O Hacquetia 15/1 • 2016, 49-65 DOI: 10.1515/hacq-2016-0002 DE GRUYTER OPEN A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) Nenad Jasprica1, *, Željko Škvorc2 & Milenko Milovic3 Keywords: Phytosociology syntaxonomy, Quercus coccifera, Quercetea ilicis, eastern Adriatic, Ionian coast. Ključne besede: fitosociologija, sintaksonomija, Quercus coccifera, Quercetea ilicis, vzhodni Jadran, ionska obala. Received: 16. 3. 2015 Revision received: 8. 9. 2015 Accepted: 15. 10. 2015 Abstract The results of a phytosociological investigation of the Quercus coccifera L. stands occurring along the Adriatic and Ionian coasts in south Albania are given. The surveyed stands are localized in the Thermo-Mediterranean and Meso-Mediterranean belts. On the basis of literature and unpublished data, Q. coccifera stands extend from the shoreline up to 680 m of altitude, within the Quercetea ilicis vegetation zone. According to numerical analysis, Q. coccifera stands in south Albania can be divided into two broad groups forming macchia of 3-4 m height and low shrublands [0.5-1.5(-2) m)] defined primarily by degree of human pressure and altitudes. In addition, relevés from Q. coccifera stands in Albania were differentiated from Croatian and Montenegrin the Fraxino orni-Quercetum cocciferae associations. Conversely, Q. coccifera stands in south Albania share several biological and ecological similarities with those of the eastern Adriatic coast: hemicryptophytes prevailed and the chorological spectrum highlights a clear dominance of the steno-Mediterraneans. With respect to indicator values, an important differentiation from the eastern Adriatic associations was shown only in higher light intensity of the stands in Albania. Izvleček Predstavili smo rezultate fitocenološke raziskave sestojev vrste Quercus coccifera L. ob jadranski in ionski obali v južni Albaniji. Preučevani sestoji se pojavljajo v termo-mediteranskem in mezo-mediteranskem pasu. Na osnovi literaturnih in neobjavljenih podatkov smo ugotovili, da so sestoji Q. coccifera razširjeni od obale do višine 680 m v vegetacijski coni Quercetea ilicis. Z numerično analizo smo sestoje razdelili v dve široki skupini: 3-4 m visoko makijo in nizko grmičevje [0,5-1,5(-2) m)], ki ju opredeljuje predvsem stopnja človekovega vpliva in nadmorska višina. Popisi sestojev z Q. coccifera iz Albanije se jasno razlikujejo od asociacije Fraxino orni-Quercetum cocciferae iz Hrvaške in Črne Gore. Nasprotno pa so sestoji Q. coccifera iz južne Albanije po bioloških in ekoloških lastnostih podobni tistim z vzhodne jadranske obale: prevladujejo hemikriptofiti, horološki spekter pa kaže močno prevlado stenomediteranskih vrst. Indikatorske vrednosti pa ločujejo sestoje iz Albanije od vzhodnojadranskih le po večji potrebi po svetlobi. 1 Institute for Marine and Coastal Research, University of Dubrovnik, P.O. Box 83, HR-20000 Dubrovnik, Croatia, *Corresponding author. E-mail: nenad.jasprica@unidu.hr 2 Faculty of Forestry, University of Zagreb, Svetošimunska 25, HR-10000 Zagreb, Croatia 3 "Antun Vrančic" Grammar School, Put Gimnazije 64, HR-22000 Šibenik, Croatia 67 O Hacquetia 15/1 • 2016, 49-65 N. Jasprica, Z. Skvorc & M. Milovic A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) Introduction Among the main forest species in Albania, oaks (Quercus) are represented by 12 taxa and their forests contribute to 32.7% of the surface area of Albania (Vangjeli & Habili 1995, Proko et al. 2005, Haska 2011). They are distributed over almost all the Albanian territory, but most of the forests occur in the north-east and south-eastern parts of Albania (Dida 2003). By contrast, Mediterranean shrubby formations were mainly found in the southern mountain region and mostly consisted of evergreen (e.g. Quercus ilex, Q. coccifera, etc.) or deciduous shrubs (Paliurus spina-christi, Ulmus foliacea, etc.) (Demiraj et al. 1996). Generally, on the eastern Adriatic coast, the area of Q. coccifera L. distribution extends from the northern Croatian island of Mali Losinj (44°35' N) to the south Albanian island of Sazani (40°29' N). It also occurs in the most south-western part of the Adriatic Basin in the Italian region of Puglia (Salento area), while its populations also continue on the Ionian coasts of both Italy (Basilicata region) and south Albania (districts of Vlore and Sarande) (Conti et al. 2005, Shuka 2010, Barina & Pifko 2011, Miho et al. 2013, etc.). The phytosociology of Q. coccifera-dominated vegetation formations has been studied across the Mediterranean region and many different associations and/or subassociations have been described (Tsiourlis et al. 2009, and references therein). The most recent and a very comprehensive description of Q. coccifera stands and their character-species on the eastern Adriatic, especially in Croatia, was given by Jasprica et al. (2015). On the eastern Adriatic, Q. coc-cifera forms a macchia (Fraxino orni-Quercion ilicis) within the Fraxino orni-Quercetum cocciferae pistacietosum lentisci in Croatia and Montenegro and also, though only sparsely, as shrubland within Fraxino orni-Quercetum cocciferae neri-etosum oleandri subassociation in south Croatia. In addition, Q. coccifera also appears as a macchia within the Erico arboreae-Arbutetum unedonis (Oleo sylvestris-Ceratonion si-liquae) association on the south Croatian island of Mljet. In Albania, stands with Q. coccifera have been previously associated with the Fraxino orni-Quercetum cocciferae association (Hoda & Mersinllari 1996, Dring et al. 2002), but it seems that this association, at least in part, is not developed in the area (Jasprica et al. 2015). Until now, very scarce data on the floristic composition and syntaxonomic characteristics of Q. coccifera stands from Albania have been available (Hoda & Mersinllari 1996). This study is focused on the known sites with Q. coccifera stands in south Albania. The objectives were: 1) to bring together the available published and recently collected data, and 2) to investigate the phytosociology and syntaxonomy of Q. coccifera-dominated formations in this area. The results will contribute to the scarce syn-taxonomic knowledge of these formations within the Quercetea ilicis vegetation zone in Albania. Study area The study area of Quercus coccifera-stands covers most of the south Adriatic coast, extending from the Strait of Otranto where it connects to the Ionian Sea (Figure 1). The investigated area stretches along the district of Vlore (AL: Rrethi i Vlores; 40° 21' N, 19° 37' E; surface area of 1,609 km2; population of 151,314 estimated 2010) in the direction NW-SE. The investigated area belongs to the Sazani and Ionian zones of External Albanides, and consists mostly of: i) Cretaceous fissured and karstified limestones, dolomitic limestones and dolostones (the Sazani-Karaburuni nature reserve, village of Dhermi); ii) Quaternary gravel, sand and poorly consolidated boulders covered by shallow clay (village of Dukati), and iii) Neogenic rocks, mainly clay-stones, silts and marlstones (town of Orikumi) (Polemio et al. 2008, Frasheri et al. 2009). In terms of soils, Euri-lithic Leptosol and Calcic Luvisol have developed in the investigated area (Zdruli 2005). Phytogeographically, the area lies within the Quercetea ilicis vegetation zone, where the climate is typically Mediterranean: mild and rainy winters, warm and dry summers, and an extended period of sunshine throughout the year. Average annual air temperature is 16.5 °C and precipitation averages 1,028 mm yr1 (Kabo 1990-1991, Nuri 1995, Frasheri et al. 2008). The highest daily average tem- Figure 1: Position of investigated localities in south Albania. Abbreviations: AL - Albania, MK - Republic of Macedonia, GR - Greece. Slika 1: Raziskovane lokacije v južni Albaniji. Okrajšave: AL - Albanija, MK - Republika Makedonija, GR - Grčija. 65 O Hacquetia 15/1 • 2016, 49-65 N. Jasprica, Z. Skvorc & M. Milovic A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) perature is 24.6 °C in August, and lowest falls below 9.2 °C in January. The absolute minimum recorded so far is -7 °C, but temperatures of zero or below occur only on 3 to 5 days per year. The highest temperature recorded during the past three decades is 39.5 °C. Eastern and south-eastern winds prevail in the area. The annual hours of sunshine are 2,734 (Nuri 1995). According to Koppen's climate classification, the coastal lowlands of Albania lie within the Csa climate zone (Koppen & Geiger 1954, Strafier 1998). Over the last decades the few remnants of natural vegetation, including Q. coccifera stands, in the Albanian karstic area have been seriously threatened by uncontrolled human activities (cf. Parise et al. 2004). Material and Methods Stands and relevé data In this study, 35 phytocoenological relevés were analysed. In July 2014, 24 phytocoenological relevés were made on the Adriatic side of south-west Albania: 1) near the town of Orikumi, 2) SW of the village of Dukati within the Sazani-Karaburuni nature reserve, and on the Ionian coast, 3) on the S-facing slopes of Llogara Pass (AL: Qafa e Llogorasë), and 4) SE of the village of Dhërmi. The sampling localities are shown in Figure 1 (see also details on the sites of relevés in the Appendix). In addition, nine relevés from Q. coccif-era shrublands in south Albania originating from the island of Sazani were used from Hoda and Mersinllari (1996). Relevés were collected using the Braun-Blanquet (1964) approach. Most of the plot-sizes were set at 100 m2, and every effort was made to achieve high ecological and physiognomic homogeneity within each plot (Moravec et al. 1994). The plots were located at various altitudes, exposures and inclinations. The system of characterizing species and the nomenclature of higher taxa were derived from Horvat et al. (1974), Rivas-Martinez et al. (2002) and Biondi et al. (2014). The nomenclature of plant species follows the Flora Europaea (Tutin et al. 1964-1980, 1993; Royal Botanic Garden Edinburgh 2014), except for Cistus creticus ssp. eriocephalus (marked with asterisks *) where the Med-Checklist was used (Greuter et al. 1984). Constant species were defined as those with a frequency > 30% in all the relevés. Dominant species were defined as those occurring in at least 10% of relevés of a vegetation unit with a cover value >25%. Biological form was verified in the field and denoted according to categories reported in Pignatti (1982), these being based on the classification of Raunkiaer (1934). Regarding chorological form, reference was also made to Jasprica et al. (2014, 2015), as well as to the monographs used for taxonomic nomenclature. The abbreviations of life- and chorological forms are given in Table 1, before the each species name. These abbreviations are denoted in Figure 3 and Table 2, respectively. Regarding taxonomy, both Q. coccifera and Q. calliprinos Webb were reported from Albania (cf. Paparisto et al. 1988, Mullaj et al. 2010, Ball 2011). In this study, we recognize one taxon that is referred to as Q coccifera and includes the possible presence of Q. calliprinos. In practice we accepted the taxonomic data in Euro+Med Plantbase (Euro+Med 2006-) and the results ofToumi and Lumaret (2010) who suggest that Q coccifera and Q calliprinos are, in fact, two morphotypes closely related genetically and constitute two components of the same species. Statistical analysis In order to obtain more complete information about Q. coccifera stands in south Albania, we compared our relevés with those from the eastern Adriatic coast (Jasprica et al. 2015). To identify vegetation types, relevés were classified by TWINSPAN (Hill 1979) using Juice 7.0 software (Tichy 2002). TWINSPAN pseudospecies cut levels for species abundances were set to 0—5—25 percentage scale units. Initially, six division levels were chosen. Later, different levels of division were accepted resulting in nine groups interpretable in terms of ecology. The resulting classification were projected onto an ordination diagram using non-metric multidimensional scaling (NMDS) performed on a matrix of Bray-Curtis dissimilarities. Ordination was calculated using the R program (R Development Core Team 2012) and its vegan package (Oksanen et al. 2012). For further interpretation of the ecological conditions of the studied vegetation types, unweighted average indicator values were used (Pignatti 2005) calculated in the JUICE 7.0. Average indicator values were presented with Box-Whiskers diagrams made in the STATISTICA 7.1 (STATSOFT inc. 2005). Results In this study, the Quercus coccifera stands in south Albania belong to the macchia or degraded scrub vegetation occurring in Mediterranean bioclimatic strata at 0-685 m a.s.l. (Table 1). Q. coccifera was also found at higher altitudes on the Llogara Pass (ca. 970 m a.s.l.) within the grassland vegetation not studied here. Generally, two groups of stands were recognized. The stands represent low [0.5-1.5(-2) m)] shrubby formations (the island of Sazani, Dukati and the S-facing slopes of Llogara Pass) or macchia of 3-4 m height (Orukumi, Dhermi). They generally occupy slopes of20-40° with vegetation cover from 30 to 100%. The habitats are exposed to disturbances. 65 O Hacquetia 15/1 • 2016, 49-65 N. Jasprica, Z. Skvorc & M. Milovic A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) ? □ Median □ 25%-75% I Min-Max Light T , Continentality . , Soil Reaction ,. , TemDerature Moisture Nutrients Figure 2: Pignatti indicator values in the Quercus coccifera stands in south Albania. Slika 2: Pignattijeve indikatorske vrednosti sestojev Quercus coccifera v južni Albaniji. The stands had from 8 to 30 plant taxa (average 17.4). Generally the highest number of taxa was recorded on the island of Sazani. Constant species were: Quercus coccifera, Pistacia lentiscus, Olea europaea var. sylvestris, Spar-tium junceum, Brachypodium retusum, Carlina corymbosa, Hyparrhenia hirta, Micromeria graeca, Phlomis fruticosa, Thymus capitatus, Erica manipuliflora, Teucrium polium, Scutellaria rubicunda and Scolymus hispanicus. Dominant species were: Quercus coccifera, Pistacia lentiscus, Phillyrea angustifolia, Phlomis fruticosa, Erica manipuliflora and Cistus creticus ssp. eriocephalus. The greatest number of characteristic species were found within the Festuco valesiacae-Brometea erecti (28), followed by the Quercetea ilicis and Thero-Brachypodietea ramosi (with 20 taxa in each class), Artemisietea vulgaris (13) and Cisto cretici-Micromerietea julianae (including Erico-Cistetea, 10 taxa). In addition to Q. coccifera, four other species displayed the greatest (>50%) presence: Pistacia lentiscus, Brachypodium retusum, Phlomis fruticosa and Thymus capitatus. Quercus coccifera stands in south Albania showed high Pignatti indicator values for light intensity and temperature (Figure 2). On the other hand, low moisture and nutrient values were indicated. With regard to continentality and substrate reaction, the stands had an intermediate position. The analysis of plant life forms showed that the stands were dominated by hemicryptophytes (35%) followed by phanerophytes (25%), while chamaephytes and thero-phytes contributed equally (17%) (Figure 3). Mediterranean floral elements (61%), mostly circum-Medi-terranean plants (38.21%), followed by a considerable proportion of Eurasian plants (9.76%), dominated in the stands (Table 2). 40 n 30- 20- 100. I Chamaephyta Geophyta Hemicryptophyta Phanerophyta Terophyta (Ch) (G) (H) (P) (T) Figure 3: Life-form spectra (in percentage) in the Quercus coccifera stands in south Albania. Slika 3: Spekter življenskih oblik (v odstotkih) v sestojih Quercus coccifera v južni Albaniji. Based on TWINSPAN, there was a clear difference in the structure between Croatian and Montenegrin associations (the mostly common subassociation Fraxino orni-Quercetum cocciferae pistacietosum lentisci) and stands in south Albania (Figure 4). In addition, stands in Albania also differed among the investigated localities. In fact, relevés from the island of Sazani were separated into two groups (4 and 5). Group 5 (corresponding to the relevés 19—21, Table 1) was dominated by QQuercus ilex and Phillyrea angustifolia while QQuercus coccifera had low abundances. Relevés from Dukati and Llogara (groups 7 and 8), and only partly those from the island of Sazani (group 4), were closely related (Figure 4). Relevés from Dukati and Llogara were characterized by the absence of almost all taxa (except Pistacia lentiscus and Juniperus oxycedrus ssp. macrocarpa) from the QQuercetea Table 2: Floral elements in the Quercus coccifera stands in south Albania. Tabela 2: Florni elementi vegetacije s Quercus coccifera v južni Albaniji. iFloral elements No. % Circum-Mediterranean (CME) 47 38.21 East Mediterranean (EME) 12 9.76 Northeast Mediterranean (NEME) 3 2.44 Southeast Mediterranean (SEME) 12 9.76 South Mediterranean (SME) 1 0.81 West Mediterranean (WME) 1 0.81 Illyrian-Adriatic (IL-ADR) 7 5.69 Illyrian-Apennine (IL-APEN) 2 1.63 Illyrian-Balkanic (IL-BALK) 1 0.81 Mediterranean-Atlantic (MEAT) 6 4.88 Mediterranean-Pontic (MEPO) 2 1.63 European-Mediterranean (EURME) 1 0.81 Euroasian (EURAS) 12 9.76 Southeast European (SEE) 3 2.44 South European-Pontic (SEPO) 5 4.07 Cosmopolitan (WS) 8 6.50 65 O Hacquetia 15/1 • 2016, 49-65 N. Jasprica, Z. Skvorc & M. Milovic A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) -1.5 -1.0 -0.5 0.0 0.5 1.0 1.5 NMDS1 Figure 4: NMDS ordination diagram with projected cluster membership of the relevés. Group abbreviations: 1. Fraxino orni-Quercetum cocciferaepistacietosum lentisci (Croatia, Montenegro); 2. Fraxino orni-Quercetum cocciferae nerietosum oleandri (Croatia); 3. Erico arboreae-Arbutetum unedonis (Croatia); Stands in south Albania: Groups 4-5. Island of Sazani (rel. 14-22, Tab. 1); 6. Orikumi (rel. 8-13, Tab. 1); 7. Dukati (rel. 23-27, Tab. 1); 8. Llogara (rel. 28-35, Tab. 1); 9. Dhërmi (rel. 1-7, Tab. 1). Slika 4: Ordinacijski diagram NMDS s prikazano pripadnostjo popisov posameznim klastrom. Okrajšave skupin: 1. Fraxino orni-Quercetum cocciferae pistacietosum lentisci (Hrvaška, Črna Gora); 2. Fraxino orni-Quercetum cocciferae nerietosum oleandri (Hrvaška); 3. Erico arboreae-Arbutetum unedonis (Hrvaška); sestoji iz Albanije: Skupini 4-5. otok Sazani (rel. 14-22, Tab. 1); 6. Orikumi (rel. 8-13, Tab. 1); 7. Dukati (rel. 23-27, Tab. 1); 8. Llogara (rel. 28-35, Tab. 1); 9. Dhërmi (rel. 1-7, Tab. 1). ilicis class, and the presence of some common taxa from the Thero-Brachypodietea ramosi (e.g. Brachypodium retu-sum), Festuco valesiacae-Brometea erecti ( Teucrium polium, Bromus erectus ssp. condensatus, etc.), and Cisto cretici-Micromerietea julianae (Phlomis fruticosa) classes. Relevés from Orikumi (group 6) and Dhërmi (group 9) showed some similarities due to the presence of the Quercetea ilicis taxa, particularly of Spartium junceum and Olea europaea var. sylvestris. Among all localities, relevés from Dhërmi were floristically the most closely related to Croatian and Montenegrin associations (group 1). Indeed, the similar physiognomic aspects of the stands were also evident. Discussion This study revealed the presence of sclerophyllous Quercus coccifera stands forming macchia or shrublands on the Adriatic and Ionian coasts of south Albania. Our results suggest that Q. coccifera stands in south Albania appear within the Thermo- and Meso-Mediterranean belts and extend from the shoreline up to 680 m of altitude within the Quercetea ilicis vegetation zone, the natural growing zone of Q. coccifera. In northern Greece, Q. coccifera formations were found from Meso-Mediterranean levels up to montane levels, and comprise at least two classes being syntaxonomically involved, i.e., Quercetea ilicis and Quercetea pubescentis (Mavrommatis 1980, Raus 1980, 1982, Bergmeier 1990, Bergmeier & Dimopoulos 2008). It has also been documented that Q. coccifera penetrates deep in the continental parts of Greece that are influenced by sub-Mediterranean climate, and can be found even within the vegetation of the Rhamno catharticae-Prunetea spinosae class (Tsiaoussi 1996). Further, in south Albania, Q. coccifera stands have already been found from 400-600 m a.s.l. in the nearby Llogara National Park -an area not included in our survey (Proko 1997). The numerical analysis suggests that the relevés collected in south Albania, except partly for those made in Dhërmi, were distinct from the Croatian and Montenegrin Fraxino orni-Quercetum cocciferae associations. However, the floristic composition of the Q. coccifera macchia in south Albania does not include the characteristic shrub (Fraxinus ornus), and some climbing species of Fraxino orni-Quercion ilicis, such as Rosa sempervirens, Lonicera implexa, L. etrusca, and Erica arborea, Viburnum tinus, Arbutus unedo, etc., commonly presented along the eastern Adriatic coast and islands (Jasprica et al. 2015). In addition, Q. coccifera stands in south Albania were mostly depleted of Quercus ilex — the taxon frequently found in maquis communities and sclerophyllous woodlands in Central Albania (Kalajnxhiu et al. 2012). This led us to suggest that i) Q. coccifera stands should not be reffered to the Fraxino orni-Quercetum cocciferae association, as has been previously reported by Albanian authors (cf. Hoda & Mersinllari 1996, Ruci et al. 2001, Dring et al. 2002, Anonymous 2004), or ii) the stands can be even considered as degraded forms of the Fraxino orni-Quercetum cocciferae association. In our opinion, Q. coccifera stands in south Albania show some floristic and ecological similarities with the Phillyreo latifoliae-Quercetum calliprini association, described within the Thermo-Mediterrane-an bioclimatic belt on the Ionian island of Cephalonia (Bolos et al. 1996). However, Greek association included the high presence of Cyclamen hederifolium, Arisarum vulagre and some others, and the majority of the typical Mediterranean evergreen shrubs, mainly the mesophilous species (Arbutus unedo, A. andrachne, etc.), not observed in our study probably as a result of combined types of human impact (cutting, grazing, wildfires). In either case, the Q. coccifera stands of south Albania, as in Greece, can be considered as degraded forms of the primary forests with Q. coccifera accompanied by Q. ilex, which currently 65 O Hacquetia 15/1 • 2016, 49-65 N. Jasprica, Z. Skvorc & M. Milovic A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) has only limited local distribution (Anonymous 2004, Tsiourlis et al. 2009). However, the dynamic potential of these stands, especially the temporal process of progressive succession and the corresponding climax forest community, are to a great extent site-dependent and are still not understood in detail (Proko 2008). In comparison with the synthetic table made by Jasprica et al. (2015) for the Mediterranean zone (Quercetea ilicis) of the eastern Adriatic, many differences in the floristic composition of our stands can be found. Quercus coccifera stands in Albania are constituted basically by some characteristic species of rocky scrubland plant communties (Phlomis fruticosa) and dry grasslands (Brachypodium re-tusum), which occurred at higher percentages, while some of the main sclerophyllous species such as Phillyrea lati-folia, Asparagus acutifolius, Juniperus oxycedrus ssp. oxyce-drus, etc., appeared at low frequencies. Among Quercetea ilicis taxa, only Pistacia lentiscus occurred at higher percentages. This is only partly consistent with the findings of Tsiourlis et al. (2009) who found that Phillyrea latifolia and Pistacia lentiscus are among the most common species in the Q. coccifera shrublands of Greece. In our study the presence of some characteristic species of the ruderal and weed vegetation (Artemisietea vulgaris, Stellarietea mediae) suggest that some of the localities (e.g. Orikumi) were linked to the close proximity of settlements, an environment highly affected by man. Further and in general, from the standpoint of the floristic and ecological characterization, there were no differences in the average number of taxa per relevé, the life-form and chorological spectra between the eastern Adriatic Fraxino orni-Quercetum cocciferae (the mostly common subassociation pistacietosum lentisci) association and the stands in south Albania. Namely, hemicrypto-phytes prevailed and the chorological spectrum highlights a clear dominance of the steno-Mediterraneans. With respect to species' ecological requirements, an important differentiation from the eastern Adriatic associations was shown only in the light intensity. This can be explained by a greater herb layer coverage and a higher species richness of herbs in the stands in south Albania. The results of the current study suggest that the Q. coc-cifera stands in south Albania can be divided into two broad groups: i) macchia of 3—4 m height (Orikumi, Dhërmi), and ii) mostly low shrublands in the other investigated south Albanian localities. First, this was defined by the degree of human pressure. In fact, south Albanian Q. coccifera shrublands have suffered dramatic changes under anthropogenic activities (deforestation and overgrazing) that have affected both their physiognomy as well as their floristic composition (Proko 2008). Second, the importance of altitude in determining the distribu- 65 tion and compostion of the communities could not be ignored. Namely, some relevés (no. 23—35) were made at higher altitudes (350—685 m a.s.l.), and they were characterized by absence of the (Quercetea ilics taxa and presence many taxa of the Festuco valesiacae-Brometea erecti class. In this case, Q. coccifera shrublands can mediate with the Carpinion orientalis communities (e.g. Phillyreo latifoliae-Carpinetum orientalis), which has already been documented in Greece (Bergmeier & Dimopoulos 2008). The highly anthropogenic distribution and composition of ( . coccifera shrublands within the study area, accompanied by differences in taxonomical concepts (cf. Karaer 2010), does not permit the provision of a precise syntaxo-nomical reference in terms of association. According to the results of Tsiourlis et al. (2009), human-induced degradation can cause differentiations in Q coccifera shrublands allowing some elements of the neighbouring flora to participate in, and often alter, both the physiognomy and the floristic composition of the shrublands. Many of the Q. coccifera dominated shrublands in the Mediterranean were classified under the Quercetum cocciferae association in its broader sense (e.g. Bolos 1959, Oliver 2004). This classification is based mainly on physiognomic criteria of the stands and their important flo-ristic variations were commonly avoided. But even if we accept this concept, Q. coccifera stands in south Albania, at least in part, had some similarity with the Quercetum cocciferae brachypodietosum identified by Wraber (1952) in southern France and by Curcó (1991) in Catalonia (Spain). Some of the localities with Q. coccifera stands in south Albania have not been covered by this study (e.g. the western slopes of the Karaburuni peninsula), and clearly further studies of the communities are required to increase the accuracy of predictions. In summary, we believe that the most significant result of this paper lies in the information and quantitative data it provides about Q. coccifera vegetation in this part of the NE Mediterranean. Nevertheless, given the relatively small area studied, the results should be read and analysed in the context of the anthropogenic influences that are occurring as a generalized phenomenon throughout the Mediterranean basin. Acknowledgements The authors thank Dr Rade Garic for preparation of the map of localities. Thanks also are extended to Jasmin Sali-jevski (NL) for technical support and to Steve Latham (UK) for improving the English. O Hacquetia 15/1 • 2016, 49-65 N. Jasprica, Z. Skvorc & M. Milovic A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) Appendix Syntaxonomic units mentioned in the text and Table 1 (in alphabetical order): Ammophiletea Braun-Blanquet & Tüxen ex Westhoff, Dijk & Passchier 1946 Artemisieta vulgaris Lohmeyer, Preising & Tüxen ex von Rochow 1951 Carpinion orientalis Horvat 1958 Asplenietea trichomanis (Braun-Blanquet in H.Meier & Braun-Blanquet 1934) Oberdorfer 1977 Cisto cretici-Micromerietea julianae Oberdorfer ex Horvatic 1958 Drypidetea spinosae Quézel 1964 Erico arboreae-Arbutetum unedonis Allier & Lacoste 1980 Erico-Cistetea Trinajstic 1985 Festuco valesiacae-Brometea erecti Braun-Blanquet & Tüxen ex Braun-Blanquet 1949 Fraxino orni-Quercetum cocciferae (Horvatic 1958) Trinajstic 1985 Fraxino orni-Quercetum cocciferae (Horvatic 1958) Trinajstic 1985 nerietosum oleandri Jasprica & Skvorc 2015 Fraxino orni-Quercetum cocciferae (Horvatic 1958) Trinajstic 1985 pistacietosum lentisci Jasprica & Skvorc 2015 Fraxino orni-Quercion ilicis Biondi, Casavecchia & Gigante ex Biondi, Casavecchia & Gigante in Biondi, Allegrezza, Casavecchia, Galdenzi, Gigante & Pesaresi 2013 Molinio-Arrhenatheretea Tüxen 1937 Nerio oleandri-Tamaricetea africanae Braun-Blanquet & O. Bolos 1958 Oleo sylvestris-Ceratonion siliquae Braun-Blanquet ex Guinochet & Drouineau 1944 Paliuretea Trinjastic 1978 Phillyreo latifoliae-Carpinetum orientalis Bergmeier 2008 Phillyreo latifoliae-Quercetum calliprini Knapp 1965 em. Barbéro & Quézel 1976 Quercetea ilicis Braun-Blanquet 1947 Querceteapubescentis Doing-Kraft ex Scamoni & Passarage 1959 Quercetum cocciferae Braun-Blanquet 1924 Quercetum cocciferae Braun-Blanquet 1924 brachypodietosum Braun-Blanquet 1935 Querco cocciferae-Pistacietum lentisci Braun-Blanquet, Font Quer, G. Braun-Blanquet, Frey, Jansen, & Moor 1935 nom. mut. propos. (art. 45) (addenda) [Querco cocciferae-Lentiscetum Braun-Blanquet, Font Quer, G. Braun-Blanquet, Frey, Jansen, & Moor 1935] Querco roboris-Fagetea sylvaticae Braun-Blanquet & Vlieger in Vlieger 1937 Rhamno catharticae-Prunetea spinosae Rivas Goday & Borja ex Tüxen 1962 Saginetea maritimae Westhoff, Leeuwen & Adriani 1962 Saliceteapurpureae Moor 1958 Stellarietea mediae Tüxen, W. Lohmeyer & Preising ex von Rochow 1951 Thero-Brachypodietea ramosi Braun-Blanquet 1947 Thlaspietea rotundifolii Braun-Blanquet 1948 Trifolio medii-Geranietea sanguinei Müller 1962 Table 1. Place and date of relevés: Rel. 1-7: Village of Dhërmi, 26 July 2014; rel. 1: 40°08'10.52" N, 19°38'49.65" E; rel. 2: 40°08'14.55" N, 19°38'48.88" E; rel. 3: 40°08'12.16" N, 19°38'49.87" E; rel. 4: 40°08'10.70" N, 19°38'50.24" E; rel. 5: 40°08'07.92" N, 