This article was downloaded by: [York University Libraries] On: 17 November 2014, At: 02:38 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T3JH, UK Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology: Official Journal of the Societa Botanica Italiana Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/1oi/ tplb20 Sand topography influences the distribution of xerohalophytic vegetation on a southern Adriatic beach in Montenegro A. Mijovic a , Z. Popovic b , I Vukov b , M. Smiljanic b , R Matic b & S Bojovic b a Institute for Nature Protection , Dr. Ivana Ribara 91, 11070, Belgrade, Republic of Serbia b Department of Ecology , Institute for Biological Research, University of Belgrade , Bulevar Despot a St ef ana 142, 11060, Belgrade, Republic of Serbia Accepted author version posted online: 26 Apr 2012.Published online: 21 May 2012. To cite this article: A. Mijovic, Z. Popovic, T. Vukov, M. Smiljanic, R. Matic &S Bojovic (2012) Sand topography influences the distribution of xerohalophytic vegetation on a southern Adriatic beach in Montenegro, Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology: Official Journal of the Societa Botanica Italiana, 146:3, 664-673, DOI: 10.1080/11263504.2012.687403 To link to this article: http:/ / dx. doi. org/10.1080/11263504.2012.687403 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the "Content") contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. 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MIJOVIC1, Z. POPOVIC2, T. VUKOV2, M. SMILJANIC2, R. MATIC2, & S. BOJOVIC2 1 Institute for Nature Protection, Dr. Ivana Ribara 91, 11070 Belgrade, Republic of Serbia and 2Department of Ecology, Institute for Biological Research, University of Belgrade, Bulevar Despota Stefana 142, 11060 Belgrade, Republic of Serbia Abstract Totally, 26 xerohalophytic species belonging to the associations Cakilo-Xanthietum italici and Echinophoro-Elymetum farcti were registered at the Veliká plaza beach (Montenegro); they were distributed following the spatial succession common in this type of habitat. Three transects of different terrain morphology (positioned perpendicularly to the shoreline) were surveyed to establish if there were any differences in the floristic composition and species distribution due to the configuration discrepancy. Moreover, each transect was divided into zones: The first zone from 0 to 30 m inland, the second zone from 30 m to 60 m inland, and the third zone from 60 m to the end of beach (75-115 m, depending on transect length). The floristic composition was relatively uniform in the different transects. However, the number of individuals per species differed significantly among these transects. An analysis of the qualitative composition of transects/zones showed high differences between zones within transects I and II, whereas species composition of zones within transect III was mostly similar. When analyzing the quantitative composition of species, a positive correlation between the distance from the shoreline and the number of individuals per species in transects I and II was registered for most analyzed species (Spearman correlation, p < 0.001). This indicates that terrain configuration affected the ability of present species to become established and propagate. Keywords: Cakilo-Xanthietum italici, Echinophoro-Elymetum farcti, psammophytic vegetation, sand micro-topography, species distribution, xerohalophytic vegetation Introduction Xerohalophytic vegetation of the sand dunes along the Adriatic coast is predominantly distributed on the western Italian forecast, where sandy shores are a frequent type of habitat (Pignatti 1959; Caniglia 1982), but to a much lesser extent in Croatia and Montenegro where rocky and steep shorelines make up more than 80% of coastal reliefs (Horvatic 1974; Jankovic & Stevanovic 1983). This vegetation type is endangered by tourism, which is the reason for special attention being given to these species and their habitats in biodiversity protection programs both in Montenegro and adjacent countries. On eastern Adriatic sandy coasts, the characteristic plant species constituting sand pioneer and dune xerohalophytic vegetation are included into plant associations from two classes: Cakiletaea maritimae Tx. et Prsg. 1950 and Ammophiletea Br.