + + + + 13 Centaurea deusta + + . . + + + 3 1 + 1 1 1 1 . 3 . 13 Leontodon crispus . . + + + . . . 1 1 1 1 1 . 1 + + 11 Ranunculus millefoliatus . + + + . . . + + . . + 1 + + . + 10 Convolvulus cantabrica + . . . 2 1 2 . . . 1 1 1 + . 1 . 9 Petrorhagia saxifraga . . . + + + + + . . + + . . + + . 9 Linum tenuifolium . + + . . + . . 1 . . + + + . . + 8 Anthyllis vulneraria subsp. polyphylla . . + . + + . . + + + + + . . . . 8 Carex caryophyllea . . . 1 . . . . . . 1 + + 1 1 1 . 7 Podospermum laciniatum + + . + + . . . . . . . . . . . + 5 Bothriochloa ischaemum . . . . . . . + 1 + 1 . . . 1 . . 5 Anacamptis morio . + . + + . . . . . + . . . . . . 4 Trifolium campestre . + . . . . + . . + . . . . . + . 4 Ophrys sphegodes . . . . . . + . + . . . . . + . . 3 Cuscuta epithymum subsp. epithymum . . . . + . . . . . + + . . . . . 3 Thymus striatus . . . . . . . . + . . . . + + . . 3 Leopoldia comosa . + . . . . . . + . . . . . . . + 3 Chenopodietea Avena barbata + + + + 1 1 1 . . 2 . . 1 . . + + 11 Valantia muralis . . 1 . + . . . . 1 . . . . + . + 5 Scandix pecten-veneris . . . . . . . + + . . + . . . . . 3 Stipo-Trachynietea distachyae Sideritis romana subsp. purpurea + + 1 + + 1 + + + + + + + + + + 1 17 Crepis sancta 1 1 + + 1 1 1 1 + 1 + + . . . 1 + 14 Arenaria leptoclados 1 . + 1 + + + . . . . . + + + + . 10 Polygala monspeliaca . + + . + + + . . + + . . + . 1 + 10 Ornithogalum collinum + . . . + + + 1 + . 1 1 2 . . . . 9 Tordylium apulum . . . + . + + + . . + . . . . . + 6 STANIŠIĆ-VUJAČIĆ M., STEŠEVIĆ D., HADŽIABLAHOVIĆ S., CAKOVIĆ D., ŠILC U. 16 ACTA BOT. CROAT. 81 (1), 2022 Relevé number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 % Plot size (m2) 25 Vegetation cover (%) 80 60 80 70 75 60 60 75 60 65 70 70 60 60 60 60 60 Petrorhagia dubia  . . + + . . . + . . . . + . . . . 4 Filago germanica + . . . . + . . . . . + . . . . + 4 Ononis reclinata + . . . + + . . . . . . . . . + . 4 Trachynia distachya . . + . + . . . . . . 1 . . . . . 3 Neatostema apulum . . . . . . . . . . . . 1 + + . . 3 Helianthemetea guttati Helianthemum salicifolium 1 + 1 + 1 1 2 2 + 1 1 1 1 . 1 1 . 15 Crupina vulgaris + + + . + + 1 . 1 + + + + + . . . 12 Asterolinon linum-stellatum + 1 + + + 1 1 . . . . . + + 1 + . 11 Galium divaricatum 1 1 + + . + . + . . . . . + . + + 9 Plantago bellardii + . . . . . + + . + . . 1 + + . . 7 Linaria pelisseriana . + . + . . . + + . . . . . . . + 5 Filago gallica . + . . . + . . . . . . + . + . . 4 Aira elegantissima . . . + . . . . . . . . . + . + + 4 Trifolium stellatum + . + . + . . . . . . . . . . . . 3 Poetea bulbosae Anthoxanthum odoratum + 2 . 1 + + + . + + . . + + . 1 + 12 Trifolium subterraneum + 1 . 1 + . + + . . 1 . + . . 1 1 10 Plantago lanceolata . + + 1 . . . + + + . . . . . + + 8 Leontodon tuberosus . . + + + . + 1 + . . . . . . . . 6 Herniaria glabra . . . . . . . . . . . . + + + + . 4 Prospero autumnale . . + . . . . . . . . . + . . + . 3 Trifolium nigrescens . + . . . . . . . 1 . . . . . . + 3 Sedo-Scleranthetea Cerastium pumilum subsp. glutinosum + + . 1 1 1 . + . . . . 1 + + 1 + 11 Viola kitaibeliana 1 1 + + + + . + . + . . + + . . . 10 Clinopodium acinos . . + . + . 1 . . . + + + + + . . 8 Draba verna + . . + . . . . . . . . + + + + . 6 Artemisietea vulgaris Daucus guttatus + + + + + 1 1 . . 2 + . + . . + + 12 Scolymus hispanicus . + + 1 . + 1 1 + + + . . . + 1 . 11 Tyrimnus leucographus 1 . . + . + + 1 . + . . . . . + + 8 Potentilla recta . . . + 1 . + 1 . . . . . + . . . 5 Sisymbrietea Geranium columbinum + + . + + + + + + + + . + + . + + 14 Erodium cicutarium + . . 1 . . . 1 . . + + 1 + + . 1 9 Bromus squarrosus 1 . . . . . 1 + . . 1 . + . + . . 6 Papaveretea rhoedis Euphorbia exigua + . . . + 1 1 . + + + + 1 1 + + + 13 Sherardia arvensis + + . . + + 1 . + . . . + . . + + 9 Anagallis arvensis . . + + + + 1 . + 1 . . . . . + . 8 Sonchus oleraceus + + . . . + + . . + + . + . . . . 7 Euphorbia helioscopia . . . . . . . . + . . + + . . + . 4 Euphorbia taurinensis . . . . . . . . . . . . + + + . . 3 Molinio-Arrhenatheretea Serapias vomeracea . . + . . . . . . . + + . + . + . 5 Hypochaeris radicata . + . + . . . . . . . . . . + + 1 5 Ononido-Rosmarinetea Carlina corymbosa . . + . 1 + 1 . . + + 1 1 + . . 1 10 ASPHODELUS RAMOSUS DOMINATED PLANT COMMUNITY ACTA BOT. CROAT. 81 (1), 2022 17 Relevé number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 % Plot size (m2) 25 Vegetation cover (%) 80 60 80 70 75 60 60 75 60 65 70 70 60 60 60 60 60 Lygeo sparti-Stipetea tenacissimae Anacamptis papilionacea + + + + + . + + + 1 1 1 1 1 + 1 + 16 Anemone hortensis 2 2 2 2 2 1 2 1 + 1 1 . . . . 2 . 12 Other species Cynodon dactylon . + + + + + + . + 1 + + + + + + + 15 Allium guttatum subsp. sardoum + 1 . 1 + . + + . . + . + . + + + 11 Cerastium ligusticum subsp. trichogynum 2 1 + 1 1 . . . + . . 1 . 1 1 . . 9 Silene italica . + + + + + . . . . + 1 . 1 . . . 8 Alkanna tinctoria . . . . . + . . . . 1 1 1 1 1 . . 6 Hainardia cylindrica . . . . . . 1 . . . . . + + . . . 3 Trifolium scabrum . . . . . . . . . + . + + . . . . 3 Table 2. Shortened synoptic table of plant communities with Asphodelus ramosus. The original classification indicated by authors is used. Communities: 1. Bromo erecti-Chrysopogonetum grylli (Montenegro, new relevés from Ćemovsko polje), 2. Asphodelo-Chrysopogonetum grylli (Albania, Fanelli et al. 2015), 3. Bromo-Chrysopogonetum grylli subass. asphodeletosum microcarpi (Croatia, Horvatić 1934), 4. Narcisso tazzettae-Asphodeletum microcarpi (Croatia, Šegulja 1970), 5. Narcisso tazzettae-Asphodeletum microcarpi (Croatia, Hećimović 1984), 6. Narcisso tazzettae-Asphodeletum microcarpi (Croatia, Jasprica et al. 2016), 7. Asphodelus ramosus community (Albania, Fanelli et al. 2015), 8. Charybdido pancratii-Asphodeletum ramosi (Italy, Biondi et al. 2016), 9. Alkanno tinctoriae-Asphodeletum ramosi (Italy, Biondi et al. 2016), 10. Euphorbio characiae-Thapsietum garganicae (Italy, Biondi et al. 2017), 11. Asphodelo ramosi-Feruletum communis (Italy, Biondi et al. 2016), 12. Asphodelino luteae-Feruletum communis (Italy, Biondi et al. 2016). Plant community 1 2 3 4 5 6 7 8 9 10 11 12 Number of relevés 17 4 12 8 10 3 3 14 3 5 2 7 Bromo erecti-Chrysopogonetum grylli Asphodelus ramosus 100 25 100 100 100 100 100 100 100 60 100 100 Cytisus spinescens . . 50 . . . . . . . . . Narcisso tazzettae-Asphodeletum microcarpi Narcissus tazetta . 25 . 100 70 100 . 7 . . . . Anacamptis papilionacea 94 . . 75 . . . 21 . 40 . 29 Chrysopogono grylli-Koelerion splendentis Carduus nutans subsp. micropterus 24 25 50 50 50 . . . . 80 . 43 Bupleurum veronense 53 . 42 . 90 . 67 . . . . . Centaurea cristata . . . 13 . . . . . . . . Salvia officinalis . 50 8 . . . . . . . . . Scorzoneretalia villosae Chrysopogon gryllus 88 100 100 88 . 67 33 . . . . . Koeleria lobata 71 25 67 88 40 67 . . . . . . Plantago holosteum 24 25 33 88 . . . . . . . . Eryngium amethystinum 88 50 58 88 . . 33 7 . 60 . 57 Salvia pratensis . 25 33 88 . . . . . . . . Medicago prostrata 24 . 25 . . . . . . . . . Festuca valesiaca . . 83 13 . . . . . . . . Seseli montanum subsp. tommasinii 41 . . . . . . . . . . . Satureja subspicata 35 . . . . . . . . . . . Potentilla heptaphylla subsp. australis . . 8 . . . . . . . . . Dorycnium pentaphyllum subsp. germanicum . . 17 . . . . . . . . . Festuco-Brometea Sanguisorba minor 100 25 17 88 . . . 29 . 20 . . Bothriochloa ischaemum 29 25 . 50 100 . 33 . . . . . Bromopsis erecta . 25 100 75 . 33 . . . . . . Hippocrepis comosa . . 17 25 . . . . . . . . STANIŠIĆ-VUJAČIĆ M., STEŠEVIĆ D., HADŽIABLAHOVIĆ S., CAKOVIĆ D., ŠILC U. 18 ACTA BOT. CROAT. 81 (1), 2022 Plant community 1 2 3 4 5 6 7 8 9 10 11 12 Number of relevés 17 4 12 8 10 3 3 14 3 5 2 7 Linum tenuifolium 47 25 8 63 . . . . . . . . Ruta graveolens . 25 17 75 . . . 36 . . . . Leopoldia comosa 18 . . . 70 . . . . . . 43 Carex caryophyllea 41 25 . 50 . . . . . . . . Satureja montana . 25 8 . . . 33 . . . . . Ranunculus millefoliatus 59 . . . . . . . . 20 . 14 Charybdido pancratii-Asphodeletum ramosi Carlina corymbosa 76 50 100 63 . . 33 50 . 80 100 29 Asparagus acutifolius 6 25 50 100 50 . . 79 100 60 100 86 Drimia pancration . . . . . . . 100 67 20 100 57 Anemone hortensis 71 25 . 63 70 . . 64 . 80 50 57 Asphodeline lutea . . . . . . . 7 . 60 . 100 Thapsia garganica . . . . . . . 57 . 100 . 100 Ferula communis . . . . . . . . . 20 100 100 Hypochaeris radicata 29 . . . . . . 29 33 . . . Asphodeline liburnica . . . . . . . 7 . . . . Other Dactylis glomerata subsp. hispanica . 25 100 88 50 100 . 93 67 . . . Plantago lanceolata 47 25 33 100 90 . . 29 . . . 71 Reichardia picroides . 25 50 75 100 67 . 43 . 40 . 43 Catapodium rigidum 6 50 75 88 100 . 100 . . . . . Crepis sancta 82 . . . . . . 43 67 . 100 43 Avena barbata 65 . 33 50 . . 67 . 33 . 100 29 Trifolium scabrum 18 25 25 25 50 . 67 . . . . . Helichrysum italicum . 25 75 50 90 67 33 7 . . . . Paliurus spina-christi 25 63 . . . . . 43 . . 100 . Olea europaea 17 . . . . . . 43 . . 100 . Pinus halepensis . . . . . . . . 33 20 . . Fig. 3. NMDS ordination of Asphodelus ramosus-dominated communities from the Adriatic region with passively projected ecological indicator values. Symbols indicate classification into the classes: – Festuco-Brometea, – Artemisietea vularis, – Charybdido pancratii-Asphodeletea ramosi. Numbers indicate centroids of relevés of particular communities (see Tab. 2). ASPHODELUS RAMOSUS DOMINATED PLANT COMMUNITY ACTA BOT. CROAT. 81 (1), 2022 19 with abundant grassland species of the class Festuco- Brometea, which are not present in the second group of edge communities from Italy. Two dimensional solution of the NMDS ordination attained a minimum stress of 0.21. The NMDS ordination clearly distinguishes the two groups along gradients of nutrients, light and soil reaction. Edge communities from Italy thrive on more nutrient-rich, shaded sites (Fig. 3). Based on floristic composition and all the comparative analyses, we decided to classify the studied stands domi- nated by Asphodelus ramosus into the already described grassland association Bromo erecti-Chrysopogonetum grylli. Discussion In recent years, several studies on the distribution and dynamics of plant communities with Asphodelus ramosus have been performed, especially in the central and western Mediterranean (Biondi et al. 2016, Biondi et al. 2017). In contrast, this type of vegetation has been poorly studied in Montenegro. The association Bromo erecti-Chrysopogonetum grylli was described for the first time in Croatia (Island of Pag, Horvatić 1934), with two subassociations (Bromo- Chrysopogonetum grylli dorycnietosum herbacei and Bromo-Chrysopogonetum grylli asphodeletosum microcarpi), and classified in the alliance ‘Chrysopogoneto-Satureion subspicatae Horvat i Horvatić 1934’ (recte: Chrysopogono grylli-Koelerion splendentis), order ‘Brometalia erecti Br. Bl.’ Subsequently, Bromo-Chrysopogonetum grylli asphodeletosum microcarpi was raised by Horvatić (1963) to the rank of association, with the name Asphodelo microcarpi-Chrysopogonetum grylli, and included in the order ‘Scorzonero-Chrysopogonetalia H-ić et Ht (1956) 1958’ (recte: Scorzoneretalia villosae). The association has so far been reported in Montenegro and Albania (Černjavski et al. 1949, Fanelli et al. 2015, Hadži- ablahović 2018). In his nomenclatural revision of the order ‘Scorzonero villosae-Chrysopogonetalia grylli Horvatić et Horvat in Horvatić 1963’, Terzi (2011) considered the asso- ciations Asphodelo-Chrysopogonetum grylli and Bromo- -Chrysopogonetum grylli to be valid, while Terzi (2015) later united them and retained the earlier valid name, Bromo- -Chrysopogonetum grylli (cf. Theurillat et al. 2021). In Montenegro, Bromo erecti-Chrysopogonetum grylli was reported by Blečić and Lakušić (1976) in the coastal part of Montenegro, as well as in the vicinity of Podgorica, and by Černjavski et al. (1949) and Hadžiablahović (2018) in the area of Skadar Lake. These publications were not supported with phytosociological studies and it was not possible to make a comparison with our results from Ćemovsko polje. Accord- ing to Černjavski et al. (1949), Bromo-Chrysopogonetum grylli is developed on stony hills with thin soils, while our researched stands are found in the lowlands of Ćemovsko polje on deeper alluvial soils. According to Černjavski et al. (1949) Bromo-Chrysopogonetum grylli is characterized by many species of shrub vegetation (Paliurus spina-christi, Salvia officinalis, Rubus ulmifolius, Euphorbia veneta, Helichrysum italicum, Ruscus aculeatus, Cyclamen neapolitanum, Arum italicum, Phlomis fruticosa, Cistus villosus, Cistus salviaefolius, Nephrodium filix-mas, Pteridium aqiulinum, etc.) and chas- mophytic vegetation (Asplenium trichomanes, Edraianthus tenuifolius, Cardamine glauca, Silene quadridentata, Ceterach officinarum, Sedum album, Moltkea petraea, etc.) that were absent from our stands from Ćemovsko polje. Further in- vestigation of the association Bromo-Chrysopogonetum grylli reported by Černjavski (1949) is needed to determine whether there are differences in relation to our community from Ćemovsko polje. Fanelli et al. (2015) reported Asphodelo-Chrysopogonetum grylli in Albania in the Buna River protected landscape and it shows high similarity with associations from Montenegro and Croatia. The species Cytisus spinescens, which is a charac teristic species of the association (Horvatić 1963) is not present in stands from Montenegro and Albania. Stands from Montenegro are also characterized by an absence of species characteristic of dry grasslands of the classes Festuco-Brometea – Salvia officinalis and Bromopsis erecta. According to Horvatić (1934, 1939), the development and distribution of Bromo-Chrysopogonetum grylli asphodeletosum microcarpi is conditioned by agro-pastoral activities, i.e., moderate grazing. In the case of Ćemovsko polje, it is grazed particularly by sheep and rarely by cattle. Intensive and permanent grazing leads to degradation of this community, which is especially evident from the absence of Asphodelus ramosus from relevés on Rab island (Horvatić 1939). Inten- sive grazing leads to a higher abundance of shrubs ( Helichrysum italicum, Cytisus spinescens, Euphorbia spinosa, Salvia officinalis) (Horvatić 1934), or ruderal and subruderal species (Fanelli et al. 2015). Overgrazed stands can be included in the order Carthametalia lanatae ( Artemisietea vulgaris) because of abundant therophytes: Carthamus lanatus, Dasypyrum villosum, Catapodium rigidum, Nigella arvensis, etc. (Fanelli et al. 2015). On the other hand, stands are abandoned, after which succession leads to Rhamno-Paliuretum Trinajstić 1996, or are turned into mowed grasslands of the Vulpio-Lotion Horvatić 1963 alliance (Hadžiablahović 2018). In several localities in Croatia along the Adriatic coast, similar vegetation types with Asphodelus ramosus have been reported. Narcisso tazettae-Asphodeletum microcarpi was described in Istria (Šegulja 1970). The same association was later reported on the islands of Bobara and Mrkan (Hećimović 1984), Supetar (Jasprica and Ruščić 2013) and Olib (Jasprica et al. 2016). Originally, it was classified into Scorzonerion villosae, but was later moved to Chrysopogono grylli-Koelerion splendentis. Ecological conditions are dif- ferent to those of Asphodelo ramosi-Chrysopogonetum grylli and it occurs on deep skeletoid soils and under the influence of salt spray (Šegulja 1969). According to Jasprica et al. (2016), the association is considered to be the most ther- mophilous grassland along the eastern Adriatic coast. The characteristic species of the association are Asphodelus ramosus, Narcissus tezzeta and Orchis papilionacea. Thermophilous grassland communities with domination of STANIŠIĆ-VUJAČIĆ M., STEŠEVIĆ D., HADŽIABLAHOVIĆ S., CAKOVIĆ D., ŠILC U. 20 ACTA BOT. CROAT. 81 (1), 2022 Asphodelus ramosus have a distribution along the eastern Adriatic, influenced by the Mediterranean climate. There is a different situation along the western Adriatic coast, where Asphodelus ramosus-dominated communities are considered to be heliophilous fringe and tall-herb veg- etation, which develops after the abandonment of agro-pas- toral activities (Tesei et al. 2020). Comprehensive studies on heliophilous edge vegetation have been performed in Italy (Gargano peninsula, central part of the Apennines, north- ern part of Sardinia) and southern Spain (Biondi et al. 2016, 2017). As a result, a new class of edge vegetation Charybdido pancratii-Asphodeletea ramosi was described, focused on areas with a Mediterranean macrobioclimate (Biondi et al. 2016, 2017). The proposal to update the EuroVegChecklist (Mucina et al. 2014) by adding this new class was provisionally re- jected by the European Vegetation Classification Commit- tee of the European Vegetation Survey working group of the International Association of Vegetation Science, due to the lack of evidence concerning a clear floristic delimitation of Charybdido pancratii-Asphodeletea ramosi from Lygeo sparti- Stipetea tenacissimae and Trifolio-Geranietea sanguinei ( Biurrun and Willner 2020). Nonetheless, we can accept the existence of a new fringe class replacing Trifolio-Geranietea in the Mediterranean, but we are of the opinion that Bromo erecti-Chrysopogonetum grylli clearly represents grassland vegetation erroneously classified as fringe vegetation by Biondi et al. (2016). This is clearly indicated by the numer- ous species of Festuco-Brometea present in stands (Tab. 2) from eastern Adriatic Asphodelus-dominated communities. Fringe communities with Asphodelus ramosus from Italy, due to overgrazing, are dominated by monocotyledons, ma- ny of them toxic to animals, while in Montenegro these ar- eas represent pastures. Another difference is the presence of shrub species with higher cover in stands of Charybdido- Asphodeletea ramosi (Paliurus spina-christi, Olea europaea, Pinus halepensis), missing in grasslands from the eastern Adriatic. According to Biondi et al. (2016) the diagnostic species of the newly described class are Asphodelus ramosus subsp. ramosus, A. fistlosus, A. tenuifolius, A. ayardii, Charybdis pancration, C. maritima, C. glaucophylla, C. aphylla, C. hesperia, Thapsia garganica, Asparagus acutifolius, Ornithogalum etruscum subsp. umbratille, Anemone hortensis, Carlina corymbosa, Hypochoeris radicata, Iris planifolia, I. bicapitata, Asphodeline liburnica, A. lutea, Ferula communis, F. communis subsp. cardonae, F. glauca, F. arrigonii and Hermodactylis tuberosus. The diagnostic species group of Charybdido-Asphodeletea should also be revised, since many of them are attributed to other vegetation classes (Festuco- -Brometea, Lygeo sparti-Stipetea tenacissimae, Ononido- - Rosmarinetea etc.) according to Mucina et al. (2014). Asphodelus ramosus, Anemone hortensis, Carlina corymbosa and Hypochoeris radicata, which are considered to be character species by Biondi et al. (2016), are also very frequent in grassland vegetation of Bromo erecti-Chrysopogonetum grylli. In terms of Natura 2000 habitat types (Council Direc- tive 92/43/EEC on the conservation of natural habitats and of wild fauna and flora, 1992) there are differences between grassland and fringe communities dominated by Asphodelus ramosus. Heliophilous Asphodelus spp. edge communities in the western and central Mediterranean do not represent any habitats of European Community interest and a pro- gressive increase in Asphodelus spp. cover in grasslands can result in the disappearance of grassland habitat types ( Tesei et al. 2020). According to a report of habitat types of Montenegro important for the European Union (Petrović et al. 2019), Bromo erecti-Chrysopogonetum grylli is classi- fied within the Eastern sub-Mediterranean dry grasslands (Scorzoneretalia villosae) habitat type (code 62A0). The same situation applies to Narcisso tazettae-Asphodeletum microcarpi in Croatia (Jasprica et al. 2016). Syntaxonomical scheme Based on the analyses performed, the following syntax- onomical scheme is proposed for Asphodelus ramosus-dom- inated communities in the Adriatic region. Festuco-Brometea Br.-Bl. et Tx. ex Soó 1947 Scorzoneretalia villosae Kovačević 1959 Chrysopogono grylli-Koelerion splendentis Horvatić 1973 Bromo erecti-Chrysopogonetum grylli Horvatić 1934 Narcisso tazettae-Asphodeletum microcarpi Šegu lja 1969 Artemisietea vulgaris Lohmeyer et al. ex von Rochow Asphodelus ramosus community Charybdido pancratii-Asphodeletea ramosi Biondi in Biondi et al. 2016 Asphodeletalia ramosi Biondi in Biondi et al. 2016 Charybdido pancratii-Asphodelion ramosi Biondi et al. 2016 Charybdido pancratii-Asphodeletum ramosi Biondi et al. 2016 Alkanno tinctoriae-Asphodeletum ramosi Biondi et al. 2016 Euphorbio characiae-Thapsietum garganicae Biondi et al. 2017 Asphodelo ramosi-Ferulion communis Biondi et al. 2016 Asphodelo ramosi-Feruletum communis Biondi et al. 2016 Asphodelino luteae-Feruletum communis Biondi et al. 2016 Acknowledgments We thank editor Massimo Terzi and two anonymous re- viewers for their work that substantial improved our manu- script. 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Asparagus acutifolius 17: +; Catapodium rigidum 6: +; Geranium molle 2: +; Trifolium angustifolium 3: +; Romulea bulbocodium 2: +; Sonchus asper 10: +; Pyrus amygdaliformis 3: 1; Carthamus lanatus 6: +; Salvia verbenaca 16: +; Teucrium chamaedrys 17: +; Geranium purpureum 7: +; Alyssum minus 10: 1; Ajuga chamaepitys 12: +, 16: +; Vicia angustifolia 9: +; Crepis foetida 2: +, 6: +; Tragopogon porrifolius 9: +, 11: 1; Bellis perennis 4: +; Parentucellia latifolia 14: +, 15: +; Aphanes arvensis 8: +, 17: +; Valerianella rimosa 6: +, 8: +; Knautia integrifolia 9: +, 10: +; Stipa pulcherrima 3: +, 4: 1; Cardamine hirsuta 4: +, 17: +; Linum bienne 5: +, 7: +; Trifolium cherleri 3: +, 5: +; Bromus madritensis 7: +, 16: +; Trifolium resupinatum 5: +; Vulpia myuros 10: +; Trigonella gladiata 1: +; Veronica arvensis 17: +; Reseda phyteuma 9: +; Bromus sterilis 11: +; Calepina irregularis 6: +; Bromus hordeaceus 3: +; Orchis ustulata 11: +; Orlaya grandiflora 9: +; Medicago rigidula 5: +; Poa an- nua 10: +; Hyacinthella dalmatica 5: +; Matthiola incana 15: 1; Lactuca viminea 3: +; Aegilops neglecta 11: +; Alyssum alyssoides 7: +; Alyssum campestre 11: 1; Astragalus illyricus 11: 2, 14: +; Matricaria chamomilla 13: +, 14: +; Onosma echioides 9: 1, 12: +. Appendix 2: Relevé dates (year/month/day) and coordinates (WGS84 reference system, in decimal degrees). 1. 2019/04/07, 42.3968090, 19.3021110; 2. 2019/04/07, 42.3964610, 19.3030050; 3. 2019/04/07, 42.3960000, 19.3040000; 4. 2019/04/07, 42.3978055, 19.3048888; 5. 2019/04/07, 42.3985277, 19.3019638; 6. 2019/04/07, 42.4001488, 19.3031944; 7. 2019/04/07, 42.4006410, 19.3023280; 8. 2019/04/07, 42.4021890, 19.3049660; 9. 2019/04/07, 42.3880040, 19.2856690; 10. 2019/04/07, 42.3803160, 19.2743840; 11. 2019/04/09, 42.3708888, 19.2303056; 12. 2019/04/09, 42.3728888, 19.2316944; 13. 2019/04/09, 42.3697222, 19.2339166; 14. 2019/04/09, 42.3705580, 19.2360490; 15. 2019/04/09, 42.3709350, 19.2383990; 16. 2019/04/10, 42.4021370, 19.3210555; 17. 2019/04/10, 42.3885277, 19.3110277. Appendix 3: Relevés from literature sources (name of the plant community, authors, table and number of relevés from original source paper). 1. Bromo-Chrysopogonetum grylli subass. asphodeletosum microcarpi (Horvatić 1934), Tab. 23, rels. 11-16, 18-23; 2. Narcisso tazettae-Asphodeletum microcarpi (Šegulja 1970), Tab. 2, rels. 1–8; 3. Narcisso tazettae-Asphodeletum microcarpi subass. sisymbrietosum officinalis (Hećimović 1984), Tab. 5, rels. 1–10; 4. Narcisso tazettae-Asphodeletum microcarpi (Jasprica and Ruščić 2013), rel. on the page 128; 5. Asphodelus ramosus community (Fanelli et al. 2015), Tab. 41, rels. 15, 70, 492; 6. Asphodelo-Chrysopogonetum grylli (Fanelli et al. 2015), Tab. 25, rels. 31–33; 7. Narcisso tazettae- Asphodeletum microcarpi (Jasprica et al. 2016), Tab. 11, rels. 1–3; 8. Charybdido pancratii-Asphodeletum ramosi ( Biondi et al. 2016, Tab. 1, rels. 1–14; 9. Alkanno tinctoriae- Asphodeletum ramosi (Biondi et al. 2016), Tab. 2, rels. 1–3; 10. Asphodelo ramosi-Feruletum communis (Biondi et al. 2016), Tab. 3, rels. 1–2; 11. Asphodelino luteae-Feruletum communis (Biondi et al. 2016), Tab. 4, rels. 1–7; 12. Euphorbio characiae-Thapsietum garganicae (Biondi et al. 2017), Tab. 6, rels. 1–5.