170 ACTA BOT. CROAT. 79 (2), 2020 Acta Bot. Croat. 79 (2), 170–184, 2020 CODEN: ABCRA 25 DOI: 10.37427/botcro-2020-017 ISSN 0365-0588 eISSN 1847-8476 Phytosociological study of submontane genistoid scrub communities from the Southeastern Balkans Georgi Kunev1*, Rossen Tzonev1, Ioannis Tsiripidis2, Kalina Pachedjieva1 1 Department of Ecology and Environmental Protection, Faculty of Biology, Sofia University “St. Kliment Ohridski”, 8 Dragan Tzankov Blvd., BG–1164 Sofia, Bulgaria 2 Department of Botany, Aristotle University of Thessaloniki, GR–54124, Thessaloniki, Greece Abstract – Genista lydia Boiss. is an endemic Balkan-Anatolian species which forms rare communities in the territory of south Bulgaria and northeast Greece. They are spread exclusively on acidic, siliceous substrates. This study presents new data on their distribution, floristic and ecological structure and phytosociological affin- ities. The research is based on 156 phytosociological relevés. Unweighted pair–group method with arithmetic averages (UPGMA) was employed and a detrended correspondence analysis (DCA) was performed prior to the syntaxonomical decision. Three new associations and the new alliance Genistion lydiae have been described and classified within the order Lavandulo stoechadis-Hypericetalia olympici Mucina in Mucina et al. 2016 of the class Cisto-Lavanduletea stoechadis Br.-Bl. in Br.-Bl. et al. 1940. The study has also emphazised the potential threats concerning a decline of the habitat area and proposed some conservation measures. Keywords: Cisto-Lavanduletea stoechadis, Genistion lydiae, grazing impact, shrub vegetation, vulnerable habitat types * Corresponding author e-mail: gorokunev@abv.bg Introduction The communities of Genista lydia Boiss. and G. rumelica Velen. represent indigenous and rare vegetation described from the Southern Balkans and mostly south Bulgaria (Bondev 1991). The dominant species are low shrubs, with erect or ascending branches, which lose their leaves dur- ing the summer drought (Kuzmanov 1976, Strid 1986). Ac- cording to the Bulgarian floristic literature G. lydia is clearly distinguished from G. rumelica at species level (Kuzmanov 1976, Evstatieva et al. 2004, Delipavlov and Chesmedzhiev 2011). However, for the purpose of the study, a broader de- scription of G. lydia (incl. G. rumelica) has been adopted, one that seems more widely accepted at present (Strid 1986, Zieliński et al. 2004, Roskov et al. 2006, Dimopoulos et al. 2013, The Plant List 2013). Those communities are floristically rich and occur in the transitional (sub-Mediterranean) zone, influenced by Medi- terranean and Continental (Temperate) climate. Because of the rarity and vulnerability of this vegetation type, it was as- sessed for the national (Tzonev and Gussev 2015) and Euro- pean (Janssen et al. 2016) Red Data Books of Habitat Types. The communities dominated by G. lydia have been pre- sented (Velčev and Bondev 1984a) as an original and typi- cal part of the Balkan vegetation. These communities are strongly influenced by anthropogenic activities and have a very limited range and for these reasons, they have been de- termined as "Endangered" (Velčev and Bondev 1984b). More data on their distribution in Bulgaria have been published in the vegetation map of Bulgaria (1:600 000) (Bondev 1991). Bondev (1991) has also concluded that those communities are of secondary origin, because they often replace destroyed xerothermic oak or black pine forests. The most detailed studies on the G. lydia communities, have been focused on their habitat types, their floristic and ecological structure and their range (Tzonev and Gussev 2015, Janssen et al. 2016, Kunev and Tzonev 2019). In those studies, this habitat type is described as mosaic vegetation composed of shrub, herbaceous and chasmophytic plants, typical of the Balkan regions, influenced by the transitional continental-Mediterranean climate. Despite comparatively detailed information at the habi- tat level, the syntaxonomic position of this vegetation type is still indefinite. In general, different plant communities dom- inated by or with a high abundance of Genista species are widespread in the Mediterranean regions of Europe. Most of GENISTA LYDIA COMMUNITIES IN THE BALKANS ACTA BOT. CROAT. 79 (2), 2020 171 these communities occupying siliceous substrates were clas- sified within the Calluno-Ulicetea, Cytisetea scopario-striati and Cisto-Lavanduletea classes (Rivas-Martínez et al. 2002; Gianguzzi et al. 2015). For example, some associations have been recorded from the Balkans, such as the association Genisto-Ericetum manipuliflorae Horvatić 1958 from the Croatian coastal region (Horvatić 1958) or the shrub com- munities of the association Genisto acanthocladae-Ouerce- tum cocciferae Dimopoulos et al. 1996 from Peloponnisos, Southern Greece (Dimopoulos et al. 1996). The species Ge- nista januensis Viv. demonstrates great morphological simi- larity and is also closely related to G. lydia. Genista januensis typically inhabits forest fringes and open woodlands domi- nated by Quercus pubescens and Carpinus orientalis in some regions of west Bulgaria (Tzonev et al. 2019). It is also diag- nostic species to the basophilic Pinus sylvestris and P. nigra forests of the Southeastern and Dinaric Alps (Genisto jan- uensis-Pinetum Tomažič 1940) (Chiapella and Longo 1987, Dakskobler 1999). However, after the first assessment for the Red List of European habitat types (Janssen et al. 2016), Bulgaria and Greece were the only countries that provided information on the scrub vegetation dominated by G. lydia. Therefore, its occurrence in other Balkan countries or Western Anato- lia continues to be unclear. The aim of this study is to propose a new syntaxonom- ic framework for G. lydia communities, based on abundant phytosociological material collected from south Bulgaria and northeastern Greece. Material and methods The principles and methods of the study follow the Braun-Blanquet phytosociological approach (Braun-Blan- quet 1964). The plant communities of Genista lydia were sampled between March and October of the years 2016 to 2018. All localities of the dominant species in Bulgaria known from the literature data and herbaria collections have been visited. The localities of the studied communities in Greece were provided by one of the authors. In these areas, relevés were set in places where the coverage of the species exceeded 30%. In total, 156 unpublished phytosociological relevés were used in the present study. Their plot sizes vary between 16 and 25 m2. At each sampling plot, a complete list of vascular plants, mosses and lichens taxa was made togeth- er with their cover–abundance values according to the 9–de- gree modified Br-Bl scale. The Braun-Blanquet categories were transformed to ordinal numbers from 1 to 9 for further statistical analysis (van der Maarel 1979). The data set was subjected to hierarchical classification by the SYN–TAX pro- gram (Podani 2001). The unweighted pair–group method with arithmetic averages (UPGMA) as an average–linkage type of clustering algorithm was used as clustering method. The floristic similarity between relevés in