Phytocoenologia Vol. 46 (2016), Issue 4, 339-355 Published online August 2016 Special Issue Halophytic vegetation Research Paper Sand dune vegetation along the eastern Adriatic coast Urban Silc, Alfred Mullaj, Antun Alegro, Alban Ibraliu, Zora Dajic Stevanovic, Milica Lukovic & Danijela Stesevic Abstract Questions: What is the current state of knowledge on the distribution of psammophytic vegetation along the eastern Adriatic coast? Which are the main vegetation types and how do they vary from a floristic and ecological point of view? Study area: The eastern Adriatic coast of Croatia, Montenegro and Albania. Methods: We collected all available vegetation releves (a total of 191, published and unpublished) and historical references of sand dune vegetation from the eastern Adriatic coast. Classification and ordination were performed on the dataset and a syntaxonomical overview of the different plant communities along a seashore-inland zonation and their geographical distribution is presented. Results: The classification of the phytosociological data show eight floristically and ecologically well-defined clusters. The first group of embryonic foredunes comprises of the Cakilo-Xanthietum, Euphorbia paralias community and Eryngio-Sporoboletum, whilst the second group is found on the more stable dunes with Eupborbio paraliae-Agropyretum junceiformis, Medicagini marinae-Am-mopbiletum australis and Scabiosa argentea-Epbedra distacbya communities. The latter is newly described and is found only along the Albanian coast. The number of plant communities is lower than in surrounding countries, particularly on fixed dunes. The presence of the EU habitat type ' Crucianellion maritimae fixed beach dunes' (2210) is newly reported for Albania. Conclusions: Sand dunes are important habitats from a nature conservation point of view and they are endangered due to strong human impact. The protection of sand dune habitat types throughout the study area is urgent, since they are still in good condition in Albania, while sand dune plant communities are fragmented in Croatia and under strong human impact in Montenegro. Keywords: Albania; Ammopbiletea; Balkan Peninsula; Cakiletea; Croatia; Crucianellion maritimae; Montenegro; psammophytes Nomenclature: For vascular plant taxa Euro+Med (http://ww2.bgbm.org/EuroPlusMed/, accessed 2 April 2016), except for Calystegia soldanella, Cistus incanus, Cistus salvifolius, Citrullus lanatus, Clematis flammula, Cucumis melo, Fumana procumbens, Heliantbemum jonium, Heliantbemum nummularium, Knautia integri-folia, Lonicera implexa, Nigella arvensis, Oenothera biennis, Paliurus spina-cbristi, Polygonum maritimum, Radiola linoides, Rumex crispus, Rumex pulcber, Scabiosa argentea, Scabiosa atropurpurea, Tamarix dalmatica, and Viburnum tinus according to Flora Europaea (Tutin et al. 1964-1980,1993); for higher syntaxa Mucina et al. (2016), for associations see the overview in this paper. Abbreviations: DCA = Detrended Correspondence Analysis; PNV = potential natural vegetation. Submitted: 25 August 2015; first decision: 28 December 2015; accepted: 9 May 2016 Co-ordinating Editors: Erwin Bergmeier and Joop Schaminee Introduction Coastal dunes are complex ecosystems occupying zones of transition between terrestrial and marine ecosystems. This is a very dynamic environment, with distinct degrees of stabilisation depending on the topography, natural disturbance and distance from the sea, and it harbours a mosaic of habitats (van der Maarel 2003). Plant com- munities on sand dunes have specific ecological requirements determining their position along the gradient from the sea inland (Carboni et al. 2009). Various ecological factors influence the distribution of plants along this gradient: sand abrasion, salt spray, burial by sand, erosion, accretion, tide level, wave attack, wind blasting and dehydration (Maun 2009; Miller et al. 2010). Zonation is very stable and a regular sequence of plant communities along 'Corresponding author's address: ZRC SAZU, Institute of Biology, Novi trg 2, 1000 Ljubljana, Slovenia and BC Naklo, Strahinj 99, Naklo, Slovenia; urban@zrc-sazu.si. Complete addresses of all authors can be found at the bottom of the paper. ©2016Gebriider Borntraeger, 70176Stuttgart, Germany www.borntraeger-cramer.de DOI: 10.1127/phyto/2016/0079 0340-269X/2016/0079 $ 10.80 340 Urban Šilc et al. the gradient can be observed worldwide (Doing 1985; Attorre et al. 2013). Vegetation of beaches and mobile dunes is considered azonal (Doing 1985), while more inland, stable dunes are covered by plant communities more adapted to local conditions (Buffa et al. 2012). Sand dune vegetation occurs all along the Mediterranean coasts and several authors have provided syntaxo-nomical schemes of this vegetation type. In the central and eastern part of the Mediterranean, overviews have been made for Italy (Gehu et al. 1984; Gehu & Biondi 1996; Brullo et al. 2001; Biondi 2007; Pirone 2014), Greece (Sykora et al. 2003) and Turkey (Gehu et al. 1989), but not for the eastern Adriatic coast along the Balkan Peninsula. The history of vegetation research along the eastern Adriatic coast varies greatly in the last century due to the division of the region into two countries with different political systems. Research on psammophytic (sand dunes) vegetation started with Beck von Mannagetta (1901) and Morton (1915), who refer to the "Formation des Dünensandes". Horvatic (1934, 1939) was the first to study and provide a deeper insight into this vegetation type according to the Braun-Blanquet method, on the islands of Pag and Rab (Croatia), and he added new data in 1963 (Horvatic 1963). Extensive studies in Croatia were carried out by Trinajstic (1974, 1989a, 1995), Trinajstic & Jasprica (1998) and Alegro et al. (2003, 2004) Trinajstic & Jasprica (1998) and in Montenegro by Trinajstic (1989b) and Mijovic et al. (2006, 2012). Vegetation research in Albania applying the Braun-Blanquet method only started in the late 1980s. One of the pioneering works was on the coastal vegetation of Albania (Mullaj 1989), while studies of halophytic and psammophytic vegetation have only recently emerged (Imeri et al. 2010; Fanelli et al. 2015). These differences in the degree of vegetation research along the eastern Adriatic coast are evident from the distribution maps of the localities of sand dunes, on which the eastern Adriatic coast must still be considered a white spot in Europe (Gehu 1989; Trinajstic 1989a). The main aims of this paper are: (a) to present the current state of knowledge on the distribution of psammophytic vegetation along the eastern Adriatic coast, (b) to determine the main vegetation types by numerical analysis and (c) to describe these vegetation types from a flo-ristic and ecological point of view. The paper is also a contribution to knowledge of the vegetation of Albania, which is still under-explored. Study area The eastern Adriatic coast, or Adriatic coast of the Balkan Peninsula, is bound by the mouth of the Soca River in the northwest and the Strait of Otranto in the southeast (Fig. 1). Sand dunes, or even sandy sea shores, are very rare along this coast, in contrast to the Adriatic coasts of Italy and the Ionian or Aegean coasts of the Balkan Peninsula. Several factors are important: (a) the tectonics of Dalmatia with its relatively steep coast that influences the production of coarse clastic material, (b) water courses are oriented towards north-northwest, and (c) the active contact zone between the External Dinar-ides and the small Adriatic microplate is narrow and does not allow the formation of "stable" sedimentation space, where fine, sorted sediment accumulates (juračic et al. 2009). The north-eastern Adriatic coast therefore consists mainly of solid and steep limestone rocks; in the northern part, sandy beaches are scarce, small and with fragmented vegetation. Towards the south-east, the number of localities with sandy shores increases and well developed low sand dunes occur southwards, from the border between Montenegro and Albania extending to the Strait of Otranto (Trinajstic 1989a; Simeoni et al. 1997). According to Koppen's climate classification, the southern and central Adriatic sea have a dry summer Mediterranean climate (Csa), whereas the northern Adriatic has a humid subtropical climate (Cfa) (Lionello et al. 2012). The predominant winter wind is the bora and sirocco (jugo) which is observed during fall and spring. Methods We collected all available published and unpublished re-levés (191) on sand dunes that were recorded using the standard Central European method (Braun-Blanquet 1964) along the eastern Adriatic coasts (Appendix, Fig. 1). The 191 relevés were entered into the TURBOVEG (Hennekens & Schaminée 2001) database. Relevés were a posteriori georeferenced. Although plot areas varied in size from 4 to 200 m2, with the majority (74%) being between 10-50 m2 (first quartile 25, median 30 and third quartile 50 m2) all relevés were used for the analyses. Ecological indicator values (Pignatti 2005) were used for interpretation of the ecological gradients. Numerical classification of vegetation plots, on the basis of their species composition, was performed with the PC-ORD 5.0 program (McCune & Grace 2002). The original Braun-Blanquet cover values (sensu van der Maarel 1979) were transformed into percentages (mean values of percentage cover classes) and then square rooted. Euclidean distance (as distance measure) and the Ward method (for group linkage) were used. Clusters were identified by their diagnostic species, using the JUICE program (Tichý 2002). Diagnostic species were determined using the phi-coefficient as a fidelity measure (Chytrý et al. 2002). The size of all groups was standardized to equal size and Fisher's exact test (p < 0.001) was applied. Species with phi-coefficient values higher than 0.10 were considered to be diagnostic. After preliminary grouping of the diagnostic species as a Sand dune vegetation along the eastern Adriatic coast 341 result of using lower or higher threshold values, the mentioned phi value was subjectively selected. The selected phi-coefficient value threshold was chosen to be both low enough for a sufficient number of diagnostic species from an ecological and phytogeographical point of view, allowing the description of clusters, and high enough to avoid many generalist species, or species occurring in more than one cluster, from being considered diagnostic species. The relation between clusters and environmental indicator values was visualized using DCA ordination, in which the square-rooted cover percentages of species were used, and rare species were down-weighted. The original releves and mean unweighted ecological indicator values calculated for each releve were used in DCA. Ordination results are presented on spider plots, in which each releve is linked to the centroid of its cluster by a line and vectors representing indicator values were plotted on the ordination diagram. Analyses were performed by means of R (R Development Core Team 2012; http://cc. oulu.fi/~jarioksa/softhelp/vegan.html) using functions in the vegan package. 342 Urban Silc et al. Results The vegetation table (191 releves) includes 190 plant species. Among the species characteristic for the class Caki-letea maritimae (sensu Mucina 1997) most common were Xantbium orientale ssp. italicum and Cakile maritima, while among the common species of the Ammopbiletea were Elytrigia juncea, Ecbinopbora spinosa, Eryngium maritimum, Euphorbia paralias and Medicago marina. There is an evident increase of some psammophytic species towards the south of studied area: Glaucium flavum, Mattbiola tricuspidata, Polygonum maritimum, Am-mopbila arenaria, Ephedra distachya, Achillea maritima and Sporobolus pungens. Rare psammophytes in the studied area were Ephedra distachya, Stachys maritima, Helichrysum italicum and Ononis variegata. Classification The results of cluster analysis are presented in a dendrogram (Fig. 2) and a synoptic table (Table 1). Diagnostic species are ordered according to the decreasing phi value. Two main groups were distinguished in the dendrogram and the groupings of the communities reflect their syn-taxonomic classification and site conditions. The first group (clusters 1-3) includes plant communities on embryonic foredunes and driftlines, while the second group comprises communities on more stable dunes (white and grey dunes) (clusters 4-8). The classification produced a dendrogram with 8 clusters that were ecologically most meaningful and also in accordance with traditional vegetation classification. The geographical distribution of the clusters is presented in Fig. 3. Cluster 1 Cakilo-Xanthietum strumarii Diagnostic species: Cakile maritima, Xanthium orientale ssp. italicum, Salsola kali aggr. This halo-therophytic plant community is first in the zonation of vegetation from the sea towards the inland and occurs on nutrient rich sandy beaches. It occurs in Croatia, Montenegro and Albania. Cluster 2 Euphorbia paralias community Diagnostic species: Euphorbia paralias, Tamarix dalmat-ica, Crepis foetida, Cistus incanus, Medicago orbicularis, Scabiosa atropurpurea, Scirpoides holoschoenus, Cistus salvifolius, Alkanna tinctoria. A special plant community dominated by Euphorbia paralias that develops on embryonic shifting dunes and was classified as a separate cluster. It occurs only in Albania and can spread over large areas, but it has not been reported from other countries in the literature. Cluster 3 Eryngio-Sporoboletum virginici Diagnostic species: Sporobolus pungens, Plantago cor- onopus, Limbarda crithmoides,]uncus acutus. This community occurs in the first part of dune systems on lower sites that are regularly inundated by the sea, or unstable dunes under the influence of salt spray. It is the most halophytic plant community of the sand dune vegetation. The association is distributed along the whole Albanian coast, with one isolated location in Croatia (island of Mljet). Clusters 4, 5 and 6 represent the broad transitional association Agropyretum mediterraneum (syn. of Euphor-bio paraliae-Agropyretum junceiformis), characterised by the presence of Ammopbiletea species (Echinophora spinosa, Elytrigia juncea, Cyperus capitatus), and it is distributed over the whole researched area. Cluster 4 Euphorbio paraliae-Agropyretum junceiformis wet variant Diagnostic species: Xanthium orientale ssp. italicum, Schoenus nigricans, ]uncus maritimus. 57 23 28 16 16 25 21 S SIS Fig. 2. Simplified dendrogram of releves of vegetation on sand dunes along the eastern Adriatic coast. The number above the dendrogram branches indicates the number of releves. 1 - Cakilo-Xanthietum strumarii, 2 - Euphorbia paralias community, 3 - Eryngio-Sporoboletum virginici, 4 - Euphorbio paraliae-Agropyretum junceiformis wet variant, 5 - Euphorbio paraliae-Agropyretum junceiformis otanthetosum mahtimi, 6 - Euphorbio paraliae-Agropyretum junceiformis typicum, 7 - Scabiosa argen-tea-Ephedra distachya comm., 8 - Medicagini mahnae-Ammophiletum australis. Sand dune vegetation along the eastern Adriatic coast 343 Table 1. Synoptic table of psammophytic vegetation along the eastern Adriatic coast. The first number in the column is frequency in percentage and the second fidelity (phi value multiplied by 100). Among other species only those occurring in more than 8 releves are presented. Cluster 1 2 3 4 5 6 7 8 No. of releves 57 28 16 23 25 21 5 16 Diagnostic species Salsola kali aggr. 58 11.1 21 6 13 8 38 25 Cakile maritima 74 16.9 46 38 9 20 38 Xanthium Orientale ssp. italicum 81 13.8 96 6.1 50 96 15.2 71 40 6 Euphorbia paralias 35 96 27.4 31 65 12 67 62 Tamarix dalmatica 5 32 19.1 Crepis foetida 2 54 18.6 4 8 10 12 Scirpoides holoschoenus 50 11.7 12 5 60 19 Alkanna tinctoria 43 10.2 24 40 25 Medicago orbicularis 29 14.8 12 Cistus incanus 29 17.6 4 Scabiosa atropurpurea 14 13.3 Cistus salvifolius 29 10.7 28 14.3 Sporobolus pungens 19 61 100 46.9 13 48 10 12 Plantago coronopus 7 38 19.7 Limbarda crithmoides 26 14 62 14.9 26 8 14 Juncus acutus 25 11.7 20 Juncus maritimus 2 25 26 11.8 Schoenus nigricans 12 30 11.9 Calicotome villosa 20 20.9 Achillea maritima 4 36 20.8 Matthiola tricuspidata 11 36 18.8 Petrorhagia saxifraga 2 36 17.8 5 Pinus halepensis 2 g 24 16.4 10 Dorycnium hirsutum 20 14.8 Cladanthus mixtus 16 14.1 Brachypodium retusum 16 13.8 Trifolium campestre 16 13.6 Cyperus capitatus 18 29 38 72 10.8 90 14.7 100 50 Echinophora spinosa 51 86 62 87 88 100 16.2 81 Medicago minima 12 33 16.1 Ephedra distachya 100 79.2 Scabiosa argentea 100 37.1 Ononis variegata 80 32.6 Plantago crassifolia 60 27.4 Echium plantagineum 60 27.4 Silene conica 11 4 80 26.9 6 Hordeum murinum 18 60 24.2 Teucrium polium 32 6 6 16 80 21.7 Pseudorlaya pumila 2 21 22 8 43 100 18.3 50 Oenothera biennis 2 10 60 14.8 19 Ammophila arenaria 46 6 35 36 24 20 100 57.3 Silene colorata 31 19.7 Anchusa undulata 25 17.6 Maresia nana 18 38 17.1 Matthiola sinuata g 14 38 16.4 Vulpia fasciculata 39 4.2 6 4 5 60 75 16.4 Glaucium flavum 4 25 16.2 Hordeum marinum 19 15.3 Ambrosia maritima 10 25 14.7 Cakiletea maritimae Atriplex prostrata 344 Urban Silc et al. Table 1. cont. Cluster 1 2 3 4 5 6 7 8 No. of releves 57 28 16 23 25 21 5 16 Euphorbia peplis 33 50 31 4 8 33 Ammophiletea Elytrigia juncea 53 75 44 100 100 100 100 81 Eryngium maritimum 53 71 75 65 64 86 80 88 Calystegia soldanella 7 57 26 48 62 56 Lagurus ovatus 7 21 12 35 44 33 80 31 Medicago marina 11 57 6 57 24 81 8.5 100 75 Pancratium maritimum 11 21 6 39 36 90 9.2 80 69 Polygonum maritimum 26 43 12 13 10 60 25 Scolymus hispanicus 11 4 12 6 Others Dittrichia viscosa 5 50 7.7 19 24 5 40 50 Cynodon dactylon 14 43 6 20 19 20 31 Anisantha tectorum 14 22 12 20 31 Crithmum maritimum 14 24 5 Parapholis incurva 4 11 12 17 16 Cuscuta sp. 14 11 6 4 4 Phragmites australis 12 7 12 4 4 5 Catapodium rigidum 4 20 5 20 25 Vulpia ciiiata 14 22 12 Arundo donax 11 11 5 Reichardia picroides 2 7 6 17 12 Imperata cylindrica 4 16 14 20 Tribulus terrestris 2 14 20 25 Xanthium spinosum 4 12 25 Medicago littoralis 14 4 25 Aegilops neglecta 11 22 9.6 Linum strictum 16 19 Species occurring in less than 7 releves (first figure is cluster number, second is frequency in percentage): Aethionema saxatile 5:8; Agave amehcana 4:9; Amaranthus albus 8:6; Amaranthus blitum 2:4; Anagallis arvensis 2:11, 4:4; Anchusa azurea 8:13; Anisantha madritensis 6:5; Anisantha rigida 4:4; Anthemis chia 8:13; Anthemis cotula 8:6; Arenaria leptoclados 8:6; Asparagus acutifolius 4:4; Asparagus maritimus 8:6; Atriplex tatarica agg. 1:4; Avena barbata 6:5; Avena fa-tua 2:4; Avena sterilis 5:8; Bituminaria bituminosa 4:4, 6:10; Blackstonia perfoliata 2:7, 4:9; Briza maxima 5:8; Bromus squarrosus 8:6; Catapodium marinum 1:4. 4:4; Chondrilla juncea 2:4, 6:19, 7:20, 8:6; Cichorium intybus 2:4, 7:20; Citrullus lanatus 6:14; Clematis flammula 7:20; Colutea arborescens 5:4; Convolvulus sp. 1:9, 6:5; Corynephorus articulatus 5:8; Crepis setosa 5:4; Crepis sp. 6:5; Cucumis melo 6:5; Cynanchum acutum 2:5, 3:4; Dasypyrum villosum 7:20, 8:6; Daucus carota ssp. major 5:8; Daucus guttatus 8:13; Daucus sp. 4:4; Diplotaxis tenuifolia 4:9; Dorycnium pentaphyllum ssp. herbaceum 1:2; Echium italicum 5:12; Elytrigia atherica 2:4, 3:13, 8:25; Erigeron canadensis 4:4, 5:4, 5, 8:19; Erodium cicutarium 2:4, 5:8; Eryngium campestre 2:4; Eryngium creticum 5:4; Euphorbia peplus 1:2; Fumana procumbens 2:14, 3:6; Fumana sp. 5:4; Halimione portulacoides 1:2; Hedypnois rhagadioloides 1:2, 4:4; Helianthemum nummularium 5:12; Heiichrysum italicum 5:8; Heliotropium curassavicum 8:13; Helminthotheca echioides 2:11; Hieracium sp. 5:8; Hordeum murinum ssp. leporinum 5:12; Hy-parrhenia hirta 5:8; Hypochaeris radicata 2:7, 5:8, 6:10; Juncus bufonius 1:2; Juniperus oxycedrus ssp. macrocarpa 2:11, 6:5; Juniperus phoenicea 5:8; Knautia integrifolia 4:4; Lactuca viminea 8:6; Leontodon sp. 4:4; Limonium cancellatum 1:2; Lolium multiflorum 1:2, 2:4; Lonicera implexa 5:4; Medicago lupulina 8:6; Me-dicago sp. 6:5; Melilotus albus 5:12; Melilotus officinalis 1:2; Ononis reclinata 5:4; Ononis sp. 4:4, 5:8; Orobanche sp. 5:4; Paliurus spina-christi 1:2; Parapholis fi-liformis 3:13; Phillyrea angustifolia 1:2, 5:8; Plantago afra 5:8; Plantago cynops 1:4; Plantago lanceolata 8:6; Polypogon monspeliensis 1:2; Portulaca oleracea aggr. 7:20; Psilurus incurvus 2:4; Radiola linoides 5:8; Raphanus raphanistrum 2:7, 3:6, 8:6; Rapistrum rugosum 1:2; Rubus ulmifolius 7:20; Rumex crispus 4:4: Rumex pulcher 2:4; Salsola soda 1:2, 5:8, 6:10; Setaria viridis 6:5; Sherardia arvensis 7:20; S/7ene vulgaris 5:4; Sinapis alba 2:4; Solanum nigrum 8:13; Sonchus arvensis 8:6; Sonchus asper 2:4; Sonchus bulbosus 6:5; Spergula arvensis 7:20; Stec/iys maritima 1:2; Statice oleifolia 2:7; Suaeda maritima agg. 2:2, 3:6; Sym-phyotrichum squamatum 2:2, 4:6, 7:40; Taraxacum Sect. Taraxacum 6:10; TonV/s arvensis 5:8; Tragus racemosus 4:9, 7:20; Trifolium angustifolium 5:4, 7:20; 7n-fo//l/m arvense 5:8; Tripidium ravennae 2:14, 5:4; 1/efibascum sinuatum 2:11, 6:5; Viburnum tinus 5:4; V/c/a sp. 5:4; Vitex agnus-castus 1:2, 7:40. Sand dune vegetation along the eastern Adriatic coast Fig. 3. Distribution maps of clusters of sand dune vegetation. Black-cluster 4, yellow-cluster 5, red-cluster 6. 346 Urban Silc et al. A wet variant of the Eupborbio paraliae-Agropyretum junceiformis with dominant and constant species Elytri-gia juncea occurs in depressions in which water is retained due to various disturbances (natural or anthropogenic) and is syndynamically related to communities from hind dunes and associations of Juncetea maritimi (Juncetum maritimo-acuti, Eriantbo-Scboenetum nigri-cantis, Holoscboenetum romani). Differential species of this variant against the subassociations otantbetosum and typicum are Scboenus nigricans and Juncus maritimus, while Cyperus capitatus is absent in stands of the wet variant. Stands are found in Croatia, Montenegro and Albania. Cluster 5 Eupborbio paraliae-Agropyretum junceiformis otantbetosum maritimi Diagnostic species: Calicotome villosa, Achillea marit-ima, Mattbiola tricuspidata, Petrorbagia saxifraga, Pinus balepensis, Dorycnium birsutum, Cistus salvifolius, Clad-antbus mixtus, Bracbypodium retusum, Trifolium camp-estre, Cyperus capitatus. The community thrives on more internal embryonic sand dunes, already stabilized and is a transition towards an Ammopbila dominated community, with differential species Achillea maritima, Matbiola tricuspidata, and Petrorbagia saxifraga. The subassociation is distributed in Croatia and particularly in Albania, with many locations. Cluster 6 Eupborbio paraliae-Agropyretum junceiformis typicum Diagnostic species: Ecbinophora spinosa, Medicago minima, Cyperus capitatus. The typical subtype is found in Croatia, Montenegro and Albania but is limited in Albania to the Gosa area. The typical subassociation is characterized by common Ammophiletea species. Cluster 7 Scabiosa argentea-Ephedra distacbya community Diagnostic species: Ephedra distacbya, Scabiosa argentea, Ononis variegata, Plantago crassifolia, Ecbium planta-gineum, Silene conica, Hordeum murinum, Teucrium capitatum, Pseudorlaya pumila, Oenothera biennis. The cluster represents a rare plant community dominated by Ephedra distacbya, found in a limited range along the coast in Albania. It occupies small depressions behind dunes, where the sand is less permeable than in the Medicagini marinae-Ammopbiletum australis. The original location at Golem (Kavaja district) was destroyed by the construction of tourist facilities, but similar stands with dominant Ephedra distacbya were recently found by Mullaj at Rana e Hedhur (Lezha district). Cluster 8 Medicagini marinae-Ammopbiletum australis Diagnostic species: Ammopbila arenaria, Silene colorata, Anchusa undulata, Maresia nana, Vulpia fasciculata, Mattbiola sinuata, Glaucium flavum, Hordeum mari-num, Ambrosia maritima. The Medicagini marinae-Ammopbiletum arundi-naceae develops on higher parts of mobile dunes that have been stabilised by the rhizomatous geophyte Ammopbila arenaria. It is the final successional stage of herbaceous sand dune vegetation and communities that are located furthest inland are very often affected by tourism or erosion. The association is well developed in Albania, and locations are known in Montenegro (without releve material). To underpin the description and allocation of the new plant community of Scabiosa argentea and Ephedra distacbya, we made a comparison of Crucianellion mariti-mae and Syntricbio ruraliformis-Lomelosion argenteae associations from the southern Adriatic and Ionian seas along the coasts of Italy, Albania and Greece (Table 2). Communities belonging to these two alliances are found on fixed beach dunes. Diagnostic species of the Ammo-pbilion alliance prevail in all associations. The characteristic and dominant species is Ephedra distacbya, also the only diagnostic species of the Crucianellion alliance in this community, while others are lacking. On the other hand, Scabiosa argentea links this community to the Syn-tricbio-Lomelosion and more fixed (grey) dunes. Due to the absence of good character species and a small number of known locations so far, we decided to describe these stands as plant community without association rank and classify it within the Crucianellion maritimae. Gradient analysis Moisture, salinity and nutrients appear to be the most important ecological factors influencing the vegetation composition of sand dunes and their zonation (Fig. 4). They show a strong correlation to the main variation in the species composition of the vegetation. Soil reaction and temperature are less important. From wet, nutrient-rich, lower acidity and higher salinity to dryer, more nutrient-poor, higher acidity, the communities are ordered from left to right as follows: Cakilo-Xantbietum italici, Eryngio-Sporoboletum virginici, Eupborbio paraliae-Agropyretum junceiformis (Agropyretum medi-terraneum s.lat.) and embryonic shifting dunes with Euphorbia paralias to Medicagini marinae-Ammopbiletum australis and the community dominated by Ephedra distacbya. These communities represent a zonation from the sand deposition zone (clusters 1 and 3) to embryonic foredunes to finally more stabilised dunes (clusters 7 and 8). The first axis is positively correlated to the most important variables, while the second axis is related to temperature. On the left side of the ordination diagram are grouped vegetation types developed in the zone of deposit of sand (clusters 1 and 3), on the far left are vegeta- Sand dune vegetation along the eastern Adriatic coast 347 Table 2. Synoptic table of fixed dune vegetation on the Adriatic and Ionian coasts. Frequency in % and range of cover values are given for each species. 1. Scabiosa argentea-Ephedra distachya comm.; 2. Crucianelletum mahtimae Br.-BI. 1933; 3. Euphorbio-Sileneetum niceen-s/s Lavrentiades 1964; 4. Artemisio variabilis-Ephedretum Brullo, Giusso Del Galdo, Siracusa & Spampinato 2001; 5. Ephedro distachyae-Sileneetum subconicae Oberd. 1952; 6. Plantagini albicantis-Scabiosetum albae Brullo, Giusso Del Galdo, Siracusa & Spampinato 2001. Group No. 1 2 3 4 5 6 No. of releves 5 6 21 11 97 7 Scabiosa argentea-Ephedra distachya community Ass. 1, 6 Scabiosa argentea Ass. 1, 4, 5 Ephedra distachya Crucianellietum maritimae Crucianella maritima Euphorbio-Sileneetum nicaeensis Silene niceensis Euphorbia terracina Artemisio variabilis-Ephedretum distachyae Artemisia campestris ssp. variabilis Ephedro distachyae-Sileneetum subconicae Jasione heldreichii Silene congesta Silene dichotoma Centaurea cuneifolia Silene subconica Plantagini-Scabiosetum albae Plantago albicans Scabiosa atropurpurea 10G 10G +-1 3-4 81 95 69 62 32 78 4 +-2 2-3 Ammophilion/Ammophiletalia Ammophita arenaria 20 + 33 + 18 + 57 +-4 Echinophora spinosa 50 +-2 82 +-2 43 + Euphorbia paralias 16 +-1 Pancratium maritimum 80 + B4 +-3 15 +-4 100 +- Calystegia soldanella 18 +-3 29 +- Cyperus capitatus 100 +-1 100 +-1 45 +-2 65 +-3 29 + Eryngium maritimum 80 +-1 17 + 45 +-2 60 +-3 Medicago marina 100 +-1 50 +-1 19 +-1 27 +-1 69 +-3 Achillea maritima 36 +-3 Sporobolus pungens 17 + 10 + 16 +-2 100 +- Elytrigia juncea 100 + 83 +-2 10 + 27 1-2 73 r-4 57 +- Helichryso-Crucianelletalia Centaurea sphaerocephala B2 +-2 Helichrysum italicum 82 +-2 Artemisio-Koelerietalia albescentis Helianthemum jonium 86 +- Phleum arenarium 2 + Others Pseudorlaya pumila 100 +-1 B7 +-3 8 + 71 + Chondrilla juncea 20 + 29 + 36 + 31 +-2 Hypochaeris radicata 19 + 18 +-1 4 1 Ononis variegata 80 +-1 B2 +-2 29 + Lagurus ovatus 80 +-1 86 +-2 14 +-1 Vulpia fasciculata B0 + 100 +-3 20 +-1 Scirpoides holoschoenus B0 + 33 + 20 +-4 Cynodon dactylon 20 + 5 + 27 +-3 348 Urban Silc et al. Table 2. cont. Group No. 1 2 3 4 5 6 No. of releves 5 6 21 11 97 7 Dasypyrum villosum 20 + 14 + 12 +-2 Teucrium polium 80 + 3 + Hordeum murinum 60 +-1 4 +-1 Dittrichia viscosa 40 +-1 1 + Cichorium intybus 20 + 4 + Tribulus terrestris 20 + 3 + Rubus ulmifolius 20 + 1 + Spergula arvensis 20 + 2 + Polygonum maritimum 60 + 11 +-1 Xanthium strumarium 40 + 24 +-1 Alkanna tinctoha 40 + g +-1 Anisantha tectorum 20 + 46 +-1 Trifolium angustifolium 20 + 10 +-1 Hedypnois cretica 100 +-4 12 +-1 Nigella arvensis 29 +-1 33 +-2 Trifolium scabrum 29 1-4 6 +-1 Cutandia maritima 29 +-1 71 +-1 Scolymus hispanicus 24 +-1 18 + Verbascum sinuatum 19 +-1 45 +-1 Lotus creticus 100 1-3 43 + Matthiola sinuata 36 + 86 +-1 u -2-10 1 DCA1 Fig. 4. DCA graph of releves of sand dune vegetation from the eastern Adriatic littoral Environmental variables were projected onto the DCA diagram. Eigenvalues: 0.3511, 0.2440, 0.2203, 0.1776, gradient length: 3.6865, total inertia: 11.56. Numbers correspond to the centroids of the classified clusters. 1 - Cakilo-Xanthietum strumahi, 2 - Euphorbia paralias community, 3 - Eryngio-Sporoboletum virginici, 4 - Euphorbio paraliae-Agropyretum junceiformis wet variant, 5 - Euphorbio paraliae-Agro-pyretum junceiformis otanthetosum maritimi, 6 - Euphorbio paraliae-Agropyretum junceiformis typicum, 7 - Scabiosa argen-tea-Ephedra distachya comm., 8 - Medicagini marinae-Ammophiletum australis. Sand dune vegetation along the eastern Adriatic coast 349 Table 3. Correlation of the DCA axes with ecological factors. Significant codes: '***' 0.001, n. s.: non-significant. Squared correlation coefficient, P values based on 999 permutations. DCA1 DCA2 r2 Pr(>r) Significance Temperature 0,642 -0,766 0,078 0,001 *** Moisture -0,910 0,414 0,381 0,001 *** Soil Reaction -0,784 0,621 0,125 0,001 *** Nutrients -0,993 0,115 0,190 0,001 *** Salinity -0,990 -0,138 0,241 0,001 *** Light 0,990 0,139 0,004 0,687 tion types on more stabilised foredunes (clusters 7 and 8), while the centre of the graph is occupied by releves of embryonic foredunes. Discussion Syntaxonomy We compiled all available vegetation releves from the eastern Adriatic coasts that have been elaborated on, taking the study area as a whole. Previous studies have been restricted to particular beaches or countries (Mullaj 1989, Alegro et al. 2004, Mijovic et al. 2012). We demonstrated that the vegetation of sandy shores belongs to six plant communities (Table 1, Fig. 2): Cakilo-Xantkietum italici (cluster 1), Euphorbia paralias community (cluster 2), Eupborbio paraliae-Agropyretum junceiformis (clusters 4-6), Eryngio-Sporoboletum virginici (cluster 3), Medicagini marinae-Ammopbiletum australis (cluster 8) and Scabiosa argentea-Epbedra distacbya community (cluster 7). Syntaxonomic overview Cakiletea maritimae Tx. et Preising ex Br.-Bl. et Tx. 1952 Tbero-Atriplicetalia Pignatti 1953 Euphorbien peplidisľx. ex Oberd. 1952 Cakilo-Xantbietum strumarii (Beg. 1941) Pignatti 1958 Ammopbiletea Br.-Bl. et Tx. ex Westhoff et al. 1946 Ammopbiletalia Br.-Bl. et Tiixen ex Westhoff et al. 1946 Ammopbilion Br.-Bl. 1921 Eupborbio paraliae-Agropyretum junceiformis Tiixen in Br.-Bl. & Tiixen 1952 corr. Dari-mont, Duvigneaud & Lambinon 1962 Eryngio-Sporoboletum virginici Gehu et Uslu 1989 Euphorbia paralias community Medicagini marinae-Ammopbiletum australis Br.-Bl. 1921 corr. F. Prieto & T.E. Diaz 1991 Helicbryso-Crucianelletea maritimae Géhu et al. in Sissingh 1974 Crucianellietea maritimae Sissingh 1974 Crucianellion maritimae Rivas Goday et Rivas-Mart. 1958 Scabiosa argentea-Epbedra distacbya community The typical zonation pattern consists of the following more or less distinct zones: a) zone without vegetation, b) zone of organic/sand deposition, c) embryonic dunes, d) mobile (white) dunes and e) stabilised (grey) dunes (Bi-ondi 2007). Vegetation of cluster 1 is found in zone of deposition, embryonic dunes are colonised by vegetation classified in clusters 2 to 6. Vegetation of cluster 8 thrives on white dunes, while cluster 7 is found on grey dunes. Such complete zonation is not developed at all localities because sand dunes along the northeast Adriatic are narrower (15-20 m; Alegro et al. (2004)) than similar beaches in the southern Adriatic or in parts of Italy and Greece. The vegetation of sand dunes is better developed and richer in species in the south (Trinajstic 1989a). The Cakilo-Xantbietum italici (cluster 1) had not previously been reported in Croatia (Trinajstic 2008), as releves from the islands of Mljet (Velika and Mala Salunara) and Lopud were grouped with typical stands of this association from Montenegro and Albania in our analysis. Horvat et al. (1974) and Alegro et al. (2004) were of the opinion that the zonation of sand dunes on beaches in the central Adriatic were not typical, the transition is very gradual (orig. clinal) and conditions do not allow the development of the Cakiletea maritimae communities, or they are difficult to be recognised as a clearly separated belt. Nevertheless, we are of the opinion that this community is well developed in Croatia, due to the presence of the majority of characteristic species and clear aggre- 350 Urban Silc et al. gation in cluster analysis with releves from the southern Adriatic, where this association is typically developed. The Euphorbia paralias community is developed only in Albania, but a similar stand is shown on a photograph from Lopar beach (island of Rab, Croatia) in Morton (1915), which indicates that this plant community was probably distributed along the whole eastern Adriatic coast in the past. Lavrentiades (1964) mentions a variant of the Agropyretum mediterraneum with the dominant species Euphorbia paralias, in Greece. The Euphorbio paraliae-Agropyretum junceiformis is found along entire east Adriatic coast and shows high flo-ristic diversity with three subcommunities in the researched area: typicum, wet variant and otanthetosum. In the typical subassociation Echinophora spinosa also occurs in stands, with high fidelity and frequency, which is a diagnostic species of the ecbinopboretosum subassociation (Sykora et al. 2003) found on unstable dune formations adhering to rock, but Peloponnisos is the westernmost border of its distribution. The subassociation otanthetosum maritimae Gehu & Biondi 1984 is found on more internal embryonic dunes, which are more stable and continuous, similar to the Italian part of the Adriatic coast (Biondi 2007) and in Greece (Sykora et al. 2003). In the zonation (PNV) of east Adriatic sand dune systems, the Medicagini-Ammophiletum is usually followed by a stabilised dune occupied by woody vegetation (Pis-tacio lentisci-Juniperetum macrocarpae), while on the southern Ionian coast, the inner slopes of dunes are colonised by Crucianellion plant communities (Lavrentiades 1964). Only in two localities at the central Albanian coast, in depressions behind stabilised dunes, was the plant community originally classified as Ephedretum dis-tachyae (we are classifying it to the rank of community), found by Mullaj (1989). The community of Scabiosa ar-gentea and Ephedra distachya is syndynamically linked to the Ammopbilion, although it has ecological similarities to and characteristic species (Ephedra distachya, Vul-pia membranacea) of the Crucianellion alliance. Mullaj (1989) considered it vicariant to the Crucianelletum maritimae Br.-Bl. 1933 of the western Mediterranean and to the Tortulo-Scabiosetum Pignatti 1952 of the northern Adriatic (Venetian coast) and similar to the association Ephedro distachyae-Silenetum subconicae Oberdorfer 1952 from Greece. Although diagnostic species of the Ammopbilion alliance prevail in all associations (Table 2) and character species of Crucianellion are rare, we decided to classify stands from Albania as a new plant community based also on ecology and physiognomy (Pignatti et al. 1995; Winner 2006) and classify it within the Crucianellion. Our results (with new data from Albania) demonstrate that the diversity of plant communities on sand dunes of the eastern Adriatic is similar to that of the Italian Adriatic and to the Ionian coast in Greece. The diversity of communities and number of characteristic species decreases towards the north, since the coast becomes rocky in Croatia and Slovenia. Sand dune vegetation is common and widespread along the coasts of Europe, including the whole Mediterranean. Vegetation of sand dunes consists of five to nine associations in Italy (Corbetta et al. 1989; Stanisci et al. 2004; Biondi 2007; Pirone 2014), nine in Greece (Lavrentiades 1964; Sykora et al. 2003), five in Turkey (Gehu et al. 1989) and also five in the eastern Adriatic (Table 4). All plant communities are usually not present at the same locality due to natural and human disturbances that influence and change zonation. All five associations are not present in Croatia, where sand dunes are rare and most impacted by human activities (Korica & Lovric 1979) but, in Albania, in the south of the researched area, they are well developed (Fig. 5), but the community of Scabiosa and Ephedra distachya occurs only locally. Comparing plant communities and their zonation on the Italian Adriatic coast, it is obvious that some are rarer or missing on the eastern coast, e.g., communities of Crucianellion maritimae and Malcolmietalia Rivas Goday 1958. Crucianellion maritimae is typical vegetation on back dunes or interdunes of mobile dunes (Stanisci et al. 2004). In Greece, stands of Crucianellion are well developed on northern coasts and on the Peloponnisos (Sykora et al. 2003), whereas along the Adriatic coasts in Italy, the alliance is known only in Puglia (Pirone 2014). For Albania, we found a Crucianellion plant community that has not previously been published and this vegetation type is not present further north along the eastern Adriatic coast. Sand dunes on the north Adriatic coast (Veneto) are specific, with various species compositions and influenced by the continental climate. Similar communities on back dunes were classified into Syntrichio ruralis-Lomelosion argenteae Biondi, Sburlino & Theuril-lat in Sburlino, Buffa, Filesi, Gamper & Ghirelli 2013 are also found in the south on the Ionian coast of Puglia, but are not present along the eastern Adriatic coast. Vegetation of the Malcolmietalia, which comprises of ephemeral therophytic dune vegetation, is present only on the Italian coast (Pirone 2014) and has a western and southern Mediterranean distribution. On the other hand, other communities that are found on interdunal depressions (Juncetum maritimo-acuti Horvatic 1934, Erian-tho-Schoenetum nigricantis (Pignatti 1953) Gehu in Gehu, Scoppola, Caniglia, Marchiori et Gehu-Franck 1984, Holoschoenetum romani Tchou 1948) are developed only in Montenegro and Albania, where sand beaches are wider, while in Croatia they are found in other habitats, e.g., salinas. Sand dune vegetation along the eastern Adriatic coast 351 Quercetea ilicis Pinus halepensis-pinea comm. shore aphytic deposit embryonic dune white dune grey dune stable dune zone zone Fig. 5. Scheme of typical zonation of sand dune vegetation along east Adriatic coast from north to the south. Conservation Coastal areas and sandy beaches are very valuable from a nature conservation point of view, since they represent a mosaic of plant communities with high biodiversity. Many sand dune ecosystems are listed in habitats of European interest in Annex I of the Habitats Directive (Heslenfeld et al. 2004). The habitat types that appear on beach and mobile dunes in the researched area are presented in Table 4. In the future some of them should be translated into new types: 'Dunes along the Mediterranean shoreline with Ammopbila arenaria' (new code 2280) and 'Mediterranean embryonic dunes' (new code 2290) proposed by Feola et al. (2011). Several other habitat types which also occur along Mediterranean coasts are found on more stabilized dunes in Albania and partly in Montenegro: 'Coastal dunes with Juniperus ssp.' (2250), and 'Wooded dunes with Pinus pinea and/or Pinus pinaster (2270*). There is an urgent need for protection of these habitats on the eastern Adriatic coast. Some steps have already been taken in Croatia, where Velika and Mala Saplunara and Blace Bay (island Mljet) have been protected in the category of Protected Landscape Area since 1965 (Alegro et al. 2004) and in Montenegro, where part of Velika plaza has been protected as a Resort of Natural Landscape since 1968 (Official Gazette SRCG 30/ 68). Furthermore, the majority of the typical sandy coast species are Red listed in Croatia (Nikolic & Topic 2005). However, these categories do not provide sufficient active protection. Table 4. Plant communities on dune systems in various countries. Association names are original as stated by authors. Italics indicate syntaxa new to country and bold indicates syntaxa present in each country and confirmed in our study. NATURA 2000 habitat type CRO (Korica & Lovrič 1979, Trinajstič 2008) MNE (Blečič & Lakušič 1976, Mijovič et al. 2012) AL (Dring et al. 2002) GR (Sýkora et al. 2003) I (Pirone 2014) Zone without vegetation Deposition zone (drift line zone) Annual vegetation of drift lines 1210 Cakilo-Xan thietum strumarii Cakilo-Xan thietum strumarii Cakilo-Xanthietum strumarii Cakilo-Xanthietum strumarii Salsolo kali-Cakiletum maritimae Salsolo kali-Cakiletum maritimae Embryonic dune Sporoboletum Echinophoro-Elymetum Eryngio-Sporoboletum Eryngio-Sporoboletum virginici Sporoboietum farcti virginici Embryonic shifting dunes Echinophoro-Elymetum Euphorbio paraliae- Cypero mucronati-Agropyretum Echinophoro-Elymetum farcti 2110 farcti Agropyretum junceiformis juncei Echinophoro-Elymetum farcti othanthetosum White dune (yellow dune) Shifting dunes along the shoreline with Ammophila arenaria 2120 Ammophiletum australis (extinct) Ammophiletum australis (no relevés) Medicagini marinae-Ammophiletum australis Medicagini marinae-Ammophiletum Echinophoro-Ammophiletum australis Sileno-Vulpietum Grey dune Crucianellion maritimae fixed beach dunes 2210 Fixed coastal dunes with herbaceous vegetation (grey dunes) 2130* Scabiosa argentea-Ephedra distachya comm. (syn. Ephedretum distachyae Mullaj nom. inedit.) Euphorbio-Silenetum nicaeensis Ephedro distachyae-Silenetum subconicae Crucianellietum maritimae Crucianellietum maritimae Artemisio variabilis-Ephedretum distachyae Plantagini albicantis-Scabiose-tum é Tortulo-Scabiosetum albae (endemic N Adriatic) Sand dune vegetation along the eastern Adriatic coast 353 There is therefore an urgent need to raise the level of protection to Special Botanical Reserve. Sandy beaches in Albania that are still in good condition but are under strong human influence, mainly by increasing tourism, are especially worth protection, particularly sand dunes in Kune-Vain, Patok and Divjake. Author contribution The text was written by U.S., who also conducted the main part of the statistical analysis. US, Z.D.S, A.I., A.M., M.L. conducted field sampling of unpublished material in Albania. All authors endorsed the presentation and interpretation of the field work data and approved the final manuscript. 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(danijela.denist@gmail.com)8 1ZRC SAZU, Institute of Biology, Novi trg 2, 1000 Ljubljana, Slovenia 2 BC Naklo, Strahinj 99, Naklo, Slovenia 3 Faculty of Agriculture and Environment, Agricultural University of Tirana, Tirana, Albania 4 University of Zagreb, Faculty of Sciences, Division of Biology, Department of Botany, Croatia 5 University of Tirana, Faculty of Natural Sciences, Tirana, Albania 'University of Belgrade, Faculty of Agriculture, Department of Botany, Nemanjina 6, Belgrade-Zemun, Serbia 'Faculty of Tourism and Hotel Management, Department of Natural Science, University of Kragujevac, Serbia 8 Department of Biology, Faculty of Natural Sciences and Mathematics, University of Montenegro, Podgorica, Montenegro Appendix Relevé data source in database: Alegro et al. (2004): 33 rel, Croatia; Alegro et al. (2003): 4 rel, Croatia; Fanelli et al. (2015): 4 rel, Albania;, Imeri et al. (2010): 21 rel., Albania; Mijovic et al. (2006): 15 rel, Montenegro; Mijovic et al. (2012): 18 rel, Montenegro; Mullaj (1989): 51 rel., Albania; Silc et al. (unpublished): 25 rel., Albania; Trinajstic (1973): 2 rel., Croatia; Trinajstic (1989b): 7 rel., Montenegro; Trinajstic (1995): 5 rel., Croatia; Trinajstic & Jasprica (1998): 6 rel, Croatia. Unpublished relevés (25) were made by U.S., M.L. and Z.S.D. in June 2014 in Gosa area in Albania of following syntaxa: Cakilo-Xantkietum strumarii, Euphorbia paralias community, Eryngio-Sporoboletum Virginia, Eupborbio paraliae-Agropyretum junceiformis, and Medicagini marinae-Ammopbiletum australis. Plot size varies between 6 and 150 m2, with average of 44 m2.