19°38'51.82" E; rel. 6: 40°08'07.55" N, 19°38'52.31" E; rel. 7: 40°08'08.73" N, 19°38'54.02" E; Rel. 8-13: Town of Orikumi, 25 July 2014; rel. 8: 40°17'35.33" N, 19°29'28.95" E; rel. 9: 40°17'37.52" N, 19°29'28.46" E; rel. 10: 40°17'38.56" N, 19°29'27.95" E; rel. 11: 40°17'39.92" N, 19°29'27.42" E; rel. 12: 40°17'42.40" N, 19°29'28.12" E; rel. 13: 40°17'40.02" N, 19°29'31.93" E; Rel. 14-22: The island of Sazani (40°29'09.84" N, 19°17'15.91" E), from Hoda & Mersinllari (1996); Rel. 23-27: Village of Dukati, 25 July 2014; rel. 23: 40°14'33.24" N, 19°33'01.23" E; rel. 24: 40°14'32.79" N, 19°33'00.06" E ; rel. 25: 40°14'31.59" N, 19°32'59.00" E; rel. 26: 40°14'34.22" N, 19°33'01.12" E; rel. 27: 40°14'33.25" N, 19°32'02.75" E; Rel. 28-35: Southern slopes bellow the Llogara Pass, 26 July 2014; rel. 28: 40°11'06.18" N, 19°35'35.79" E; rel. 29: 40°11'06.54" N, 19°35'36.46" E; rel. 30: 40°11'06.84" N, 19°35'36.68" E; rel. 31: 40°11'07.38" N, 19°35'40.90" E; rel. 32: 40°11'06.92" N, 19°35'41.77" E; rel. 33: 40°11'06.62" N, 19°35'42.14" E; rel. 34: 40°11'07.97" N, 19°35'40.55" E; rel. 35: 40°11'05.79" N, 19°35'36.81" E. 65 O Hacquetia 15/1 • 2016, 49-65 N. 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FE LF No of relevés 1 2 3 4 5 6 7 8 9 10 11 No of group in Figure 4 9 9 9 9 9 9 9 6 6 6 6 No of species 25 19 15 12 11 14 17 30 15 14 15 Altitude (m) 5 5 15 20 16 19 38 66 64 64 65 Slope (°) 40 40 40 40 40 40 40 10 10 10 10 Aspect S SW SW S S S S NE NE NE NE Vascular plant cover (%) 95 100 100 100 100 100 100 100 100 100 100 Surface (m2) 100 100 100 100 100 100 100 50 50 50 100 Height of vegetation (m) 4 4 4 4 4 4 4 4 3-4 3-4 3-4 Char. Ass. P CME Quercus coccifera L. (incl. Q. calliprinos Webb) 5 5 5 5 5 5 5 3 5 4 5 Quercetea ilicis Oleo sylvestris-Ceratonion siliquae P CME Pistacia lentiscus L. 1 + + + + + + P CME Olea europaea L. var. sylvestris Brot. 1 + + + + + + + + + + P CME Myrtus communis L. + + + + + + + P CME Juniperus oxycedrus L. ssp. macrocarpa (Sibth. & Sm.) Ball Ch CME Prasium majus L. + P CME Calicotome villosa (Poiret) Link + + + P WME Phillyrea angustifolia L. Fraxino orni-Quercion ilicis P CME Spartium junceum L. 1 + + + + + + 2 1 1 1 P CME Smilax aspera L. + + + + + P MEAT Rubus ulmifolius Schott + + + + G CME Asparagus acutoflius L. + + + P CME Pistacia terebinthus L. 1 P CME Rubia peregrina L. P EME Cupressus sempervirens L. (juv.) + + + P CME Juniperus oxycedrus L. ssp. oxycedrus + + + P CME Quercus ilex L. Ch CME Teucrium flavum L. P CME Clematis flammula L. + + P CME Phillyrea latifolia L. + H EURAS Rubia tinctorum L. Companions Thero-Brachypodietea ramosi H CME Brachypodium retusum (Pers.) Beauv. + + 1 1 + + + H CME Carlina corymbosa L. + + + + H CME Hyparrhenia hirta (L.) Stapf + + + + + + + + + Ch IL-ADR Micromeria graeca (L.) Bentham ex Reichenb. G CME Asphodelus aestivus Brot. + T CME Pallenis spinosa (L.) Cass. + T MEAT Blackstonia perfoliata (L.) Hudson H WS Centaurium erythraea Rafn + 65 O Hacquetia 15/1 • 2016, 49-65 N. Jasprica, Z. Skvorc & M. Milovic A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 6 6 5 4 4 4 4 5 5 5 4 7 7 7 7 7 8 8 8 8 8 8 8 8 15 23 11 20 25 24 19 8 20 20 28 21 19 12 13 14 20 17 16 17 17 13 15 15 64 66 100 180 250 150 200 150 130 180 270 358 367 380 356 349 655 8 5 6 661 678 675 673 5 8 6 650 10 10 30 25 25 30 30 25 25 35 25 20 20 20 20 20 30 30 30 30 30 30 30 30 NE NE W SW WSW S W SW NW W SW E E NW NW NW W W W W W W W W Fr. 100 100 - - - - - - - - - 50 90 30 40 40 100 100 100 100 100 90 90 90 100 100 - - - - - - - - - 100 100 100 100 100 100 100 100 100 100 100 100 100 3-4 3-4 - - - - - - - - - 0.5 1-2 2 1.5 1.5 0.8 0.8 0.9 0.9 0.8 0.8 0.7 0.6 % 4 4 5 5 3 5 5 4 1 5 3 4 3 3 3 5 5 5 4 4 5 5 5 100 + + + 1 1 5 11 + . 1 + 11 4 1 1 + + + + + + 51 ......43 ......29 26 14 14 14 + + + + + + + 1 + 11 + 1 . 2 11 . 3 5 4 3.11 37 20 17 14 11 11 9 9 9 9 6 3 3 + + . + + 11 + + 11. . 1 . + . 1 1 . 111 + ..1+ 111111 +. + + +. +. +. + + + . + + + + 1 . + 1 . . 1 + + + + + . . + + + + + + + 57 + 43 43 31 17 14 14 14 65 + + + + + + + ++ + 2 + O Hacquetia 15/1 • 2016, 49-65 N. Jasprica, Z. Skvorc & M. Milovic A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) FE LF No of relevés T IL-ADR Bupleurum báldense Turra ssp. gussonei (Arcangeli) Tutin H SEME Piptatherum miliaceum (L.) Cosson H CME Andropogon distachyos L. T CME Brachypodium distachyon (L.) Beauv. G SEME Allium sphaerocephalon L. T WS Anagallis foemina Miller H NEME Helictotrichon convolutum (C.Presl) Henrard G CME Urginea maritima (L.) Baker T SEPO Avena sterilis L. T CME Briza maxima L. T CME Lagurus ovatus L. H IL-APEN Asyneuma limonifolium (L.) Janchen Cisto cretici-Micromerietea julianae + Erico-Cistetea P EME Phlomis fruticosa L. Ch CME Thymus capitatus (L.) Hoffmanns. & Link Ch EME Erica manipuliflora Salisb. P EME Cistus creticus L. ssp. eriocephalus (Viv.) Greuter & Bürdet* Ch CME Dorycnium hirsutum (L.) Ser. Ch CME Fumana scoparia Pomel Ch CME Micromeria juliana (L.) Bentham ex Reichenb. Ch CME Fumana thymifolia (L.) Spach ex Webb Ch CME Anthyllis hermanniae L. P EME Cistus salvifolius L. Festuco valesiacae-Brometea erecti Ch MEPO Teucrium polium L. H SEME Bromus erectus Huds. ssp. condensatus (Hack.) Asch. & Graebn. Ch SEE Thymus longicaulis C. Presl aggr. H EURAS Melica ciliata L. Ch EURAS Origanum vulgare L. H SEME Petrorhagia saxifraga (L.) Link H SEME Koeleria splendens C. Presl H NEME Stachys cretica L. ssp. salviifolia (Ten.) Rech. fil. H SEME Galium corrudifolium Vill. T MEAT Desmazeria rigida (L.) Tutin H IL-ADR Anthyllis x rubicunda Wender. T SEME Bromus hordeaceus L. T CME Echinaria capitata (L.) Desf. H SEPO Centaurea salonitana Vis. Ch IL-APEN Onosma echioides L. Ch SEPO Teucrium chamaedrys L. H EURAS Poa bulbosa L. H Festuca sp. Ch EURME Ononis spinosa L. H SEME Chondrilla juncea L. T EME Petrorhagiaprolifera (L.) P.W. Ball & Heywood H SEE Dorycniumpentaphyllum Scop. ssp. pentaphyllum H CME Asperula aristata L.f. ssp. scabra (J.Presl & C.Presl) Nyman 23456789 10 11 + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + 65 O Hacquetia 15/1 • 2016, 49-65 N. Jasprica, Z. Skvorc & M. Milovic A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 . . . . . . . . . + + + + + 14 + + . 14 1 1 + 2 . . . . . . . . . 1 + .1 1......... 1 1 . + 1 .. 2 + . + 1 . . . . . . + . + . + 1.. 1 1 + ......... 9 6 + 1......... 6 1 + ......... 6 + + ......... 6 57 51 40 26 23 11 11 9 9 3 2 + + + + + + + + + + + + + + + + + 1 1 1 . . 1 1 . . . 1 1 + 1 1 . 1 + 111 111 + + + .11 . 1 + + + + 11 + + + + + + . 40 1 26 + 23 . 20 + 20 . 17 . 17 . 14 + 14 1 1 1 1 1 1 9 9 9 9 6 6 6 6 79 + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + O Hacquetia 15/1 • 2016, 49-65 N. Jasprica, Ž. Škvorc & M. Milovic A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) FE LF No of relevés H EURAS Phleumpratense L. ssp. bertolonii (DC.) Bornm. Ch IL-ADR Genista pulchella Vis. ssp. pulchella Ch EURAS Thymuspulegoides L. aggr. H EURAS Scabiosa columbaria L. Ch IL-ADR Salvia officinalis L. Thlaspietea rotundifolii + Drypidetea spinosae H EME Scutellaria rubicunda Hornem. Ch IL-BALK Cerastium grandiflorum Waldst. & Kit. Artemisietea vulgaris H CME Scolymus hispanicus L. H CME Verbascum sinuatum L. H CME Dittrichia viscosa (L.) W. Greuter H IL-ADR Daucus carota L. ssp. major (Vis.) Arcangeli T MEAT Echium plantagineum L. H CME Foeniculum vulagre Mill. T MEPO Dasypyrum villosum (L.) P. Candargy H WS Cichorium intybus L. H MEAT Centaurea calcitrapa L. H CME Picnomom acarna (L.) Cass. H SEPO Eryngium campestre L. H SEPO Centaurea solstitialis L. T CME Sonchus asper (L.) Hill Querco roboris-Fagetea sylvaticae P EME Coronilla emerus L. ssp. emeroides (Boiss. & Spruner) G WS Pteridium aquilinum (L.) Kuhn G CME Cyclamen hederifolium Aiton P SEE Lonicera caprifolium L. P NEME Quercus trojana Webb P EME Quercus macrolepis Kotschy Asplenietea trichomanis P SME Putoria calabrica (L.f.) DC. Ch EME Inula verbascifolia (Willd.) Hausskn. P CME Ficus carica L. H SEME Ceterach officinarum DC. Paliuretea P EME Paliurus spina-christi Miller P IL-ADR Rhamnus intermedius Steudel & Hochst. Quercetea pubescentis+Rhamno-Prunetea P SEME Pyrus amygdaliformis Vill. H WS Clinopodium vulgare L. Trifolio medii-Geranietea sanguinei H EME Origanum heracleoticum L. Stellarietea mediae T EURAS Avena fatua L. T EURAS Veronica arvensis L. T WS Bromus sterilis L. T WS Sisymbrium officinale (L.) Scop. T CME Anthemis arvensis L. 62 23456789 10 11 + . + + 11 + + + + + + + + + + + + + + + + + + + + + + + + + O Hacquetia 15/1 • 2016, 49-65 N. Jasprica, Z. Skvorc & M. Milovic A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 + + + + + + .1 + .11 +.....1 + . + 1 . . 1 34 + +.........+11 + +........31 + + +.....................20 ........................11 ........................11 ......+ . +...............11 . + . . . . . . . . . . . . . . . . . . . . . . 6 . + . . . . . . . . . . . . . . . . . . . . . . 6 ...........+ +...........6 ...........+ +...........6 ........................ 3 . +......................3 . +......................3 ........+............... 3 20 14 6 6 3 3 2 + + + + + .......... +..+..++ 17 17 6 + + + + + +.........+ +...........14 ++....++.. 14 1+ 11 9 6 3 3 81 + + O Hacquetia 15/1 • 2016, 49-65 N. Jasprica, Z. Skvorc & M. Milovic A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) FE LF No of releves T CME Lathyrus cicera L. Molinio-Arrhenatheretea H EURAS Dactylis glomerata L. H SEME Hypericum perforatum L. H SEME Lotus corniculatus L. H EURAS Anthoxanthum odoratum L. H WS Plantago lanceolata L. Nerio oleandri-Tamaricetea africanae P CME Vit ex agn us-castus L. Ammophiletea G CME Elymuspycnanthus (Godr.) Melderis Salicetea purpureae P EURAS Salix eleagnos Scop. Saginetea maritimae T MEAT Parapholis filiformis (Roth) C.E.Hubbard 23456789 10 11 + + + + + 65 O Hacquetia 15/1 • 2016, 49-65 N. Jasprica, Z. Skvorc & M. Milovic A phytosociological analysis of the Quercus coccifera L. stands in south Albania (NE Mediterranean) 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 3 . . . . 1.1.....+...........9 + +......................6 ...........+ +...........6 ................1.1..........6 ...........+............3 +.............3 65