the dissimilari- ty matrix was calculated by using Horn’s index (see Podani 2001). Five relevés from the dendrogram of the cluster anal- ysis were not considered in further analysis since they repre- sented strongly ruderalized stands. Some of these plots were set within the built-up area of the town of Plovdiv, while oth- ers were placed in abandoned vineyards. Diagnostic species of each cluster were determined on the basis of their fidelity (phi) measures using the phi coefficient of association be- tween groups (Chytrý et al. 2002) in the JUICE 7.0 program (Tichý 2002). The size of target groups was fixed at 25% of the total data set. The threshold phi value for the diagnostic species was set at 0.30 and Fisher’s exact test was calculated to exclude those species from diagnostic ones when their fi- delity was not statistically significant (P < 0.001). The results of the classification are summarized in Tab. 1, in which the diagnostic species are ranked according to decreasing fidel- ity. Species are considered as constant if their frequency ex- ceeded 50% for a given group, and species with cover higher than 50% in a minimum of 5% of relevés were considered to be dominant. New syntaxa are named according to the rules of the 3rd edition of the International Code of Phytosocio- logical Nomenclature (Weber et al. 2000). Syntaxonomic no- menclature follows Mucina et al. (2016). Vegetation groups are described on the basis of their di- agnostic, constant and dominant species in the “Results” sec- tion. The values of fidelity, relative frequency and number of relevés for dominants with at least 50% cover are giv- en in brackets for diagnostic, constant and dominant spe- cies respectively. In addition to the syntaxa names of the groups, their geographical names are also included accord- ing to the proposal on the subdivision of the habitat S3-4 Balkan-Anatolian submontane genistoid scrub from the EU- NIS habitat classification into three subhabitat types: S3-41 Balkan-Range submontane genistoid scrub, S3-42 Thracian submontane genistoid scrub and S3-43 Rilo-Rhodopean submontane genistoid scrub (Schaminée et al. 2019; Kunev and Tzonev 2019). For the visualization and interpretation of the relevé groups, a detrended correspondence analysis (DCA) was per- formed using the vegan 2.5–3 package for R (https://cran.r– project.org/web/packages/vegan/index.html). All 151 rele- vés, together with the diagnostic species given in the synoptic table and selected ecological variables, were projected onto two–dimensional ordination space of DCA, with centroids calculated for each of the four clusters. Species ecological in- dicator values (EIVs) for temperature, light, moisture, conti- nentality, soil reaction and nutrients (Pignatti et al. 2005) and selected bioclimatic variables from the WorldClim database (Fick and Hijmans 2017) were used as explanatory ecological variables. EIVs were calculated as unweighted average indi- cator values of taxa for each relevé. All bioclimatic variables available in the WorldClim database were extracted for each relevé on the basis of their geographical coordinates and with the use of the finest spatial resolution provided (30 seconds; ~1 km2). The possible effect of climatic variables on the vege- tation composition was tested by using canonical correspon- dence analysis (CCA) in CANOCO 5.0 software (Ter Braak and Šmilauer 2012) using the global Monte-Carlo permuta- tion test with 999 permutations. Forward selection together with Monte-Carlo permutation test was used to indicate the KUNEV G, TZONEV R, TSIRIPIDIS I, PACHEDJIEVA K 172 ACTA BOT. CROAT. 79 (2), 2020 Tab. 1. Frequency–fidelity synoptic table of the communities of Genista lydia. Frequencies of species are presented as percentages with phi values multiplied by 100 in superscript (group sizes were equalized before the calculation of phi values). Diagnostic species of each cluster are shaded and ranked by decreasing fidelity. Only species with phi values higher than 0.30 are considered to be diagnostic. Species with less than 20% frequency in the whole data set have been omitted. Group No. 1 2 3 4 No. of releves 79 54 14 4 No. of species with less than 20% occurrence 329 272 108 61 Diantho pinifolii-Genistetum lydiae Centaurea rhenana ssp. rhenana 82 82.2 . --- 14 --- . --- Sedum grisebachii 24 43.8 . --- . --- . --- Asperula aristata ssp. scabra 70 40.2 31 --- . --- 25 --- Anthemis tenuiloba 23 37 . --- 7 --- . --- Thymus longicaulis 49 32.8 11 --- 43 --- . --- Dianthus pinifolius ssp. pinifolius 39 31.6 11 --- 14 --- . --- Romuleo graecae-Genistetum lydiae Romulea linaresii ssp. graeca . --- 70 80 . --- . --- Tuberaria guttata . --- 50 65.5 . --- . --- Cerastium gracile . --- 50 65.5 . --- . --- Trifolium tenuifolium . --- 41 58.3 . --- . --- Erodium botrys . --- 37 55.3 . --- . --- Dianthus corymbosus . --- 26 45.6 . --- . --- Galio flavescentis-Genistetum lydiae Anthemis cretica . --- . --- 71 80.8 . --- Sempervivum erythraeum 4 --- . --- 57 65 . --- Hypericum montbretii . --- 4 --- 50 61.3 . --- Galium flavescens . --- . --- 43 60 . --- Verbascum humile 9 --- . --- 50 52.6 . --- Plantago subulata 41 26.6 2 --- 71 42.7 . --- Lembotropis nigricans . --- . --- 21 41.2 . --- Genista lydia and Satureja pilosa community Satureja pilosa . --- . --- 7 --- 75 81.6 Crucianella angustifolia 3 --- . --- 7 --- 75 78.6 Linaria simplex . --- . --- . --- 50 65.5 Cionura erecta . --- . --- . --- 50 65.5 Geranium purpureum . --- . --- . --- 50 65.5 Genistion lydiae Genista lydia 100 --- 100 --- 100 --- 100 --- Minuartia hirsuta ssp. falcata 35 30.3 6 --- 29 --- . --- Centaurea cuneifolia 8 --- 48 44.8 7 --- 25 --- Viola tricolor ssp. macedonica 20 --- 26 --- . --- 25 --- Rorippa thracica 25 --- 11 --- 21 --- . --- Lavandulo stoechadis-Hypericetalia olympici Mucina et 2016 and Cisto-Lavanduletea stoechadis Br.-Bl. in Br.-Bl. et al. 1940 Hypericum olympicum 32 --- 24 --- 21 --- . --- Genista carinalis 22 --- 11 --- . --- . --- Vicia lathyroides 16 --- 13 --- . --- . --- Stachys angustifolia 19 --- 4 --- 7 --- . --- Campanula lingulata 18 --- 4 --- . --- . --- Cistus creticus . --- 19 38.2 . --- . --- Thymus sibthorpii 10 --- 2 --- . --- . --- Festuco-Brometea Br.-Bl. et Tx. ex Soó 1947 Eryngium campestre 58 --- 87 31.4 . --- 100 --- Festuca valesiaca 81 42.8 41 --- 7 --- . --- Sanguisorba minor 48 --- 70 25.2 . --- 50 --- Teucrium chamaedrys 58 24.8 41 --- . --- . --- Cruciata pedemontana 56 --- 41 --- . --- . --- Hieracium hoppeanum 49 --- 28 --- 43 --- . --- Chrysopogon gryllus 23 --- 76 51.6 7 --- . --- Euphorbia cyparissias 43 --- 24 --- 21 --- 100 --- Scabiosa triniifolia 39 --- 24 --- 29 --- . --- GENISTA LYDIA COMMUNITIES IN THE BALKANS ACTA BOT. CROAT. 