-Bl. & Tiixen ex Westhoff, Dijk & Passchier 1946 (Mijovic & Stevanovic 1988; Pulevic 1973, 1982). Plants at foreland habitats are exposed to specific stresses, such as salt spray, episodic over-wash, highly unstable substrate, low field capacity, high air and soil temperatures, drought, and strong wind (Barbour et al. 1985; Rozema et al. 1985; Clark 1986; Hesp 1991; Kumler 1997; Randall & Scott 1997). In accordance with their basic ecological features, such as persistence and ability to overcome water shortage, xerohalophytic species show more or less regular zonal distribution perpendicular to the shoreline (Maun 1994). Thus, the draft zone is mostly occupied by the "pioneers of dunes", i.e. annuals with extensive root systems (Salisbury 1952). The zone which follows is mostly inhabited by the same but more frequent species and partly by perennials. The perennial species associated with Correspondence: Zorica Popovic, Department of Ecology, Institute for Biological Research, University of Belgrade, Bulevar Despota Stefana 142, 11060 Belgrade, Serbia. Tel: +381 11 2078 360. Fax: +381 11 2761 433. Email: zorica.popovic@ibiss.bg.ac.yu ISSN 1126-3504 print/ISSN 1724-5575 online © 2012 Societa Botanica Italiana http://dx.doi.org/10.1080/11263504.2012.687403 Xerohalophytic plant distribution along sand beach 665 lower sand salinity inhabit the third zone, which meets with another type of vegetation - a typical garrigue (Salisbury 1952; Ranwell 1972). However, in the investigated area, instead of garrigue, stands of the class Thero-Brachypodietea Br.-BL 1947 are present (Mijovic & Stevanovic 1988). Specific depositional topography alters the physico-chemical properties of the sand base and, consequently, can affect the distribution and number of species (Costa et al. 1996). Therefore, the aim of our study was to demonstrate the distribution of xerohalophytic vegetation in relation to terrain geo-morphology. Materials and methods Study area Field data were collected from the Veliká plaza beach near Ulcinj (location given in Figure 1) during the growth season of 2002. This is the most southern point of the Montenegrinian coast, and the largest beach in this part of southern Adria (approximately 11.5 km in length); it extends into the Albanian foreland south of the delta of the Bojana River. The study area is characterized both by high patchiness and fragmentation, with various landscape elements such as mobile sand dunes and semi-fixed dunes. Meteorological data for the Ulcinj area over a 30-year period (1969-1999), obtained from the Hydro-meteorogical Service of the Republic, are given in Table I. Figure 1. Geographical position of Veliká plaza. Vegetation survey The survey was made by total plant census along the three strip transects (each 3-m wide) of unequal length perpendicular to the shoreline and randomly selected quadratic plots. The starting point of all three transects is the shoreline (0 m from the sea). Each transect survey was conducted along different shore configurations to establish if there were differences in the floristic composition due to configuration discrepancy. The first transect was made across rising terrain (from sea level going inland), the second one was made along the dune where the terrain first rose and then descended going inland, and the third one was made along the depression (i.e. the terrain descended and then rose going inland) (Figure 2). The length of transects depended on the floristic composition (transect census was made as long as the species of the associations Cakilo-Xanthietum italici and Echino-phoro-Elymetum farcti were present). When the floristic composition changed, i.e. species from another type of vegetation predominated, the survey was stopped. Each transect was divided into three zones: The first zone from 0 to ca. 30 m inland; the second zone from ca. 30 to 60 m inland; and the third zone from ca. 60 m to 114, 99 and 75 m inland in the first, second, and third transect, respectively. Eighteen plots were selected randomly (size of plot 25 m2) at different distances from the sea (from 30 to 120 m). Total census of individuals present in the each square was carried out during the survey. The standard keys for determination of plants were used (Tutin et al. 1964a, 1964b, 1972, 1976, 1980; Domac 1984). A combined scale was used for evaluation of species' abundance and cover, as proposed by Westhoff and van der Maarel (1973). Substrate analyses Along each transect, surface soil samples were taken, from each zone, three times during the growth season (April, June, and September; Mijovic 1997). Soil samples (0-20 cm depth) from each transect/zone square were taken at three different points and mixed into a composite sample for a specific transect/zone combination. The sand samples were immediately placed into 30-ml containers and taken back to the laboratory, weighed, dried at 105°C and weighed again to calculate the water content. The air-dried subsam-ples were passed through a 2-mm sieve. The electrical conductivity (EC) was measured in a 1:5 soikwater extract (Rhoades et al. 1999). Nitrogen (N) determination was done by semi micro-Klejdahl analysis according to Markham (1942). Samples for analysis were digested with sulfuric acid at 350°C for 4 h. Results of substrate analyses are given in Table II. 666 A. Mijovič et al. Table I. Meteorological data for Ulcinj area for 30-year period (1969-1999). I II III IV V VI VII VIII IX X XI XII Yearly VI-X T 6.9 10.4 10.4 13.7 18 21.6 24.4 24.2 21.1 16.9 12.3 8.5 15.5 20.5 P 157.7 138.8 116 113.6 62.2 52.1 29.8 50 85.5 142 173 157.4 1278.1 393 I 116.1 119.2 164.8 194.3 253.6 289.1 337.7 312.2 247.8 191.8 120.7 106.4 204.5 275.7 Months from January (I) to December (XII); T, temperature (°C); P, precipitation (%); and I, duration of sunshine (hours). Class Thcio-Braehypodictca Br.-BI. 1947 Transccl i SEA Zone I Zoos H Zone III Transccl I Transecl ; 3: I -i (A) SEA (B) Figure 2. Surveyed transects: (A) horizontal profile and (B) vertical profile. Table II. Characteristic of soil probes taken from surveyed transects/zones (mean values + SD). Probe Water content (%) Total N content (ppm) pH Electrical conductivity (mmol/L) T1Z1 2.7 + 0.02 68 + 0.4 7.30 + 0.06 0.71 + 0.005 T1Z2 4.5 + 0.03 74 + 0.5 7.88 + 0.06 0.84 + 0.006 T1Z3 5.1 + 0.03 88 + 0.6 8.79 + 0.07 0.46 + 0.004 T2Z1 2.8 + 0.02 116 + 0.9 7.35 + 0.06 0.73 + 0.006 T2Z2 3.8 + 0.02 72 + 0.6 7.65 + 0.06 0.32 + 0.003 T2Z3 5.7 + 0.03 91 + 0.07 7.90 + 0.07 0.91 + 0.007 T3Z1 3.1 + 0.02 81 + 0.07 7.81 + 0.07 0.66 + 0.004 T3Z2 4.1 + 0.02 98 + 0.09 8.05 + 0.06 0.23 + 0.001 T3Z3 5.9 + 0.03 153 + 1.1 8.20 + 0.07 0.19 + 0.001 Data analysis Correspondence analysis was performed to evaluate differences in the qualitative composition of transects and of the zones within them (for a total of nine samples: Three transects*three zones in each transect). The output of the analysis was coordinates of rows (transect*zone) and columns (species) on correspondence axes superimposed on the scatter diagram. Spearman's rank test was used to test relations between the distance from the sea and number of individuals per species. Statistical relevance of differences between transect pairs was verified with a test for the difference between two proportions. Bray-Curtis cluster analysis based on root-root transformed quantitative data for trans-ect*zone samples was performed to establish levels of similarity between samples. Statistical analysis was performed using the statistical packages Statistica 5 for Windows and BioDiversity Pro (McAlecee et al. 1997). Canonical correlation analysis (CCA) was performed to explore linear relationships between two sets of variables (species abundance and substrate characteristics). The results of the CCA were plotted as a two-dimensional graph with species as points and substrate variables as arrows originating from the center of the graph. Canonical correlation analysis (CCA) was performed using the SAS 9.1 package (SAS, SAS Institute, Gary, NC). Results Plots A total of 26 species from the associations Cakilo-Xanthietum italici and Echinophoro-Elymetum fared Xerohalophytic plant distribution along sand beach 667 were recorded in eight plots. It is evident that the abundance of species of the association Cakilo-Xanthietum italici decreases with increasing distance from the sea, while the abundance of species of the association Echinophoro-Elymetum farcti increases with increasing distance from the sea (Table III). Transects Qualitative composition of transects. A total of 10 species of the associations Cakilo-Xanthietum italici and Echinophoro-Elymetum farcti were analyzed for their distribution along the transects (other recorded species were too few to participate in further analyses). Four of these species were members of Cakilo-Xanthietum italici ass.: Xanthium italicum Mor., Cakile maritima Scop., Salsola kali L., and Euphorbia peplis L. and six were members of Echinophoro-Elymetum farcti ass.: Euphorbia paralias L., Eryngium maritimum L., Elymus farctus (Viv.) Runemark ex Melderis, Echinophora spinosa L., Ammophila arenaria (L.) Link, and Pancratium maritimum L. Ten, nine, and six species were recorded in the first, second, and third transect, respectively. Differences in the qualitative composition of transects and of the zones within them are illustrated in Figure 3. Two groups of samples can be distinguished along the second correspondence axis. The first group contains all three zones of the third transect e (III-1, III-2, III-3) and the second transect-second zone sample (II-2). The second group contains the first and second zone of the first transect (1-1, 1-2) and the first zone of the second transect (II-l). The number of species increases with increasing distance from the sea in transects I and II (four species in zone 1, five species in zone 2 for both transects I and II, 10 species in zone 3 of transect I, and nine species in zone 3 of transect II). Transect III is specific because there is no increase in number of species along zones (five species in zones 1 and 2 and three species in zone 3). Quantitative composition of transects. Spearman's rank test showed a statistically significant positive correlation between the distance from the sea and the number of individuals per species for most analyzed species in transects I and II (Table IV).Statistically relevant differences in the quantitative composition of transects are presented in Table V. In transect I, the largest number of individuals have E. farctus (38.45%) and X. italicum (31.00%), in transect II X. italicum (68.80%) and E. farctus (22.77%), and in transect III X. italicum (55.06%) and E. peplis (29.96%). Better insight into the quantitative composition was obtained when the transects were subdivided into zones (Table VI). X. italicum out- numbered all species in seven samples (zones), E. farctus had the largest number of individuals in zone 3 of transect I (40.35%) and E. peplis in zone 1 of transect III (61.36%). Similarities and dissimilarities between samples (transect*zone) obtained by cluster analysis of the quantitative data set (Figure 4) are compatible with the results of the correspondence analysis based on the qualitative data set (see above). Relationship of species' abundance and substrate characteristics The first canonical correlation axis was the only significant axis and accounted for 80% of the species' abundance/substrate characteristics relationship. The strongest correlations were between the first axis and pH (Figure 5). Other substrate characteristics (electrical conductivity EC, N content, water content) showed weaker correlations along the graph axes. Discussion Plots Spatial succession of the two analyzed plant communities occurred, with an overlapping zone where species of both communities were established. Characteristic species of the association Cakilo-Xanthietum italici became less abundant, whereas the species of the association Echinophoro-Elymetum farcti became more frequent with increasing distance from the sea. Generally, there are two difficulties for vegetation development on mobile sand dunes (Li et al. 1997): (1) the extremely barren soil limiting plant growth and (2) the unstable substrate (wind erosion) which prevents the propagules of most plants from becoming established. Thus, only specific ecophysiological adaptations (e.g. salt and drought tolerance) and reproductive features (e.g. mode of seed dispersal) enable the establishment of particular species on these sites. Annual species with an extensive root system, which are more tolerant to sea spray and seawater inundations (ass. Cakilo-Xanthietum italici), are common in the zone nearest the sea (Barbour & Dejong 1977; Maun 1994). The dispersion of these species in deeper landward zones is attributable to wind action (van der Valk 1974) or specific seed features aiding animal dispersal (e.g. X. italicum which is present far inland). A gradual transition to the other association (i.e. from Cakilo-Xanthietum italici to Echinophoro-Elymetum farcti) is noticeable by the increasing number of Elymus farcti (already at 50 m). The tussocks of this grass species are the main agents of dune evolution since these "immobilize" sand movements, and become a Downloaded by [York University Libraries] at 02:38 17 November 2014 Os 00 Table III. Species abundance in eight plots. Plot code 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Distance from sea (m) 30 50 50 60 70 70 80 80 80 80 90 90 100 100 120 120 120 120 Size of releve (m2) 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 Coverage (%) 15 40 40 30 45 45 40 35 40 40 60 60 65 65 60 60 60 60 Characteristic species of ass. Xanthium italicum Mor. 5 5 5 7 7 7 5 5 5 5 3 3 5 5 3 3 3 3 Cakilo-Xanthietum italici Cakile maritima Scop. 2 2 2 1 Characteristic species of class Cakiletea Salsola kali L. Euphorbia peplis L. Polygonum maritimum L. Artiplex hastata L. 1 1 1 2 2 2 1 1 1 1 Characteristic species of ass. Elymus faraus (Viv.) Runemark. 4 5 3 4 5 7 7 7 7 8 8 8 8 8 8 8 8 Echinophoro-Elymetum fared Echinophora spinosa L. 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 Characteristic species of Eryngium maritimum L. 1 2 2 1 3 3 2 3 2 2 1 1 class Ammophiletea Euphorbia paralias L. Medicago marina L. 1 1 1 1 2 2 2 2 2 1 1 1 1 1 1 Others Ammophila arenaria (L.) Link. Pancratium maritimum L. Calystegia soldanella (L.) R. Br. Pseudorlaya pumila (L.) Grande Aegilops ovata L. Blackstonia perfoliata (L.) Huds. Bromus tectorum L. Cuscuta sp. L. /7?w/tz crithmoides L. Juncus maritimus Lam. Parapholis incurva (L.) C. E. Hubb. Reichardia picroides (L.) Roth. Schoenus nigricans L. Vulpia ciliata Dumort. Lagurus ovatus L. Xerohalophytic plant distribution along sand beach 669 nucleus for further accretion and eventual dune formation (Harris & Davy 1986). Finally, the full establishment of Ammophilla arenaria (after 100 m) indicates a significant decrease of sand salinity. This species has a significant role in the development and stabilization of sand dune systems on European coasts (Salisbury 1952; Ranwell 1972; Deshmukh 1977), but it is usually present only on broader beaches (Alegro et al. 2004). Qualitative composition of transects. When the most numerous species from the total census were analyzed separately for their presence in particular transects/zones, certain effects due to length of transects were observed. Thus, specimens of E. farctus and A. arenaria were present only in 1-3 and II-3 samples, whereas P. maritimum was registered only in 1-3. In the grouping of samples along the second correspondence axis, it could be noted that in transect III, zonation of species was almost absent. Moreover, the species present in transect III were 11 ' □ in g • S. kali X. italicum « species □ transect*zone C. maritima A. arenaria n Tn O E. farctus p^timum n □ ' E, maritimum IIB m E. spinosa*a mi □ III2 E. paralias • E. peplis • -0.5 0 0.5 1.0 1.5 D1M1 Figure 3. CA ordination graph of transects (I, II, III, the first, second, and third transect, respectively) and zones within (1, 2, 3, the first, second, and third zone, respectively). similar to those of sample II-2. On the other hand, samples 1-1, 1-2, and II-1 were grouped. The segregation of these samples was due to the presence or absence of C. maritima. Apparently, this species is more invasive landwards along the continuously rising terrain of transect I. Quantitatively, the most frequent species were X. italicum and E. maritimum (present in nine and seven samples, respectively, and not affected by transect length or topography). Transects I and II are characterized by the gradual shift from species belonging to one association to species from the other association: In zone 1 species of the ass. Cakilo-Xanthietum italici prevail, while zone 2 has a similar number of species of both associations; in zone 3 species of the ass. Echino-phoro-Elymetum farcti prevail. Transect III is once again distinct from the others because there is no such obvious shift in species composition. Quantitative composition of transects. In general, the number of species increased with increasing distance from the sea in transects I and II, but not in transect III. Also, the number of individuals per species increased according to the same pattern (in transects I and II, but not in III). There were several exceptions to this rule (S. kali in transects I and II; C. maritima and E. peplis in transect II). The pattern of increasing number of individuals with increasing distance from the sea is applicable for the cumulative number of individuals of both associations for transects I and II as well. In transect III, the rule fits for ass. Echinophoro-Elymetum farcti, but not for ass. Cakilo-Xanthietum italici. The better soil properties landwards, especially in zone 3, favored an enhanced establishment of most species, which resulted in an increase of their number. This regularity is confirmed by the positive correlation between the distance from the sea and the number of individuals for each species. Table IV. Spearman correlation between distance from the sea and number if individuals per species. Transect I Transect II Transect III Species Spearman R P Spearman R P Spearman R P X. italicum 0.85 *** 0.76 *** 0.71 C. maritima 0.65 ** 0.23 ns / S. kali -0.13 ns -0.09 ns -0.30 E. peplis 0.12 ns 0.03 ns 0.09 E. paralias 0.40 * 0.54 ** 0.05 E. maritimum 0.68 *** 0.61 *** 0.59 E. farctus 0.78 *** 0.62 *** / E. spinosa 0.78 *** 0.48 ** 0.72 A. arenaria 0.13 ns 0.39 * / P. maritimum 0.40 * / / / *P < 0.05; **P < 0.01; ***P < 0.001; ns - non-significant. 670 A. Mijovič et al. Table V. Quantitative composition of transects with statistical relevance of difference between transect pairs. Transect I Transect II Transect III I/II I/III II/III Species A. N. % A. N. % A. N. % P P P X. italicum 954 31.00 955 68.80 147 55.06 *** *** *** C. maritima 419 13.62 5 0.36 0 0.00 *** *** ns S. kali 8 0.26 6 0.43 3 1.12 ns * ns E. peplis 3 0.10 4 0.29 80 29.96 ns *** *** E. paralias 91 2.96 20 1.44 5 1.87 ** ns ns E. maritimum 206 6.69 16 1.15 8 3.00 *** * * E. farctus 1183 38.45 316 22.77 0 0.00 *** *** *** E. spinosa 191 6.21 54 3.89 24 8.99 ** ns *** A. arenaria 8 0.26 12 0.86 0 0.00 ** ns ns P. maritimum 14 0.45 0 0.00 0 0.00 * ns ns A.N. - absolute number of specimens; *P < 0.05; **P < 0.01; ***P < 0.001; ns - non- ■significant. Table VI. Quantitative composition of zones within transects. Species A.n. Zl (%) A.n. Z2 (%) A.n. Z3 (%) Transect I X. italicum 14 51.85 52 44.07 888 30.29 C. maritima 10 37.04 29 24.58 380 12.96 S. kali 2 7.41 3 2.54 3 0.10 E. peplis 0 0.00 0 0.00 3 0.10 E. paralias 0 0.00 2 1.69 89 3.04 E. maritimum 1 3.70 32 27.12 173 5.90 E. farctus. 0 0.00 0 0.00 1183 40.35 E. spinosa 0 0.00 0 0.00 191 6.51 A. arenaria 0 0.00 0 0.00 8 0.27 P. maritimum 0 0.00 0 0.00 14 0.48 Transect II X. italicum 8 53.33 89 82.41 858 67.83 C. maritima 2 13.33 0 0.00 3 0.24 S. kali 2 13.33 3 2.78 1 0.08 E. peplis 0 0.00 2 1.85 2 0.16 E. paralias 0 0.00 0 0.00 20 1.58 E. maritimum 0 0.00 1 0.93 15 1.19 E. farctus 0 0.00 0 0.00 316 24.98 E. spinosa 3 20.00 13 12.04 38 3.00 A. arenaria 0 0.00 0 0.00 12 0.95 P. maritimum 0 0.00 0 0.00 0 0.00 Transect III X. italicum 12 27.27 113 62.43 22 52.38 C. maritima 0 0.00 0 0.00 0 0.00 S. kali 3 6.82 0 0.00 0 0.00 E. peplis 27 61.36 53 29.28 0 0.00 E. paralias 1 2.27 4 2.21 0 0.00 E. maritimum 0 0.00 1 0.55 7 16.67 E. farctus 0 0.00 0 0.00 0 0.00 E. spinosa 1 2.27 10 5.52 13 30.95 A. arenaria 0 0.00 0 0.00 0 0.00 P. maritimum 0 0.00 0 0.00 0 0.00 A.n. - absolute number of specimens. Zl - zone 1; Z2 - zone 2; Z3 - zone 3. The transects were significantly different considering the number of individuals of analyzed species. Thus, the largest differences in quantitative composition were between the transects I and II, somewhat smaller, but still large differences were between the transects I and III. When transects were divided into zones, more subtle differences were observed. X. italicum was very numerous in all nine samples (zones) but outnumber all species in seven samples (zones). E. farctus had the largest number of Xerohalophytic plant distribution along sand beach 671 individuals in 1-3 sample and E. peplis in III-l sample. When the cluster analysis based on quantitative data set was applied, we confirmed the previously presented results of the correspondence analysis based on quantitative data set. The morphology of the terrain at the sand beaches and the physico-chemical conditions of substrate are the most important factors determining the presence, distribution, number and abundance of plant species (Alvarez-Rogel et al. 2000, 2007; Jafari et al. 2004; Lortie & Cushman 2007; Bornman et al. 2008). Soil characteristics estimated in this study were used for establishing the relationship between these variables II-; I- 3 II- 1 ■11-3 «1-2 Ii-: ■n 1-2 1-1 0. % Similarity 50. 100 Figure 4. Bray-Curtis dendrogram of transect*zone samples (I, II, III, the first, the second, and the third transect, respectively; 1, 2, 3, the first, the second, and the third zone, respectively). and species abundance in CCA analysis. These characteristics only partially reflect the conditions at certain microsites, but still we can deduce some conclusions comparable with previous findings. Species' abundances were mostly affected with pH values, the number of specimens increased with increasing pH (third transect is excluded from this conclusion, because of lack of correlation between the abundances and pH values). Soil water regime and EC affected the abundance of few species (A. arenaria and X. italicum), whereas soil nitrogen content had the least impact to the plants' abundance. High heterogeneity of estimated soil characteristics, as a consequence of constant human activity and disturbance, has resulted in a poor correlation between the soil variables and species' abundances. Overall, our results show that two factors affect the plant distribution and species abundance along the shoreline. One is the distance from the sea, and we confirmed the gradual distribution of species, and the more or less regular shift of plant communities with increasing distance from the sea, as previously shown in earlier studies (Pakeman & Lee 1991; Acosta et al. 2007; He et al. 2007). Another is the set of soil characteristics, which was highly heterogeneous and, in relation to the distance from the sea, particularly irregular. Similar results were obtained from the studies on plant distribution along coastal dunes (Alvarez-Rogel et al. 2006, 2007, Dasti et al. 2010), where it has been shown that the soil type and vegetation positioning were connected with terrain P. maritimum C. maritima s *E. maritimum E. paralias . E. spinosa E. farctus Figure 5. CCA ordination diagram showing the distribution of 10 most abundant species and the environmental variables. N - nitrogen content, EC - electrical conductivity. 672 A. Mijovic et al. configuration, regardless of the distance to the shoreline. Lortie and Cushman (2007) also showed that wind is an important factor affecting abiotic factors (soil moisture, N content) along the sand dune shoreline. The fluctuating environment of coastal foredunes, characterized by a variety of geomorphological features, soil properties, and environmental factors (Seelinger 1992), causes specific vegetation composition. Generally, foredune vegetation is dominated by few species, which are limited to beaches and coastal foredunes, and which show a wide geographical distribution, with a number of many subordinate species with a more limited distribution (van der Maarel & van der Maarel-Versluys 1996). Foredune morphology may play a key role for certain species by offering temporary refuges where more stable populations survive, becoming the source for future propagules (Davy & Figueroa 1993). Variable topography implies differences in edaphic factors, and consequently affects the presence, number and relative abundance of species (Jafari et al. 2004, Alvarez-Rogel et al. 2006), and relatively small differences in elevation may produce large differences in species composition (Barrett 2006). The alternations in dune morphology are particularly important in rare or heavily disturbed communities, where these may cause changes in coastal vegetation, even the disappearance of some species/communities (Acosta et al. 2007; Cutini et al. 2010). Adaptations to a broad array of micro-environmental factors are demonstrated in a number of structural and functional plant characteristics, making them recognizable as specific functional types (Garcia-Mora et al. 1999). Low competitive pressure at sand shorelines, accompanied by eutrophication and human pressure, allows the colonization of alien species of suitable functional types, which may, consequently, alter community composition (Stanis-ci et al. 2010). In summary, the xerohalophytic species belonging to the associations Cakilo-Xanthietum italici and Echinophoro-Elymetum farcti were more or less regularly zonated at the Velika plaza beach, following the spatial succession of species common for this type of habitat. With regard to the different micro-topography along the investigated transects, significant differences in number of individuals of plant species among these transects were observed. However, the floristic composition was relatively uniform between transects. This indicates that terrain configuration significantly affects the ability of present species to establish and propagate. X. italicum, which is a representative of the plant association typically distributed in draft zones, showed the widest distribution along the investigated beach, indicating the great ecological amplitude and dispersal advan- tages of this species. Strong anthropogenic pressure on touristically attractive beaches of the southern Adriatic foreland contributes to changing sand micro-topography, thereby disturbing the vegetation communities occurring on them. For the purposes of conservation, sand sieving and gardening of gradually arising dunes may improve the establishment of plants over the entire beach width. Acknowledgment This work was supported by the Ministry of Science, Republic of Serbia, Grant #173011. References Acosta A, Ercole S, Stanisci A, De Patta Pillar V, Blasi C. 2007. Coastal vegetation zonation and dune morphology in some Mediterranean ecosystems. J Coastal Res 236: 1518-1524. Alegro A, Biljakovic M, Bogdanovic S, Borsic I. 2004. Psammo-halophytic vegetation on the largest sand area on the Croatian cost: The island of Mljet, southern Adriatic. Biologia (Bratislava) 59(4): 435-445. Alvarez-Rogel J, Alcaraz F, Ortriz R. 2000. Soil salinity and moisture gradients and plant zonation in Mediterrenean salt marshes of Southeast Spain. Wetlands 20: 357-372. Alvarez-Rogel J, Carrasco L, Marin CM, Martinez-Sanchez JJ. 2007. Soil of a dune coastal salt marsh system in relation to groundwater level, microtopography and vegetation under a semiarid Mediterranean climate in SE Spain. Catena 69: 111-121. Alvarez-Rogel J, Martinez-Sanchez JJ, Blazquez LC, Marin Semitiel CM. 2006. A conceptual model of salt marsh plant distribution in coastal dunes of Southeastern Spain. Wetlands 26(3): 703-717. Barbour MG, De Jong TM. 1977. Response of West Coast beach taxa to salt spray, seawater inundation and soil salinity. B Torrey Bot Club 104: 29-34. Barbour MG, De Jong TM, Pvlik BM. 1985. Marine beach and dune plant communities. In: Chabot BF, Mooney HA, editors. Physiological ecology of North American plant communities. New York: Chapman and Hill. pp. 296-322. Barrett G. 2006. Vegetation communities on the shores of a salt lake in semi-arid Western Australia. J Arid Environ 67: 77-89. Bornman TG, Adams JB, Bate GC. 2008. Environmental factors controlling the vegetation zonation patterns and distribution of vegetation types in the Olifants Estuary, South Africa. S Afr J Bot 74: 685-695. Caniglia G. 1982. La vegetazione del litorale del lido di Volano (Ferrara). Bollettino del Museo Civico di Storia Naturale di Venezia 33: 37-61. Clark JS. 1986. Coastal forest tree populations in changing environment, SE Long Island, New York. Ecol Monogr 5: 97-126. Costa CSB, Cordazzo CV, Seelinger U. 1996. Shore disturbance and dune plant distribution. J Coastal Res 12: 133-140. Cutini M, Agostinelli E, Acosta TRA, Molina JA. 2010. Coastal salt-marsh zonation in Tyrrhenian central Italy and its relationship with other Mediterranean wetlands. Plant Biosyst 144: 1-11. Dasti AA, Saima S, Mahmood Z, Athar M, Gohar S. 2010. Vegetation zonation along the geomorphological gradient at Eastern slope of Sulaiman range, Pakistan. Afr J Biotechnol 9(37): 6105-6115. Xerohalophytic plant distribution along sand beach 673 Davy AJ, Figueroa ME. 1993. The colonization of strandlines. In: Miles J, Walton DWH, editors. Primary succession on land. Oxford: Blackwell. pp. 113-131. Deshmukh JK. 1977. Fixation accumulation and release of energy by Ammophila arenaria in a sand dune succession. In: Jefferies RL, Davy AJ, editors. Ecological processes in coastal environments in a sand dune succession. London: Blackwell Scientific Publications, pp. 353-362. Domac R. 1984. Mala flora Hrvatske i susjednih područja. Zagreb: Školská knjiga. Garcia-Mora MR, Gallego-Fernández JB, García-Novo F. 1999. Plant functional types in coastal foredunes in relation to environmental stress and disturbance. J Veg Sci 10: 27-34. Harris D, Davy AJ. 1986. Strandline colonization by Elymus farctus in relation to sand mobility and rabbit grazing. J Ecol 74(4): 1045-1056. He MZ, Zheng JG, Li XR, Qian XL. 2007. Environmental factors affecting vegetation composition in the Alxa Plateau, China. J Arid Environ 69: 473-489. Hesp PA. 1991. Ecological processes and plant adaptations on coastal dunes. J Arid Environ 21(2): 165-191. Horvatic S. 1974. Contribution to study of psamophytic vegetation of Eastern-Adriatic forecast. IV Congress of Biologists of Yugoslavia. Sarajevo, pp. 39-40. Jafari M, Zare Chahouki MA, Tavili A, Azarnivand H, Zahedi Amiri Gh. 2004. Effective environmental factors in the distribution of vegetation types in Poshtkouh rangelands of Yazd Province (Iran). J Arid Environ 56: 627-641. Jankovič MM, Stevanovič V. 1983. Contribution to study of marshy vegetation of Boka Kotorská. Povremena izdanja muzeja grada Šibenika 10: 377-396. Kumler ML. 1997. Critical environmental factors in dry coastal ecosystems. In: van der Maarel E, editor. Dry coastal ecosystems. General aspects. Amsterdam: Elsevier, pp. 387- 409. Li SG, Zhao AF, Chang XL. 1997. Several problems about vegetation succession of Horquin Sandy Land. J Desert Res 17(1): 25-32. Lortie JC, Cushman JH. 2007. Effects of a directional abiotic gradient on plant community dynamics and invasion in a coastal dune system. J Ecol 95: 468-481. Markham R. 1942. A steam distillation apparatus suitable for micro-Kjeldahl analysis. Biochem J 36(10-12): 790-791. Maun MA. 1994. Adaptations enhancing survival and establishment of seedlings on coastal dune systems. Vegetatio 111: 59-70. McAlecee N, Lambshead PJD, Paterson GLJ, Gage LG. 1997. Biodiversity Professional. Beta-Version. London: The Natural History Museum and The Scottish Association for Marine Sciences. Mijovic A. 1997. Ecological and phytoceonological chacteristics of vegetation of South Montenegrinian foreland (Dissertation). Faculty of Natural Sciences, University of Belgrade, Belgrade (in Serbian). Mijovic S, Stevanovič V. 1988. Phytocoenological characteristics of vegetation of Southern-Montenegrian forecast. IV Congress of Ecologists of Yugoslavia. Ohrid. pp. 344-345. Pakeman RJ, Lee JA. 1991. The ecology of the strandline annuals Cakile maritime and Salsola kali. I. Environmental factors affecting plant performance. J Ecol 79: 143-153. Pignatti S. 1959. Richerche sull'ecologie e sul populamento della dune del litoral di Venezia. Bollettino del Museo Civico di Storia Naturale di Venezia 12: 61-142. Pulevic V. 1973. Contribution to flora of Montenegro. Glasnik republičkog zavoda za záštitu prirode i prirodnjačkog muzeja u Titogradu 6: 77-83. Pulevič V. 1982. Protection of flora and vegetation in Montenegro. MacedAcad Sci Arts (MANU) 1: 109-124. Randall RE, Scott GAM. 1997. Communities of sand and shingle beaches. In: van der Maarel E, editor. Dry coastal ecosystems. General aspects. Amsterdam: Elsevier, pp. 263-274. Ranwell DS. 1972. Ecology of salt marshes and sand sunes. London: Chapman and Hall. Rhoades JD, