152 ACTA BOT. CROAT. 77 (2), 2018 Acta Bot. Croat. 77 (2), 152–160, 2018 CODEN: ABCRA 25 DOI: 10.2478/botcro-2018-0017 ISSN 0365-0588 eISSN 1847-8476 Subalpine vegetation in Giresun Mountains (Turkey) Rena Hüseyinova1, Erkan Yalçin2* 1 Şebinkarahisar School of Applied Sciences, Giresun University, Giresun, Turkey 2 Department of Biology, Faculty of Arts and Science, Ondokuz Mayıs University, Samsun, Turkey Abstract – In this study, the subalpine vegetation in the Giresun Mountains of northern Turkey was investigat- ed. The study area included north- and south-facing slopes at altitudes ranging between c. 2000 and 2500 me- ters. For vegetation classification and for describing the relationships between vegetation and environment, tra- ditional Braun-Blanquet methods and multivariate analysis techniques were used. The vegetation mainly consisted of subalpine grasslands and coniferous cushion scrubs. Caricetea curvulae and Astragalo microcepha- li-Brometea tomentelli were found to be dominant syntaxa in the vegetation of the study area. Land topography, soil physical and chemical factors and species richness have important impacts on the development of subalpine vegetation according to the results of multivariate analysis. Three associations and two subassociations were newly determined and classified. Hemicryptophytes, chamaephytes and geophytes participated in the floristic composition of these syntaxa. EUNIS habitat code and names for described syntaxa were also proposed. Keywords: environment, grasslands, life form, phytosociology, species richness, syntaxonomy * Corresponding author, e-mail: eryalcin@omu.edu.tr Introduction Mountainous areas are mainly located in the northern hemisphere (Vogiatzakis 2012) and orographic, edaphic and climatic factors have drawn an alpine borderline in these mountainous areas (Holtmeier and Broll 2005). Alpine and subalpine belts of mountains form specific ecological con- ditions for vascular flora and vegetation due to geographi- cal isolation, glaciation, the existence of microhabitats, cli- matic and historical changes (Uysal et al. 2011). Subalpine vegetation is greatly influenced by different types of natural and anthropogenic disturbance (Tappeiner et al. 1998) and is rather sensitive to global climate change (Cannone et al. 2007). It is expected that the subalpine plant composition will be changed by upward species migration due to ongoing global climate changes. These changes have been observed by the GLORIA monitoring network (GLORIA-Europe = Global Observation Research Initiative in Alpine Environ- ments and GLORIA-worldwide) in high mountain regions. Subalpine and alpine grasslands are considered biodiversity hotspots and included in the NATURA 2000 network as an EU priority habitat type (Pfeiffer et al. 2016). Syntaxonomic and ecological investigations on alpine and subalpine belts have been carried out by many researchers in Europe and Asia. The first vegetation investigation on al- pine vegetation in western and southern Anatolia was done by Quézel and Pamukçuoğlu (1970) and Quézel (1973). Re- hder et al. (1994) and Hein et al. (1998) researched new al- pine plant syntaxa in Uludag-Bursa province from Turkey. In these studies, the alpine vegetation was mostly included in the class Astragalo microcephali-Brometea tomentelli Quézel 1973 em. Parolly and its five orders. These orders include fol- lowing vegetation types; 1- Astragalo microcephali-Brometalia tomentelli Quézel 1973: Oreal to subalpine xerophytic grass- lands, dwarf-shrub and the thorny-cushion communities of Anatolia, 2- Drabo-Androsacetalia Quézel 1979: Alpine to subnival mat-forming communities, 3- Hyperico linarioidis- Thymetalia skorpilii Akman, Quézel, Yurdakulol, Ketenoğlu, Demirörs 1987: Xerophytic grasslands, dwarf-shrub and thorny-cushion communities of the West and Middle Black Sea Mountains, 4- Onobrychido armenae-Thymetalia leucos- tomi Akman, Ketenoğlu, Quézel & Demirörs 1984: Xerophyt- ic grasslands, dwarf-shrub and the thorny-cushion commu- nities of Central Anatolia, 5- Trifolio anatolici-Polygonetalia arenastri Quézel 1973: Hygrophytic to mesophytic vegetation of dolines, snow-patch and the melt water communities of the Taurus range (Hamzaoğlu 2006). SUBALPINE VEGETATION IN GIRESUN MOUNTAINS ACTA BOT. CROAT. 77 (2), 2018 153 Turkish phytosociologists and ecologists have mostly concentrated on the forest and steppe vegetation in moun- tainous areas in Turkey (Parolly 2004). Alpine and subal- pine vegetation studies were mostly carried out under the leadership of foreign researchers in Turkey. However, some of the Turkish botanists such as Akman et al. (1987), Tatlı (1987), Düzenli (1988), Kılınç and Karakaya (1992), Vural (1996), Uysal et al. (2011) partly focused on alpine and sub- alpine vegetation. Vural (1996) published a comprehensive syntaxonomic scheme about the alpine belts of mountains of the Eastern Black Sea, while Hein (1998) and Parolly (1998, 2004) did important revision for some of the syntaxa in the alpine vegetation of the West and Eastern Taurus Moun- tains. Although the syntaxonomic classification of vegeta- tion is very important, the relationships between vegetation and environment have not been sufficiently investigated for alpine and subalpine vegetation in Turkey. The phytosocio- logical approach of vegetation classification considered that each community is evolved by a site-specific range of envi- ronmental factors leading to differences in species composi- tion (Vonlanthen et al. 2006). The aim of this study was to describe syntaxa and the relationships between vegetation and environmental prop- erties of the subalpine communities in Giresun Mountains from northern Turkey. Materials and methods Study area The study area is situated at A7 according to the grid system of Davis (1965, 1985), and is approximately located between N 40°29'27'' and E 38°25'41''. The study area is surrounded by high mountains. In the study area, the subalpine belt extends from c. 2000 to 2500 m upwards on south and north-facing slopes. Giresun Mountains are a system of mountains that extend up to the peaks on Mt Karadağ, 3391 m, in the east and on the Karagöl plateau with 3095 m in the west. The study area is located in the north of Şebinkarahisar, between the Eastern Pontide Metallogenic Belt and the North Anatolian Fault system (Yavuz et al. 2008). The Turkish Pontides comprise the Cretaceous to Eocene igneous record of the convergence between Eurasia and Gondwana (Boztuğ et al. 2004) The nearest district to the study area (Şebinkarahisar) has a Mediterranean type of climate with 525 mm mean annual precipitation (P), and a drought period is observed in July with 0.5 mm precipitation according to the Şebinkarahisar State Meteorological Station. The mean annual temperature is 11.3 °C. Summer rainfall (PE) is 37 mm. The mean maximum for the hottest month (M) and mean minimum for the coldest month (m) are 30.3 and -16.1 °C, respectively. Index of xericity (S=PE/M) is 1.8. Pluviometric quotient (Q= 2000 P/ [(M + m + 546.4) * (M-m)] is 40.7, and the precipitation regime is sub-Mediterranean (spring, autumn, winter, summer). Northeast winds predominate, thus north facing slopes are partly windward, whereas the south facing slopes are leeward. The north facing slopes are snow covered for a longer time than the south facing slopes. The vegetation mostly consists of Festuca-dominated grasslands and traditionally has been grazed by sheep, cows and goats since the 19th century. However, grazing pressure has gradually decreased over the last two decades due to de- clining livestock production. Vegetation sampling The taxonomy of vascular plants follows Davis (1965, 1985), Davis et al. (1988), Tutin et al. (1964, 1980), Güner et al. (2000) and Güner et al. (2012), respectively. All plant samples were preserved in the Herbarium of the Faculty of the Arts-Sciences of Ondokuz Mayıs University (OMUB). Non vascular plants were omitted. For sampling procedure, we considered the study area according to the north and the south facing slopes. Both of them occupied altitudes ranging between 2000 and 2500 m. Five floristically and environmentally homogeneous sample plots were established with 100 m altitudinal intervals on each slope. Plot size was selected as 20 and 50 m2 (van der Maarel 2005). In total, 60 plots were obtained during the spring and summer months of 2015 and 2016. Phytosocio- logical records were carried out by using the cover/abun- dance values proposed by Braun-Blanquet (1964) in each plot. In each plot, the slope degrees (%), altitude (meters above the sea level), aspect and GPS coordinates were also logged. Aspect in degrees was transformed using the formula of Beers et al. (1966). Plant associations were named according to the Interna- tional Code of Phytosociological Nomenclature (ICPN) (We- ber et al. 2000). Syntaxonomic interpretations were made by using the available phytosociological literature (Quézel 1973, Akman et al. 1987, Vural 1996, Mucina 1997, Parolly 1998 and 2004, Onipchenko 2002, Mucina et al. 2016). Plant life forms were identified according to Mueller-Dombois and El- lenberg (2002). EUNIS habitat code and names were identi- fied according to Davies et al. (2004) and EUNIS habitat type hierarchical view (EEA 2017). Soil sampling and analysis From each plot, soil samples were taken at a depth of 20 cm from the topsoil. Soil samples were then air dried and sieved through a 2 mm mesh prior to analysis. Soil organ- ic matter (%) was determined using the Walkley and Black method (Black et al. 1965). Soil texture was determined us- ing the Bouyoucus hydrometer method, and the clay con- tent was expressed as %. The pH values were measured in deionized water (1:1) and the soil nitrogen (%) was deter- mined by the micro Kjeldahl method. The soil phosphorus (ppm) was determined spectrophotometrically following the extraction by ammonium acetate. Soil potassium (cmol(+)/ kg) was determined using a Petracourt PFP-7 flame photom- eter after nitric acid wet digestion. The CaCO3 (%) concen- trations were determined using a Scheibler calcimeter. For the determination of soil moisture, freshly weighed soil sam- ples were air dried for 48 h to calculate wet to dry mass ratios. HÜSEYINOVA R., YALÇIN E. 154 ACTA BOT. CROAT. 77 (2), 2018 These values were used to calculate soil moisture (gravimet- ric method) (Kacar 2009). Data analysis Prior to data analysis, 10 vegetation plots were subjec- tively excluded due to high heterogeneity and their uncertain positions. The resulting dataset of 50 plots and 160 species was classified by the PC-ORD program (McCune and Grace 2002), using Ward’s method and the Jaccard similarity index as a resemblance measure. “Crispness of the classification” was used for the optimal number of clusters (Botta-Dukát et al. 2005). Diagnostic taxa for each group were defined in the JUICE program (Tichý 2002) by calculating the fidelity of each species to each group (Chytrý et al. 2002) using the φ-coefficient as the fidelity measure. Species with a fidelity of above 50 (φ > 50) were considered as diagnostic. To detect gradients in species composition and species- environment relations, canonical correspondence analysis (CCA) was performed by using the ECOM 2.1.3.137 version software programme (Seaby and Henderson 2007). Canon- ical correspondence analysis includes two matrices, one of which has 50 plots, classified into syntaxa by phytosociologi- cal analysis, × 160 species (average % cover values of species corresponding to the transformations of the Braun-Blanquet scale as proposed by van der Maarel (1979) were r = 1, + = 2, 1 = 3, 2m = 4, 2a = 5, 2b = 6, 3 = 7, 4 = 8, 5 = 9) and second matrix 50 plots × 7 environmental parameters (soil mois- ture (SM), soil pH, soil clay content (Clay), total soil nitro- gen ratio (N), aspect, altitude and slope). The interpretation of the results was made by the first two axes because only these two axes were statistically significant. Statistically sig- nificant variables that derived from CCA were shown by a bold number. Species diversity was calculated as the Shannon–Wiener index at log base 10 (Magurran 2004) performed by using Biodiversity version 2.0 (McAlleece et al. 1997) software pro- gramme, respectively. Moreover, the comparison of growth form and species diversity parameters for the described com- munities was visualized by a Box-Whisker diagram prepared in SPSS 21.0 version (IBM Corp. 2012). The data of the Şebinkarahisar (Giresun) State Meteoro- logical Station, the nearest meteorology station to the study area, were used to explain the climatic properties. Results Cluster analyses yielded that vegetation of Eğribel Pass was shown by four main clusters (Fig. 1). Crispness of clas- sification value verified that the data set was classified in four clusters. The most prominent split within the four-clusters concerned the cluster C, which is clearly separated from the other clusters due to syntaxonomic distinction on the basis of the summed up average cover values of diagnostic species. We defined two grassland associations and subassocia- tions dominated by Festuca, and a cushion Juniperus scrub on the different slopes. The south facing slopes had drier soil conditions than the north facing slopes. Cluster A contained the plots sampled in the southern slopes in the study area. Festuca pinifolia var. pinifolia, Bunium microcarpum subsp. bourgaei, Veronica peduncularis and Lotus corniculatus var. alpinus were constant. We defined two subassociations from the cluster. Subassociations were formed by different envi- ronmental variables such as altitude, slope, soil organic mat- ter and nitrogen ratios (Tab. 1). We propose the following name for the association and subassociations: Bunio micro- carpae-Festucetum pinifoli ass. nova hoc loco (holotypus: plot 14 in On-line Suppl. Tab. 1), Bunio microcarpae-Festucetum pinifoli subass. oxytropetosum lazicae subass. nova hoc loco (holotypus:plot 10 in On-line Suppl. Tab. 1) and Bunio mi- crocarpae-Festucetum pinifoli subass. verbascetosum froedinii subass. nova hoc loco (holotypus: plot 16 in On-line Suppl. Tab. 1), respectively. Oxytropis lazica and Verbascum froe- dinii were dominant taxa in the subassociations. Oxytropis lazica spread on mesic slopes while Verbascum froedinii oc- cupied drier slopes. Cluster B included plots where the vegetation was dom- inated by Festuca amethystina subsp. orientalis var. turcica, Lotus corniculatus var. corniculatus and Scorzonera cana var. radicosa in northern slopes. We named cluster B Lotus cor- niculati-Festucetum turcici ass. nova hoc loco (holotypus:plot 33 in On-line Suppl. Tab. 1). Bunio microcarpae-Festucetum pinifoli and Lotus corniculati-Festucetum turcici were differ- entiated by the bulk of Caricetea curvulae Br.-Bl. 1948 and Caricetalia curvulae Br.-Bl. in Br.-Bl. et Jenny 1926 species in these clusters puts them in these syntaxa. At the alliance level, their affiliation is clearly with Agrostio lazicae-Sibbal- dion parviflorae Vural 1996 owing to the species diagnostic of alliance. Moreover, Molinio-Arrhenatheretea R. Tx. 1937 and Astragalo microcephali-Brometea tomentelli Quézel 1973 em. Parolly species participated in the floristic composition of both of them. Cluster C which was coniferous cushion scrublands comprised of the plots sampled in the southern slopes in Fig. 1. Dendrogram of vegetation plots (50 plots and 160 species) of the Giresun Mountains. A: Bunio microcarpae-Festucetum pinifolii, 1: oxytropetosum lazicae, 2: verbascetosum froedinii, B: Lotus cornic- ulati-Festucetum turcici, C: Alysso pseudo-mouradici-Juniperetum saxatilis stands. SUBALPINE VEGETATION IN GIRESUN MOUNTAINS ACTA BOT. CROAT. 77 (2), 2018 155 Tab. 1. The mean values environmental, soil parameters and species richness in the associations and their comparison with ANOVA test. The difference in letters indicates significant difference (P < 0.05) between means according to Tukey’s (HSD) test among associations and subassociations. BG – between groups, WG – within groups; BFol – Bunio microcarpae-Festucetum pinifoli subass. oxytropetosum lazicae, BFvfi – Bunio microcarpae-Festucetum pinifoli subass. verbascetosum froedinii, LF – Lotus corniculati-Festucetum turcici, AJ – Alysso pseu- do-mouradici-Juniperetum saxatilis. Parameter Associations Mean±std. error Sum of squares df Mean square F P Aspect BFol 0.68±0.21 b BFvfi 0.32±0.09 b 15.27 (BG) 3(BG) 5.09(BG) 17.13 <0.01 LF 1.62±0.08 a 15.16 (WG) 51(WG) 0.29(WG) AJ 1.44±0.14 a Slope BFol 24.33±2.22 b BFvfi 42.00±0.81 a 2794.05(BG) 3(BG) 931.35(BG) 17.92 <0.01 LF 27.75±1.96 b 2649.58(WG) 51(WG) 51.95(WG) AJ 39.50±0.89 a Altitude BFol 2290.86±16.69 a BFvfi 2113.10±14.16 b 206244.59(BG) 3(BG) 68748.19(BG) 6.19 <0.01 LF 2257.70±35.55 a 566279.33(WG) 51(WG) 11103.51(WG) AJ 2245.50±10.13 a Species richness BFol 23.13±0.63 a BFvfi 26.90±1.90 a 929.97(BG) 3(BG) 309.99(BG) 16.85 <0.01 LF 17.50±1.09 b 937.73(WG) 51(WG) 18.38(WG) AJ 27.30±0.86 a Soil moisture BFol 6.23±0.96 ab BFvfi 3.59±1.06 b 396.25(BG) 3(BG) 132.08(BG) 10.71 <0.01 LF 9.32±0.93 a 628.87(WG) 51(WG) 12.33(WG) AJ 2.54±0.09 b pH BFol 4.65±0.12 b BFvfi 5.10±0.19 b 7.22(BG) 3(BG) 2.40(BG) 11.21 <0.01 LF 4.76±0.10 b 10.95(WG) 51(WG) 0.21(WG) AJ 5.65±0.01 a Silt BFol 27.86±1.35 ab BFvfi 27.73±1.06 b 527.07(BG) 3(BG) 175.69(BG) 8.04 <0.01 LF 32.69±1.24 a 1113.96(WG) 51(WG) 21.84(WG) AJ 24.39±2.17 b Sand BFol 59.30±1.34 b BFvfi 58.41±1.73 b 738.76(BG) 3(BG) 246.25(BG) 7.86 <0.01 LF 55.42±1.54 b 1596.41(WG) 51(WG) 31.30(WG) AJ 65.91±0.64 a CaCO3 BFol 2.79±0.24 a BFvfi 2.47±0.29 ab 5.97(BG) 3(BG) 1.99(BG) 2.88 0.04 LF 2.57±0.19 ab 35.13(WG) 51(WG) 0.68(WG) AJ 1.82±0.06 b Organic matter BFol 11.18±1.32 b BFvfi 4.78±0.53 c 1123.63(BG) 3(BG) 374.54(BG) 28.98 <0.01 LF 15.65±0.82 a 659.14(WG) 51(WG) 12.92(WG) AJ 5.45±0.19 c Total soil nitrogen BFol 0.54±0.06 b BFvfi 0.27±0.04 c 2.27(BG) 3(BG) 0.75(BG) 24.66 <0.01 LF 0.75±0.03 a 1.56(WG) 51(WG) 0.03(WG) AJ 0.27±0.09 c the study area and, dominated by Juniperus communis var. saxatalis, Alyssum pseudo-mouradicum and Dianthus zederbaueri. Cluster C is named as Alysso pseudo-mouradici- Juniperetum saxatilis ass. nova hoc loco (holotypus: plot 58 in On-line Suppl. Tab. 1). In the association, Astragalo microcephali-Brometea tomentelli Quézel 1973 em. Parolly, Hyperico linarioidis-Thymetalia scorpilii Akman, Quézel, Yurdakulol, Ketenoğlu, Demirörs In all, 1987 species are present. The diagnostic species of Peduicularo comosa- Asterion alpini Akman, Quézel, Yurdakulol, Ketenoğlu, Demirörs 1987 were represented in this association such as Aster alpina and Jasione supina subsp. pontica. HÜSEYINOVA R., YALÇIN E. 156 ACTA BOT. CROAT. 77 (2), 2018 The syntaxonomic scheme for syntaxons in the study area is as follows: Class: Astragalo microcephali-Brometea tomentelli Quézel 1973 em. Parolly Order: Hyperico linarioidis-Thymetalia scorpilii Akman, Quézel, Yurdakulol, Ketenoğlu, Demirörs 1987 Alliance: Peduicularo comosa-Asterion alpini Akman, Quézel, Yurdakulol, Ketenoğlu, Demirörs 1987 Association: Alysso pseudo-mouradici-Juniperetum saxatilis ass. nova hoc loco Class: Caricetea curvulae Br.-Bl. 1948 Order: Caricetalia curvulae Br.-Bl. in Br.-Bl. et Jenny 1926 Alliance: Agrostio lazicae-Sibbaldion parviflorae Vural 1996 Association 1: Bunio microcarpae-Festucetum pinifoli ass. nova hoc loco Subassociation 1.1: oxytropetosum lazicae subass. nova hoc loco Subassociation 1.2: verbascetosum froedinii subass. nova hoc loco Association 2: Lotus corniculati-Festucetum turcici ass. nova hoc loco Hemicryptophytes were the dominant life form in all syntaxa, while chamaephytes and geophytes also contribut- ed to the life form composition (Fig. 2). Alysso pseudo-mou- radici-Juniperetum saxatilis had the highest species richness and most diverse syntaxon while Lotus corniculati-Festuce- tum turcici was the syntaxon with lowest species richness and the least diverse syntaxon (Figs. 3 a,b). The environmental factors and species richness of syntaxa were statistically dif- ferent (Tab. 1). Lotus corniculati-Festucetum turcici contained species related to plots with more humid, nitrogenous, silty soil, rich in organic matter, while Alysso pseudo-mouradici- Juniperetum saxatilis had species related to plots with sandy soil and plots of greater inclination. The first two CCA axes were significant (P < 0.001) ac- cording to the Monte-Carlo permutation test. Eigenvalue of axis 1 was 0.70 and explained 13.75% of total variance, while eigenvalue of axis 2 was 0.51 and explained 23.87% of total variance (On-line Suppl. Tab. 2). According to CCA ordi- nation axes, aspect, slope, altitude, soil moisture, clay and nitrogen contents were important environmental parame- ters for the development of different vegetation types in the study (On-line Suppl. Tab. 2). The slope, proportion of soil moisture, clay and nitrogen were negatively correlated with the first and second CCA axes, while aspect and altitude were positively correlated with the second axis (Fig. 4). In the CCA ordination, Alysso pseudo-mouradici-Juniperetum saxatilis preferred higher soil moisture, clay and nitrogen Fig. 2. Life form spectra for associations and subassociations. BFol – Bunio microcarpae-Festucetum pinifoli subass. oxytropeto- sum lazicae, BFvfi – Bunio microcarpae-Festucetum pinifoli subass. verbascetosum froedinii, LF – Lotus corniculati-Festucetum turcici, AJ -Alysso pseudo-mouradici-Juniperetum saxatilis. Fig. 3. Species richness (a) and Shannon diversity index values (b) for associations and subassociations. The difference in letters indi- cates the significant difference (P < 0.05) between means accord- ing to Tukey’s (HSD) test among associations and subassociations. Abbreviations for associations and subassociations are as in Fig. 2. SUBALPINE VEGETATION IN GIRESUN MOUNTAINS ACTA BOT. CROAT. 77 (2), 2018 157 content in spite of the spread of Lotus corniculati-Festucetum turcici on the highest altitude slopes. Caricetea curvulae Br.- Bl. 1948 plots were located in a separate position towards the right margin of the ordination space and showed high corre- lations with topographical factors such as altitude, slope and aspect (Fig. 4). Bunio microcarpae-Festucetum pinifoli and Alysso pseudo-mouradici-Juniperetum saxatilis spread on the south facing slopes in spite of Lotus corniculati-Festucetum turcici occupying the north facing slopes. The plots of Bunio microcarpae-Festucetum pinifoli subass. oxytropetosum lazi- cae spread on the landscapes that had higher altitudes and lower slopes than Bunio microcarpae-Festucetum pinifoli sub- ass. verbascetosum froedinii plots. Discussion In the Eastern Black Sea region, detailed research deal- ing with vegetation of the highest mountains above the tim- berline began in the 1980s. In the first studies, Turkish au- thors (Düzenli 1988, Kılınç and Karakaya 1992, Vural 1996) recorded a number of vegetation plots from different alpine and subalpine communities. Düzenli (1988) investigated the vegetation in the alpine zone of Mount Tiryal (Artvin). He represented the alpine vegetation as associations Alchemi- llo-Campanuletea tridentate Quézel et Düzenli 1979 and Al- chemillo-Campanuletalia tridentatae Quézel et Düzenli 1979, respectively. Vural (1996) studied the high mountain vegeta- tion of Rize. In his paper, subalpine vegetation was consid- ered within one class and two orders. Kılınç and Karakaya (1992), suggested that subalpine vegetation in the Ordu- Çambaşı Plateau should be included in the class Molinio-Ar- rhenatheretea R. Tx. 1937, order Arrhenatheretalia and class Alchemillo retinervis-Sibbaldietea parviflorae Vural 1996. Aspect, slope and altitude influence the plant commu- nities and environmental conditions in the Giresun Moun- tains. Consequently, in this study, different plant associa- tions and subassociations were defined. Vural (1996), in his study, formerly described Alchemillo retinervis-Sibbaldietea parviflorae Vural 1996, Alchemillo retinervis-Sibbaldietalia parviflorae Vural 1996 and Swertio ibericae-Nardetalia stric- tae Vural 1996. But Parolly (2004) proposed that all of them should be included in Caricetea curvulae Br.-Bl. 1948 and Caricetalia curvulae Br.-Bl. in Br.-Bl. et Jenny 1926. Onip- chenko (2002) supported a similar syntaxonomic scheme for the subalpine vegetation of the Teberda Reserve, the North- western Caucasus. We agreed with this viewpoint and firstly classified the subalpine Festuca grasslands within these syn- taxa in Turkey. At the alliance level, Agrostio lazicae-Sibbal- dion parviflorae Vural 1996 was accepted by Parolly (2004) for subalpine grassland vegetation from northern Turkey. We also considered that alliance for the syntaxa dominated by Festuca in this study. A coniferous cushion scrub commu- nity spread at 2200-2300 meters altitude was dominated by Juniperus communis var. saxatilis in this study. Akman et al. (2014) previously reported a similar community on Il- gaz Mountain. In the present study, it was found that the bulk of Astragalo microcephali-Brometea tomentelli and Hy- perico linarioidis-Thymetalia scorpilii species in the floristic composition of the coniferous cushion scrub association was dominated by Juniperus communis var. saxatilis. At the alli- ance level, the association affiliated with Peduicularo como- sa-Asterion alpini Akman, Quézel, Yurdakulol, Ketenoğlu, Demirörs 1987. Cluster groups in dendrogram of plots that also reflected a floristic assemblage characterized by a mix- ture of species of Caricetea curvulae and Astragalo micro- cephali-Brometea tomentelli. Hemicryptophytes were dominant in accordance with general characterisation of subalpine vegetation in this study. Hemicryptophytes were fairly common life form in alpine and subalpine landscape, which showed adaptations to snowbed environmental conditions by the strong per- sistence via plurennial stocks or dense turfs, above-ground renewal buds over winter, lateral spreading over short dis- tances and generalistic diaspore dissemination (Komac et al. 2015, Scheepens et al. 2015). Nevertheless, considerable per- centages of particular plant types (like therophytes, various kinds of chamaephytes, succulents, evergreens and berry- producers) generated a highly diversified alpine belt (Illa et al. 2006). Chamaephytes also presented low percentages in the floristic composition that were related to various stress conditions in this study. These plants mainly grew in infer- tile soils and were able to persist for many years with small above-ground lignified perennial structures (Sanz‐Elorza et al. 2003, Schweingruber 2007). In this study, the environmental factors influenced vege- tation differentiation at different physiognomic and syntaxo- nomic levels (Onipchenko and Semenova 1995, Onipchenko 2002). Etzold et al. (2016) reported that altitude, slope and aspect were also the main topographical factors and formed different syntaxa in subalpine and alpine grassland vegetation in the northeastern Greater Caucasus of Azerbaijan. Noroozi et al. (2010) also reported that slope and aspect were also the main driving topographical factors on vegetation differentia- Fig. 4. Canonical correspondence analysis ordination diagram of the dendrogram groups related to environmental factors in the study area. Abbreviations for associations and subassociations are as in Fig. 2. SM – soil moisture ratio, N – total soil nitrogen ratio. HÜSEYINOVA R., YALÇIN E. 158 ACTA BOT. CROAT. 77 (2), 2018 tion in the high alpine vegetation of the Tuchal Mountains (Central Alborz, Iran). Likewise, new plant associations and subassociations were described depending on altitude and aspect gradients, in our study. The position of Caricetea cur- vulae Br.-Bl. 1948 probably reflected a moisture gradient in terms of the longer persistence of snow. Altitude decreases temperature while increases precipitation that indirectly in- fluence vegetation (Walther et al. 2005). Aspect could indi- rectly alter soil moisture and mineralization due to their ef- fect on the solar radiation (Winkler et al. 2016). Therefore, we found that different plant associations grew on different slopes and humid soils. Gottfried et al. (1998) and Pauli et al. (1999) showed increasing elevation limits of alpine grassland from south-western to south-eastern slopes on Schrankogel in the central Eastern Alps. Besides, this study showed that south-exposed mountain slopes favour local-scale species richness, compared to the northern sides of the same moun- tains. This is consistent with the species-energy hypothesis (Wright 1983) and other temperature-related diversity hy- potheses (Currie et al. 2004), suggesting temperature-driven processes as decisive determinants of vascular plant species richness (Winkler et al. 2016). CCA reflected soil clay content being one of the impor- tant environmental factors for the floristic composition of subalpine vegetation. The soil particle size is determined by microtopography and fine-textured soils (mainly clay and silt) have higher water-holding capacity (Michalet et al. 2002). Zanelli et al. (2007) hypothesised that vegetation change has led to changes in soil chemistry and soil miner- alogy. Clay minerals are often a weathering product of the near-surface that is dependent on the precursor minerals and the surrounding environmental conditions (Egli et al. 2008). In addition to this, the clay contents in soils are favourable for the immobilization of nutrient ions and enzymes. Sub- alpine soils have high contents of total nitrogen, which also explains the lower activities and microbial biomass in these soils. Nitrogen is generally a limiting factor for soil biologi- cal processes (Margesin et al. 2009). We also found that soil clay and nitrogen content had an impact on vegetation dif- ferentiation in subalpine belts in the Black Sea Mountains. New subalpine syntaxa were identified in the Black Sea Mountains in Turkey. They were included in two classes Caricetea curvulae Br.-Bl. 1948 and Astragalo microcephali- Brometea tomentelli Quézel 1973 em. Parolly. Astragalo mi- crocephali-Brometea tomentelli Quézel 1973 em. Parolly was widespread on the different altitudes and bedrocks in east- ern and inner Anatolia. This study revealed that this class penetrated into the Black Sea high mountain chains. It also showed that Caricetea curvulae Br.-Bl. 1948 and Astragalo microcephali-Brometea tomentelli Quézel 1973 em. Parolly were distributed together on different altitudes and aspects in the subalpine belt in Black Sea Mountains in Turkey. How- ever, these classes diverged in some environmental param- eters such as altitude, slope, aspect, soil nitrogen, moisture and clay contents. The EUNIS habitat classification is important for species, habitat types and designated sites compiled in the framework of Natura 2000 (Davies et al. 2004, Özüdoğru and Duygu 2009). Festuca-dominated grasslands could be included in Pontic alpine grasslands as EUNIS habitat code E4.441. The Juniperus communis dominated syntaxa were classified in EUNIS by the habitat code and name F3.16 and Juniper- us communis scrub in Europe, respectively. We newly de- scribed a Juniperus scrub syntaxon which could not be in- cluded in this habitat code and name. Therefore, we have proposed a new EUNIS habitat code and name for this syn- taxon as F3.165 and Black Sea subalpine juniper thickets, respectively. Acknowledgement This study was funded by the Research Council of Gire- sun University (Project number: FBA 250414-79). Spe- cial thanks are due to Rıdvan Kızılkaya (PhD), Ali Kavgacı (PhD), and to Sıddık Yüksel (English specialist) for their valuable contributions. 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