79 (2), 2020 173 Group No. 1 2 3 4 Achillea coarctata 48 44.6 2 --- 36 --- . --- Hypericum perforatum 39 --- 7 --- 64 --- . --- Dichanthium ischaemum 32 --- 33 --- . --- . --- Thymus glabrescens 29 --- 35 --- . --- . --- Koeleria macrantha 53 67.8 . --- . --- . --- Luzula campestris 20 --- 31 --- . --- . --- Phleum montanum 38 44.1 . --- 21 --- . --- Helianthemetea guttati Rivas Goday et Rivas-Mart. 1963 Aira elegantissima 33 --- 69 36.3 7 --- . --- Trifolium arvense 41 --- 31 --- 14 --- 75 --- Cerastium brachypetalum 27 --- 46 --- . --- . --- Cynosurus echinatus 29 --- 39 --- 14 --- . --- Galium divaricatum 35 --- 31 --- . --- 25 --- Linaria pelisseriana 14 --- 41 32.7 . --- . --- Teesdalia coronopifolia 16 --- 35 --- 7 --- . --- Hypochaeris glabra 8 --- 37 36.5 7 --- . --- Vulpia myuros 13 --- 31 --- . --- . --- Crupina vulgaris 18 --- 20 --- 14 --- . --- Trifolium strictum 8 --- 30 31.2 . --- . --- Ornithopus compressus . --- 39 56.8 . --- . --- Myosotis stricta 8 --- 26 27.2 . --- . --- Vulpia ciliata . --- 35 53.8 . --- . --- Trifolium cherleri 1 --- 26 41.6 . --- . --- Trifolium striatum ssp. striatum 16 --- 4 --- . --- . --- Alyssum minutum 13 --- 4 --- . --- 50 --- Sedum rubens 5 --- 13 --- . --- 75 --- Koelerio–Corynephoretea Klika in Klika et Novák 1941 and Sedo–Scleranthetea Br.–Bl. 1955 Cladonia foliacea 65 --- 52 --- 100 39.4 . --- Rumex acetosella 34 --- 59 --- 64 --- . --- Trifolium campestre 57 35.9 24 --- 7 --- . --- Potentilla argentea 44 --- 30 --- . --- . --- Bromus squarrosus 46 27.5 20 --- . --- 50 --- Potentilla neglecta 46 39.8 11 --- 7 --- . --- Myosotis ramosissima 32 --- 26 --- 14 --- . --- Moenchia mantica 11 --- 50 41.1 14 --- . --- Scleranthus perennis 29 --- 15 --- 43 --- . --- Petrorhagia prolifera 14 --- 35 --- 7 --- 50 --- Filago vulgaris 22 --- 26 --- . --- . --- Chondrilla juncea 24 --- 15 --- . --- 100 --- Cerastium pumilum 34 48.2 2 --- . --- . --- Racomitrium canescens 24 --- 9 --- 14 --- . --- Veronica praecox 16 --- 19 --- . --- . --- Scabiosa argentea 16 --- 15 --- . --- . --- Other species with high frequency Poa bulbosa 51 --- 83 30.5 36 --- 50 --- Plantago lanceolata 51 --- 52 --- 7 --- . --- Anthoxanthum odoratum 30 --- 69 35.2 29 --- . --- Galium verum 33 --- 41 --- . --- . --- Agrostis capillaris 54 58.3 . --- 21 --- . --- Euphorbia niciciana 16 --- 44 27.2 . --- 100 --- Geranium columbinum 22 --- 39 --- 7 --- . --- Leontodon crispus 32 29 11 --- . --- . --- Poa compressa 28 --- 15 --- . --- . --- Sherardia arvensis 4 --- 50 57.2 . --- . --- Hieracium bauhinii 23 --- 19 --- 14 --- . --- Dianthus giganteus ssp. giganteus 33 35.9 4 --- 14 --- . --- Tab. 1. Continued KUNEV G, TZONEV R, TSIRIPIDIS I, PACHEDJIEVA K 174 ACTA BOT. CROAT. 79 (2), 2020 bioclimatic variables most effective on species composition. Rare species were down-weighted in CCA. In both ordina- tion techniques the digital cover–abundance values (numbers from 1 to 9) were used without any further transformation. The three bioclimatic variables with the highest explanatory value calculated in CCA were selected and together with the EIVs according to Pignatti et al. (2005) were passively pro- jected onto the DCA-ordination diagram. The taxonomic nomenclature of the vascular plants fol- lows Delipavlov and Chesmedzhiev (2011), except for the genera Juniperus L., Genista L., Cistus L., Anthemis L., Tol- pis Adans. and Tripleurospermum Sch. Bip., for which some additional sources were used, mostly from other Bulgari- an floras (Jordanov et al. 1963-2012; Assyov and Petrova 2012) or occasionally, on-line databases (Roskov et al. 2006, The Plant List 2013, Euro+Med 2006-2018). Nomenclature of mosses follows Hill et al. (2006); lichens are given after Mayrhofer et al. (2005). Chorological and life-form spectra were calculated for each vegetation unit and are presented as percentages on the basis of presence of a species (On–line Suppl. Tab. 1). The chorotypes were initially indicated in ac- cordance with Assyov and Petrova (2012) and then grouped in 10 larger categories for better representation of the general trends in the phytogeographical relationships. The choro- types of lichens and mosses have not been included in these calculations due to insignificant species richness of these taxa in the communities of Genista lydia. The biological spectrum is presented following the classification of Raunkiær (1934). The endemic taxa are given after Petrova and Vladimirov (2010), Assyov and Petrova (2012) and Euro+Med PlantBase (2006-2018). The soil types and composition of the bedrock outcrops are presented in accordance with the soil (Ninov 2002) and the geological map of Bulgaria (Cheshitev and Kânčev 1989). Analysis of representative soil samples taken from selected relevés were performed in an authorized labo- ratory – NIK Agro Service Ltd. The samples were analyzed (see Teoharov et al. 2009) in accordance with ISO standards as follows: pH (ISO 10390), specific electrical conductivity (ISO 11265), total organic carbon (ISO 10694) and carbon- ate content (ISO 10693). Results The floristic composition of the communities of Ge- nista lydia is very rich and includes 580 taxa. The average number of species per plot is 46. The mean vegetation cov- er is 80.14%. The elevation is significantly variable, ranging between 150 and 1500 m a.s.l. The slope inclination var- ies between 1°- 60° (mean 10°), while the exposure is most frequently southern. The soil sample analysis (Tab. 2) dem- onstrates that in most localities the soils have a sandy texture. They are slightly to moderately acidic (pH 5.02 – 7.13), with conductivity ranges between 0.02 and 0.13 dS m-1, while the carbonate content is represented only as traces. The amount of soil organic carbon varies significantly (0.24 to 10.03%), although it rarely exceeds (3%), which is an indicator for their low nutrition content. The cluster analysis resulted in the distinction of four groups (Fig. 1). Two of them are found only in the territory of Bulgaria, while the rest are also found in Northern Greece. These four units are geographically, ecologically and floris- tically well defined. The differences between them are also supported by the DCA-ordination diagram (Fig. 2). The in- terpretation of the DCA-diagram is assisted by the passively projected EIVs and the three bioclimatic variables with the highest explanatory value. Analysis with forward selection in CCA provided a ranking of the relative importance of the tested bioclimatic variables (On–line Suppl. Tab. 2). All 19 bioclimatic variables were statistically significant (P < 0.001) and explained 31.19% of the total variance of the data set. The three with strongest effect on vegetation composition are precipitaton of warmest quarter (Bio 18), precipitation seasonality (Bio 15) and mean temperature of coldest quar- ter (Bio 11). Tab. 2. Soil properties of representative plots occupied by the communities of Genista lydia. EC – specific electrical conductivity (dS m-1), and C org – total organic carbon content (%). Group 1 – Diantho pinifolii-Genistetum lydiae, Group 2 – Romuleo graecae-Genistetum lydiae, Group 3 – Galio flavescentis-Genistetum lydiae. Relevé no. Soil depth (cm) pH EC (dS m–1) C org (%) CaCO₃ (%) Group 1 59 15–30 6.66 0.02 0.45 0.41 65 15–30 6.48 0.02 0.4 Traces 70 0–5 5.08 0.09 9.22 Traces 71 0–5 6.09 0.03 1.85 0.03 100 5–15 6.65 0.02 0.44 0.08 102 5–15 6.22 0.02 0.7 Traces 111 0–5 6.8 0.02 0.46 Traces 112 15–30 5.38 0.02 1.79 Traces 119 5–15 7.13 0.02 0.24 Traces Group 2 13 15–30 5.39 0.02 1.65 Traces 114 0–5 6.52 0.04 0.51 Traces 133 5–15 5.62 0.02 2.61 Traces Group 3 128 0–5 5.34 0.13 8.9 9.57 135 0–5 5.02 0.07 10.03 Traces GENISTA LYDIA COMMUNITIES IN THE BALKANS ACTA BOT. CROAT. 79 (2), 2020 175 The first axis on Fig. 2 can be interpreted as an altitudi- nal gradient, positively correlated also with continentality and precipitation, the latter being represented by the pre- cipitation of the warmest quarter. The other two bioclimat- ic variables plotted in the ordination space are more or less negatively correlated with the first DCA axis. These are the temperature of the coldest quarter and precipitation sea- sonality. The second DCA axis is probably related to soil moisture as revealed by the corresponding environmental parameter used in the DCA-diagram, as well as from the discrimination at the top of this axis of the Genista lydia- Satureja pilosa community (group 4) which grows on well drained substrates of sandy gravel bars. Description of the communities Group 1: Association Diantho pinifolii-Genistetum lydiae ass. nova hoc loco (On–line Suppl. Tab 1), holotypus rel. 55 (Appendix, Fig. 3a) (Rilo–Rhodopean submontane genistoid scrub) Diagnostic species: Centaurea rhenana ssp. rhenana (82.2), Sedum grisebachii (43.8), Asperula aristata ssp. scabra (40.2), Anthemis tenuiloba (37), Thymus longicaulis (32.8), Dianthus pinifolium ssp. pinifolius (31.6) Constant species: Genista lydia (100), Centaurea rhenana ssp. rhenana (82), Festuca valesiaca (81), Asperula aristata ssp. scabra (70), Cladonia foliacea (65), Teucrium chamae- drys (58), Eryngium campestre (58), Trifolium campestre (57), Cruciata pedemontana (56), Agrostis capillaris (54), Koeleria macrantha (53), Poa bulbosa (51), Plantago lanceolata (51) Dominant species: Genista lydia (20) Distribution: Mesta (Nestos) River valley in Bulgaria and Greece, Central Rhodope Mts., 600 – 1500 m a.s.l. This association is distributed on volcanic rocks and soils with a sandy structure. Its altitudinal range corresponds to submontane and montane belt on slopes with more pro- nounced inclinations (mean 14.5°). The soils are Umbric Leptosols and less often Dystric Cambisols (especially in the Central Rhodope Mts.). This group is characterized by its more frequent occurence at higher altitudes and in sites with higher annual precipitation, lower annual temperatures and less pronounced summer drought, as compared to all other groups (On–line Suppl. Tab. 3). The influence of cold- er and more humid mountain climate is a reason for more significant participation of some European and Boreal spe- cies as well as more xeromesophytes or mesophytes, such as Agrostis capillaris, Brachypodium sylvaticum, Arrhenatherum elatius, Vicia incana and Trifolium medium. The herbaceous layer is dominated by hemicryptophytes (46.8%), although therophytes are still well represented (34.5%) (Figs. 4, 5). There is also high number of Balkan endemics and suben- demics (13.04%), such as Viola aetolica, Sedum grisebachii, S. stefčo, Silene velenovskyana and Anthemis macedonica. Fig. 1. Cluster dendrogram of Genista lydia communities. Group 1 – Diantho pinifolii-Genistetum lydiae, Group 2 – Romuleo graecae-Geniste- tum lydiae, Group 3 – Galio flavescentis-Genistetum lydiae, Group 4 (G4) – Genista lydia-Satureja pilosa communities, X – excluded plots. KUNEV G, TZONEV R, TSIRIPIDIS I, PACHEDJIEVA K 176 ACTA BOT. CROAT. 79 (2), 2020 Group 2: Association Romuleo graecae-Genistetum lydiae ass. nova hoc loco (On–line Suppl. Tab 1), holoty- pus rel. 131 (Appendix, Fig. 3b) (Thracian submontane ge- nistoid scrub) Diagnostic species: Romulea linaresii spp. graeca (80), Tuberaria guttata (65.5), Cerastium gracile (65.5), Trifolium tenuifolium (58.3), Erodium botrys (55.3) and Dianthus cor- ymbosus (45.6) Constant species: Genista lydia (100), Eryngium camp- estre (87), Poa bulbosa (83), Chrysopogon gryllus (76), Ro- mulea linaresii spp. graeca (72), Sanguisorba minor (70), An- thoxanthum odoratum (69), Aira elegantissima (69), Rumex acetosella (59), Plantago lanceolata (52) and Cladonia folia- cea (52) Dominant species: Genista lydia (15) Distribution: Eastern Rhodope Mts., both in Bulgaria and Greece, on schist, sandstones, tuffs, 150 – 600 m a.s.l. The association occupies sandy soils, sandstones or min- ing deposits composed of white zeolites. The soil types are mostly Lithic and Umbric Leptosols. The precipitation re- gime has a winter maximum in November and a well pro- nounced summer drought between July and September (On–line Suppl. Tab. 3). The Mediterranean climate influ- ence is significant in this association, and possibly is one of the reasons for it having the highest species richness amongst all groups. This is justified by the high frequency of therophytes (47.18%), and Mediterranean and sub-Med- iterranean species (38%) (Figs. 4, 5). In early spring, the her- baceous layer is dominated by geophytes like Romulea lina- resii ssp. graeca, Ornithogalum spp. and Crocus chrysanthus. Therophytes with a typical Mediterranean origin dominate from the late April till the end of May, such as many annual clovers, Vulpia ciliata, Tuberaria guttata, Molineriella minu- ta, Aira elegantissima and Aegilops neglecta. In the summer period, the flowering aspect is influenced by yellow flower- Fig. 2. Detrended correspondence analysis (DCA) spider-plot of the 151 relevés classified in the four groups distinguished. The length of the first DCA axis is 3.31 SD units, while of the second axis 3.80 SD units. Numbers refer to the centroids of vegetation groups correspond- ing to: 1 – Diantho pinifolii-Genistetum lydiae, 2 – Romuleo graecae-Genistetum lydiae, 3 – Galio flavescentis-Genistetum lydiae, 4 – Genista lydia-Satureja pilosa communities. Ecological indicator values and bioclimatic variables are represented by arrows (temp – temperature, light – light, pH – soil reaction, cont – continentality, moist – moisture, nutr – nutrients, Bio 15 – precipitation seasonality, Bio 11 – mean temperature of coldest quarter, Bio 18 – precipitation of warmest quarter). Diagnostic species for the relevant groups are represented by dots and abbreviated names as follows: Anthcre – Anthemis cretica, Anthten – Anthemis tenuiloba, Aspesca – Asperula aristata ssp. scabra, Centrhe – Centaurea rhenana ssp. rhenana, Ceragra – Cerastium gracile, Cionere – Cionura erecta, Crucang – Crucianella angustifolia, Di- ancor – Dianthus corymbosus, Dianpin – Dianthus pinifolius ssp. pinifolius, Erodbot – Erodium botrys, Galifla – Galium flavescens, Genicar – Genista carinalis, Genilyd – Genista lydia agg., Gerapur – Geranium purpureum, Hypemon – Hypericum montbretii, Hypeoly – Hypericum olympicum, Lembnig – Lembotropis nigricans, Linasim – Linaria simplex, Plansub – Plantago subulata, Romugrae – Romulea linaresii ssp. graeca, Satupil – Satureja pilosa, Sedugri – Sedum grisebachii, Sempery – Sempervivum erythraeum, Thymlon – Thymus longicaulis, Triften – Trifolium tenuifolium, Tubegut – Tuberaria guttata, Verbhum – Verbascum humile. GENISTA LYDIA COMMUNITIES IN THE BALKANS ACTA BOT. CROAT. 79 (2), 2020 177 ing Compositae species like Hypochaeris glabra, H. cretensis, H. radicata and Crepis setosa. The endemics are not so fre- quent/abundant, as compared to the other groups. Balkan endemics recorded in the association are Armeria rumelica, Chamaecytisus jankae, Daucus guttatus ssp. zahariadii, Di- anthus corymbosus, Anthemis virescens and Romulea linare- sii ssp. graeca. Group 3: Galio flavescentis-Genistetum lydiae ass. nova hoc loco (On–line Suppl. Tab 1), holotypus rel. 41 (Appen- dix, Fig. 3c) (Balkan–Range submontane genistoid scrub) Diagnostic species: Anthemis cretica (80.8), Sempervivum erythraeum (65), Hypericum montbretii (61.3), Galium fla- vescens (60.0), Verbascum humile (52.6), Plantago subulata (42.7) and Lembotropis nigricans (41.2) Constant species: Genista lydia (100), Cladonia foliacea (100), Polytrichum pilifermum (71), Plantago subulata (71), Anthemis cretica (71), Rumex acetosella (64), Hypericum per- foratum (64), Festuca dalmatica (64) and Viola arvensis (57) Dominant species: Genista lydia (7) Distribution: Southern slopes of East Balkan Range (Sliv- en district) and East Rhodope Mts., 300 – 1000 m a.s.l. The communities of this association mostly occur on acidic rocks such as andesites, diorites, conglomerates. They are found on rock cliffs and terraces with a predominantly southern exposure. The soils are shallow Umbric Leptosols. Although this association demonstrates a more pronounced continental character, the number of Mediterranean spe- cies in their floristic composition continues to be signifi- cant (34.93%) (Fig. 4). On the other hand, the annual species (28.77%) are fewer in number than other groups. Anoth- er specific feature of the group is the higher frequency and abundance of shrub and juvenile tree species (Fig. 5) such as Syringa vulgaris, Lembotropis nigricans and Fraxinus ornus. This is due to the abandonment of grazing practices, since this association is mostly located in some protected areas such as Sinite Kamuni Nature Park. These communities are also rich in Balkan endemics and subendemics (17.81%), like Sesleria latifolia, Sempervivum erythraeum, Verbascum humile, Chamaecytisus calcareus, Scabiosa triniifolia and Si- lene lerchenfeldiana. Group 4. Genista lydia-Satureja pilosa community (Fig. 3d) Diagnostic species: Satureja pilosa (81.6), Crucianella an- gustifolia (78.6), Linaria simplex (65.5), Geranium purpure- um (65.5) and Cionura erecta (65.5) Constant species: Genista lydia (100), Euphorbia nicici- ana (100), Euphorbia cyparissias (100), Eryngium campestre (100), Chondrilla juncea (100), Vicia cracca (75), Trifolium arvense (75), Senecio vulgaris (75), Sedum rubens (75), Sat- ureja pilosa (75), Orlaya grandiflora (75), Melica ciliata (75), Medicago minima (75), Linaria genistifolia ssp. genistifolia Fig. 3. Physiognomy of Genista lydia communities: a – Diantho pinifolii-Genistetum lydiae, b – Romuleo graecae-Genistetum lydiae, c – Galio flavescentis-Genistetum lydiae, d – Genista lydia-Satureja pilosa community. KUNEV G, TZONEV R, TSIRIPIDIS I, PACHEDJIEVA K 178 ACTA BOT. CROAT. 79 (2), 2020 (75), Koeleria nitidula (75), Geranium rotundifolium (75), Galium lucidum (75), Euphorbia myrsinites (75), Crucianel- la angustifolia (75), Buglossoides arvensis (75) and Asperula purpurea (75) Dominant species: Genista lydia (25) Distribution: East Rhodopes Mts., 150 – 300 m a.s.l. This community is distributed on the alluvial sandy ter- races of Arda River and its tributaries. Typical species are many psamophytes as well as some species that inhabit the gravel bars like Satureja pilosa, Cionura erecta, Euphorbia niciciana, etc. The highest proportion of therophytes and the absence of geophytes in this community could be explained through its development on unstable (pioneer) substrates, influenced by river floods (Fig. 5). The floristic composition is also highly influenced by trampling by cattle and sheep that take water from the streams of the Arda River. area of occupancy (AOO) – 26 (2600 km²) and extent of oc- currence (EOO) – 9670 km² (Kunev and Tzonev 2019). Their distribution in Greece has not been completely investigated. However, the data presented in the habitat's description from the European Red List of Habitats (Janssen et al. 2016) shows a wider distribution area, which is 20 km² for the territory of Greece and 60 km² in Bulgaria. These data suggests that the occupied area in the Red List is probably overestimated. The studied shrub communities could develop as pri- mary vegetation on extreme sites, such as sheer rocks, rock crevices and terraces, where the soil is often absent or very shallow. However, secondary expansion of these commu- nities has been also observed in some areas. It is initiated largely due to habitat degradation as a result of overgrazing, deforestation and following soil erosion. Those secondary communities are distributed mostly close to the settlements and in extensively grazed areas. The soil layer is more devel- oped there, although it rarely reaches 30 cm in depth. The present distribution of the studied vegetation has probably resulted also from some historical events. These communities occur mostly on the slopes of the Mesta (Nes- tos) and Arda River Valleys in Southern Bulgaria and North- ern Greece and also penetrate into the valleys of their trib- utaries. Before the 1950s, these river valleys were used as transition route during the annual migration of the livestock from the winter grazing sites (in the lowlands) to the sum- mer pasturelands (in the mountains). This type of grazing, known as “transhumance”, was widespread until 19th century in Bulgaria (Stoynov 2008). Therefore, probably the recent distribution and abundance of G. lydia communities in these areas is also related with the trampling and seed dispersal by the grazing animals. This fact is also supported by the abun- Fig. 4. Proportion of chorotypes within the different units of the Genista lydia communities: 1 – Diantho pinifolii-Genistetum lydiae, 2 – Romuleo graecae-Genistetum lydiae, 3 – Galio flavescentis-Ge- nistetum lydiae, 4 – Genista lydia-Satureja pilosa community, 5 – The last column represents the proportion of chorotypes within all community types from the dataset. Chorotypes abbreviation: Msm – Mediterrenean and sub-Mediterrenean, Eur – European, Pal – Palearctic, Bor – Boreal, Psp – Pontic and sub-Pontic, Be – Balkan endemic, Bse – Balkan subendemic, Kos – Cosmopoli- tan. The Adventive and Holarctic elements are insignificant part of compared chorotypes, thus they are not marked in the figure. Fig. 5. Biological spectrum of the Genista lydia communities: 1 – Diantho pinifolii-Genistetum lydiae, 2 – Romuleo graecae-Geniste- tum lydiae, 3 – Galio flavescentis-Genistetum lydiae, 4 – Genista lydia-Satureja pilosa community, 5 – The last column represents the proportion of life-forms of community types 1, 2, 3 and 4. Life- form abbreviations: Ph – Phanerophytes, Ch – Chamaephytes, H – Hemicryptophytes, G – Geophytes, Th – Therophytes. Discussion The largest areas occupied by Genista lydia communities are on the slopes of Pirin, Rila and the Rhodope Mountains. (in both Bulgarian and Greek parts). The northernmost lo- calities are on the southern slopes of the Eastern Balkan Mountains. (Fig. 6). The communities of G. lydia are known to have very lim- ited distribution in Bulgaria covering a total area of 5.2 km², GENISTA LYDIA COMMUNITIES IN THE BALKANS ACTA BOT. CROAT. 79 (2), 2020 179 Fig. 6. Distribution map of Genista lydia communities: ● – Diantho pinifolii-Genistetum lydiae, ○ – Romuleo graecae-Genistetum lydiae, ▲ – Galio flavescentis-Genistetum lydiae, ▲ – Genista lydia-Satureja pilosa community. dance of G. lydia communities mostly close to the settle- ments where these grazing practices are still maintained. In the mountainous areas, these communities are often found near shieling sites. At the same time, in the sites with low or zero grazing intensity, the phytocoenoses have lower density and patchy distribution (Kunev and Tzonev 2019). The information for the distribution of the studied com- munities on the territory of other Balkan countries and Tur- key is very limited. G. lydia is reported mostly as an accom- panying species in various vegetation types, but mostly forests and grasslands. For example, this species in Serbia is distributed in the scrublands or forest fringes of the mountain regions (Diklić 1972). The species also has been reported to participate in the floristic composition of the association Trifolio-Trisete- tum flavescentis N. Ranđelović 1975 (Chrysopogono-Dantho- nion calycinae Kojić 1957 of Festuco-Brometea) from Central Serbia (Aćić et al. 2014). After revision of herbaria colections Teofilovski (2011) concluded, based on Zieliński et al. (2004), that part of the specimens previously cited by Micevski (2001) as G. lydia, actually belong to G. januensis. Therefore, the occurrence of G. lydia in North Macedonia should be accepted as un- certain. Genista lydia in Greece is distributed mainly in the north- eastern parts of the country (Strid 1986, Zieliński et al. 2004). It has been reported as a common species in the forest clear- ings from the Rhodope Mountains., as well in the shrub layer of forests dominated by Betula pendula and Pinus sylvestris (Theodoropoulos et al. 2003, Eleftheriadou et al. 2009). It is also reported as diagnostic species to the associations Cen- taureo affinis-Festucetum koritnicensis Karagiannakidou et al. 2001, especially in its high–altitudinal variant described from Pangeon Mt., where it occupuies schistose substrates at altitudes of 1500–1800 m a.s.l. However, this association is classified in the alliance of oromediterranean dry grass- lands and thorhy–cushion dwarf shrub – Astragalo angusti- folii-Seslerion coerulantis Quézel 1964 from the class Daph- no-Festucetea Quézel 1964 (Karagiannakidou et al. 2001). The association Minuartio hirsutae-Dianthetum pinifo- lii (variant with G. lydia) has been described from Nomos of Chalkidiki, Greece (Bergmeier et al. 2009). As diagnos- tic species are referred Centaurea diffusa, C. grisebachii, Di- anthus pinifolius, Hypericum olympicum, Minuartia hirsuta ssp. falcata, Phleum phleoides, Psilurus incurvus, Rumex ace- tosella and Thymus sibthorpii. Bergmeier et al. (2009) also noted that these communities inhabit slightly acidic, skeletal soils and classified them into the alliance Diantho pinifolii- Jasionion heldreichii Bergmeier et al. 2009 of Sedo-Scleran- thetea. These communities differ from the studied vegeta- tion by poorer species composition and lower abundance of therophytes. The distribution range of G. lydia in Turkey includes its European parts as well as the western Asia Minor, and es- pecially mountainous areas near the coastline (Gibbs 1970, Zieliński et al. 2004). The species occurs mostly on schistose substrates and participates in the floristic structure of mac- chia and phryganic vegetation, as understory of degraded mixed deciduous or coniferous forests and even in subalpine shrub communities (Quézel and Pamukçuoǧlu 1970, Qué- zel 1986, Uğurlu and Senol 2005, Kaya and Aladaǧ 2009). KUNEV G, TZONEV R, TSIRIPIDIS I, PACHEDJIEVA K 180 ACTA BOT. CROAT. 79 (2), 2020 However, the community Genista lydia-Hypericum li- narioides described from the Mount Uludağ (Quézel and Pamukçuoǧlu 1970) is the only unit reported from Tur- key with definitive dominance or codominance of G. lyd- ia. These dwarf shrubs occur on schistose substrates, on 1800–2200 m a.s.l., at the fringes of Abies bornmuelleriana forests. From the pointed diagnostic species only Juniperus nana and Thymus sibtorpii are presented also in the Bulgar- ian and Greek communities of G. lydia, but with a low con- stancy. Quézel and Pamukçuoǧlu (1970) have also noted the secondary origin of the community, which probably is relat- ed to the grazing in the subalpine zone. The proposed classi- fication of the unit in the alliance Bruckenthalion (Vaccinio- Piceetalia) may be considered as provisional because it was based on four relevés only. The diagnostic value of G. lydia communities for other, hierarchically higher, syntaxa is very poorly known. For ex- ample, according to Stefanov (1921), G. lydia (referred as G. rumelica) is an evergreen species from the pseudomaquis formations in the Western Thrace region and Northeastern Greece. The dominants in this type of vegetation are also Pis- tacia terebinthus, Juniperus oxycedrus, J. excelsa, Cistus cre- ticus, Calicotome villosa, etc. Specific characteristic of these vegetation formations are also the abundance of annuals and geophytes, especially in more disturbed locations. Typ- ical species mentioned by Stefanov (1921), such as Teesdalia coronopifolia, Trifolium subterraneum, Ornithopus compres- sus, Crepis zacintha, Plantago bellardii, Orchis papilionacea, Ranunculus paludosus, Campanula phrygia, are also found in the stands of G. lydia studied here. The shrub communities, dominated by Cistus incanus from the southern parts of Bulgaria demonstrate also very similar ecological and floristic structure to those of G. lyd- ia. Gussev (2015) suggested that they have some interme- diate characteristics between the two plant associations de- scribed from neighbouring countries: Diantho-Cistetum incani Micevski et Matevski 1984 and Calicotomo villosae- Cistetum cretici Oberd. 1954. For example, the first associ- ation from the southern regions of North Macedonia was classified within the alliance Trifolion cherleri Micevski 1972 of Festuco-Brometea (Micevski and Matevski 1984, Čarni et al. 2010). Some of its diagnostic species such as Cistus in- canus, Dianthus pinifolius, Micropyrum tenellum, Anthemis macedonica, Briza maxima, Scabiosa triniifolia, Lupinus an- gustifolius have been also found in the communities of G. lydia. A common feature to both vegetation types is the high constancy of leguminosae and especially Trifolium species. On the other hand, the association Calicotomo villosae- Cistetum cretici Oberd. 1954 from Thessaloniki, Thrace and Thessaly, North Greece, dominated by the more thermoph- ilous C. incanus ssp. creticus, could be a good example for communities synvicariant to G. lydia. The communities oc- cur on silicate substrates and some of them are maintained by grazing. The association is characterized by the presence of widespread mesophilous species such as Hypericum per- foratum, Brachypodium sylvaticum, Plantago lanceolata and Teucrium chamaedrys. Some early spring annuals like Aira elegantissima, Cerastium brachypetalum, Cynosurus echina- tus and Vulpia ciliata also participate in its floristic struc- ture (Oberdorfer 1954, Čarni et al. 2010). Additionally, in the mosaics of these shrub communities, Oberdorfer (1954) sampled grassland fragments with high abundance of Poa bulbosa and other therophytes and geophytes - Romulea li- naresii ssp. graeca, R. bulbocodium, Ornithogalum spp., Ga- gea spp. According to Oberdorfer (1954), these grasslands represent a regresion stage of the shrub communities, such as Calicotome villosa-Cistetum cretici Oberd. 1954 caused by overgrazing and trampling. Although this association is dominated by some typical Mediterranean species not pres- ent in the G. lydia communities, both units have a similar appearance, floristic and ecological structure. Different vegetation types, occuring in the transitional zone between the Temperate and Mediterranean climatic regions of the Southern Balkans were also reported to have such a complex floristic structure (Stoyanov and Achtarov 1951, Sopotlieva and Apostolova 2014, Čarni et al. 2018). From these data, it could be concluded that complexity and intermediate state (mixture of low shrubs, perennial and an- nual herbs with Continental-Temperate and Mediterranean origin) of the communities of G. lydia are common phenom- enon in these regions. Their specific structure is determined by the transitional position between the two climatic types as well as the variability of their altitudinal range. Other impor- tant factors influencing the studied communities are grazing pressure, deforestation, construction and mining activities. The complexity and the mixed structure of the open shrubland communities in these transitional areas make them difficult to be classified on the higher than the associ- ation level. For example, Oberdorfer (1954), who described the association Calicotomo villosae-Cistetum cretici, classified it in the alliance Cistion orientale Oberd. 1954 of the class of "eastern Mediterranean hedgehog heaths and low–grown broom phryganas” – Cisto-Micromerietea julianae Oberd. 1954 (see also Čarni et al. 2010). However, the name Cistion orientale was illegitimate and thus was replaced by Hyperico olympici-Cistion cretici (Oberd. 1954) R. Jahn et Bergmeier (Mucina et al. 2009). According to Mucina et al. (2016), this alliance belongs to the class Cisto-Lavanduletea stoechadis Br.-Bl. in Br.-Bl. et al. 1940. The similarities of the association Calicotome villosa-Cistetum cretici with the western mediter- ranean phryganic communities of Cisto-Lavanduletea have been also noted by Oberdorfer (1954). Genista lydia is al- so a low, evergreen to semi–evergreen shrub, which domi- nates communities developed mostly on siliceous substrates. This indicates the affiliation of such communities to the class Cisto-Lavanduletea stoechadis and its Eastern Mediterranean order Lavandulo stoechadis-Hypericetalia olympici. The diag- nostic species of the order, found in the studied communi- ties, are Cistus creticus, Anthemis cretica, Dianthus pinifolius, Hypericum olympicum, Genista carinalis and Stachys angus- tifolia. Moreover, it was recently reported from the south- ern regions of the Crimean peninsula (Ryff 2018), a fact that presents the order as notstrictly Mediterranean. The order consists of three alliances, from which Hyperico olympici- Cistion cretici Mucina et al. (2009) was described as ther- GENISTA LYDIA COMMUNITIES IN THE BALKANS ACTA BOT. CROAT. 79 (2), 2020 181 mo-meso-Mediterranean silicicolous phrygana of Northern Hellas. This alliance is the most floristically, biogeographi- cally and ecologically similar to the studied communities. From the characteristic species of the alliance, Campanula lingulata, Vicia lathyroides and Thymus sibthorpii (Čarni et al. 2010) have been also recorded in the G. lydia communi- ties. However, the studied vegetation could be determined as submediterranean silicicolous garigue distributed in south- ern parts of Central and Eastern Balkans, as well as locally in West Anatolia. Their floristic composition, structure and richness in Balkan endemics are sufficient to differentiate them from the already described syntaxa on the alliance level and to justify the establishment of a new independent alli- ance, which is described below. Alliance: Genistion lydiae all. nov hoc loco Holotypus hoc loco: Romuleo graecae-Genistetum lydiae Kunev et all. nov. hoc loco Diagnosis: Low-growing shrub communities dominat- ed by the evergreen to semi-evergreen species G. lydia dis- tributed in the south-eastern part of the Balkan Peninsula and probably Western Anatolia. These communities are aci- dophilus and occupy eroded places with shallow, poor soils or also rocky sites. The bedrock is of volcanic origin (andes- ites, rhyolites, volcanic schist, zeolites, gneisses), but also of sandstones. These communities occur as primary vegetation on rocky areas, but may also have a secondary origin re- placing other communities on overgrazed grasslands. They are often open and with rich floristic composition which in- cludes different plant life-forms. Many typical herbaceous species of the class of perennial grasslands Festuco-Brom- etea (in the foothill and mountain areas) and of the class of the therophytic pseudosteppes - Helianthemetea guttati (in the areas with a stronger Mediterranean climatic influ- ence) participate in the stands of G. lydia. Such species are Festuca valesiaca, F. dalmatica, Agrostis capillaris, A. castel- lana, Poa bulbosa, Chrysopogon gryllus, Koeleria macrantha, Aira elegantissima, Eryngium campestre, Sanguisorba minor, Trifolium cherleri and T. subterraneum. In the canopy gaps of more open stands or on steep slopes of river valleys ma- ny ecologically flexible species, which generally belong to Koelerio-Corynephoretea and Sedo-Scleranthetea classes can be also found, such as Rumex acetosella, Bromus squarosus, Scabiosa argentea, Scleranthus perennis, Erysimum diffusum, Myosotis ramosissima, Sedum spp., etc. Characteristic species: Genista lydia, Satureja pilosa, Thy- mus atticus, Hypericum montbretii, Anthemis tenuiloba, Ga- lium flavescens, Minuartia hirsuta ssp. falcata, Centaurea cu- neifolia, Viola tricolor ssp. macedonica and Rorippa thracica. Transgressive and differential species: Asperula arista- ta ssp. scabra, Carlina corymbosa, Centaurea rhenana ssp. rhenana, Chamaecytisus absinthioides, Daucus guttatus, Di- anthus pinifolium ssp. pinifolius, Erodium botrys, Romulea li- naresii spp. graeca, Sedum grisebachii, Tuberaria guttata, Tri- folium tenuifolium and T. strictum. This new alliance is similar and close to the alliance Hy- perico olympici-Cistion cretici, but it is differentiated by its bio- geographical range (Southeast Balkans and adjacent regions of Asia Minor), its transitional position between Mediterra- nean and Continental (Temperate) climatic types and its flo- ristic composition rich in Balkan endemics and subendemics. Proposed syntaxonomic scheme Class Cisto-Lavanduletea stoechadis Br.-Bl. in Br.-Bl. et al. 1940 Order Lavandulo stoechadis-Hypericetalia olympici Mu- cina in Mucina et al. 2016 Alliance Genistion lydiae Kunev, Tzonev, Tsiripidis et Pachedjieva all. nov. hoc loco Association Diantho pinifolii-Genistetum lydiae Kunev, Tzonev, Tsiripidis et Pachedjieva ass. nova hoc loco Association Romuleo graecae-Genistetum lydiae Kunev, Tzonev, Tsiripidis et Pachedjieva ass. nova hoc loco Association Galio flavescentis-Genistetum lydiae Kunev, Tzonev, Tsiripidis et Pachedjieva ass. nova hoc loco Genista lydia-Satureja pilosa community Conclusion The current study is the first attempt to describe appro- priately the syntaxonomy of Genista lydia communities. The establishment of an endemic Southeast Balkan-West Anato- lian alliance of sub-Mediterranean low evergreen to semi-ev- ergreen scrub vegetation is proposed. The G. lydia commu- nities have conservation significance on the European scale (Janssen et al. 2016). The major threat to the habitat is the abandonment of traditional grazing and pasture manage- ment as well as the changes in the land use. The latter threat concerns the transformation of pasturelands into crop fields or artificial forest plantations. If grazing is suspended, rapid successional changes will lead to the complete replacement of the shrub communities with hemicryptophytic grasslands or tall scrublands. Therefore, certain conservation measures, such as specific grazing maintenance, should be applied in order to prevent the decline of the communities. The inclu- sion of the “Balkan–Anatolian submontane genistoid scrub” habitat in the Annex I of the Habitat Directive is considered as a necessary policy measure that can ensure the conserva- tion of this rare and vulnerable habitat type. Acknowledgements The research was partly financed by the project: “Syn- taxonomic characteristics of the Rumelian (Genista rumelica Velen.) and Lydian (Genista lydia Boiss.) greenweed com- munities in Bulgaria and North Greece” – Contract №80.10– 65/19.04.2018 funded by the Scientific Research Fund of the Sofia University “St. Kliment Ohridski”. We are grateful to Assoc. Prof. Anna Ganeva, PhD for the determination of the bryophytes and also to Assist. Prof. Veselin Shivarov, PhD, for the determination of the lichenized fungi in all studied communities. We are also grateful to Gabriela Petrova, PhD student, for the preparation of the distribution map on the studied vegetation type. KUNEV G, TZONEV R, TSIRIPIDIS I, PACHEDJIEVA K 182 ACTA BOT. CROAT. 79 (2), 2020 References Aćić, S., Šilc, U., Jovanović, S., Kabaš, E., Vukojičić, S., Dajić Stevanović, Z., 2014: Nomenclatural revision of dry grass- land syntaxa of the Central Balkan. Tuexenia 34, 355–390. Assyov, B., Petrova, A. 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Kunev, 23.06.2016; plot size 25 m²; alt. 739 m a.s.l.; slope: 20°, S; veg- etation cover: 70%; latitude: 41.82577°, longitude: 23.62033° Shrub species: Genista lydia 3; Herbaceous species: Festu- ca valesiaca 2a, Sanguisorba minor 2a, Thymus longicaulis 2a, Dianthus pinifolius ssp. pinifolius 1, Agrostis capillaris 1, Bro- mus squarrosus 1, Centaurea rhenana ssp. rhenana 1, Chryso- pogon gryllus 1, Crupina vulgaris 1, Cynosurus echinatus 1, Eryngium campestre 1, Euphorbia cyparissias 1, Euphrasia pectinata 1, Geranium columbinum 1, Hieracium bauhinii 1, Hypericum rumeliacum 1, Koeleria macrantha 1, Leontodon crispus 1, Melica ciliata 1, Orlaya daucoides 1, Phleum monta- num 1, Plantago lanceolata 1, Plantago subulata 1, Potentilla neglecta 1, Stachys angustifolia 1, Stipa pulcherrima 1, Tae- niatherum caput-medusae 1, Teucrium chamaedrys 1, Trifo- lium campestre 1, Trifolium hirtum 1, Trifolium striatum ssp. striatum 1, Xeranthemum annuum 1, Achillea coarctata +, Acinos rotundifolius +, Aegilops triuncialis +, Anagallis ar- vensis +, Anthemis tenuiloba +, Asperula aristata ssp. sca- bra +, Campanula lingulata +, Cerastium brachypetalum +, Cerastium pumilum +, Cuscuta approximata +, Hypericum olympicum +, Logfia arvensis +, Nigella arvensis +, Onobry- chis gracilis +, Orlaya grandiflora +, Potentilla argentea +, Psi- lurus incurvus +, Scleranthus perennis +, Sedum caespitosum +, Sedum rubens +, Thesium arvense +, Trifolium arvense +, Valerianella coronata +, Valerianella dentata + 2. Romuleo graecae-Genistetum lydiae ass. nova hoc loco Holotypus hoc loco: Bulgaria, Dedets; Relevé no. 131 (On– line Suppl. Tab 1); relevé collector: G. Kunev, 22.04.2018; plot size 25 m²; alt. 382 m a.s.l.; slope: 3°, SW; vegetation cover: 80%; latitude: 41.38857°, longitude: 25.22668° Shrub species: Genista lydia 3, Cistus creticus 2a, Juniper- us communis ssp. communis 1, Juniperus deltoides 1, Chamae- cytisus albus +, Rosa turcica +; Herbaceous species: Chryso- pogon gryllus 2a, Hieracium hoppeanum 2a, Poa bulbosa 2a, Aira elegantissima 1, Anthoxanthum odoratum 1, Carex caryophyllea 1, Centaurea cuneifolia 1, Cerastium gracile 1, Chondrilla juncea 1, Crepis sancta 1, Daucus guttatus 1, Ero- dium botrys 1, Eryngium campestre 1, Hieracium bauhinii 1, Hypericum cerastoides 1, Hypericum olympicum 1, Hypo- chaeris glabra 1, Jasione heldreichii 1, Leontodon crispus 1, Moenchia mantica 1, Ornithopus compressus 1, Parentucellia latifolia 1, Petrorhagia illyrica 1, Potentilla recta 1, Romulea linaresii ssp. graeca 1, Rumex acetosella 1, Sanguisorba minor 1, Sherardia arvensis 1, Spergula pentandra 1, Teesdalia coro- nopifolia 1, Thymus atticus 1, Thymus longicaulis 1, Trifoli- um nigrescens ssp. petrisavii 1, Trifolium strictum 1, Trifolium tenuifolium 1, Tuberaria guttata 1, Anthemis arvensis +, Cir- sium vulgare +, Cnicus benedictus +, Dianthus corymbosus +, Geranium molle +, Lupinus angustifolius +, Moenchia erecta +, Orchis morio +, Scabiosa argentea +, Scilla autumnalis +, Scleranthus verticillatus +, Thesium dollineri; Cryptogams: Cladonia foliacea 1, Cladonia furcata agg. 1, Polytrichum pil- iferum 1, Racomitrium canescens 1, Cetraria aculeata agg. + 3. Galio flavescentis-Genistetum lydiae ass. nova hoc loco Holotypus hoc loco: Bulgaria, Sliven, NP “Sinite kamаni”; Relevé no. 41 (On–line Suppl. Tab 1); relevé collector: G. Kunev, 19.06.2016; plot size 16 m²; alt. 612 m a.s.l.; slope: 7°, SW; vegetation cover: 60%; latitude: 42.70712°, longitude: 26.34363° Shrub species: Genista lydia 3, Amelanchier ovalis +, Lem- botropis nigricans +; Herbaceous species: Plantago subulata 2a, Achillea coarctata 1, Anthemis cretica 1, Anthemis tinc- toria 1, Festuca dalmatica 1, Galium flavescens 1, Hieracium hoppeanum 1, Hypericum perforatum 1, Koeleria nitidula 1, Logfia arvensis 1, Phleum montanum 1, Rumex acetosella 1, Scabiosa triniifolia 1, Scleranthus perennis 1, Sempervivum erythraeum 1, Stachys angustifolia 1, Thymus callieri ssp. uru- movi 1, Verbascum humile 1, Veronica verna 1, Viola arvensis 1, Achillea millefolium +, Allium sphaerocephalon +, Alyssum alyssoides +, Avenula compressa +, Buglossoides arvensis +, Crucianella angustifolia +, Crupina vulgaris +, Cuscuta ap- proximata +, Hypericum montbretii +, Linaria genistifolia ssp. genistifolia +, Micropyrum tenellum +, Orlaya grandiflora +, Ornithogalum kochii +, Psilurus incurvus +, Scandix pectin- veneris ssp. macrorhyncha +, Viscaria vulgaris ssp. atropur- purea +; Cryptogams: Cladonia foliacea 1, Cladonia rangi- formis 1, Polytrichum piliferum 1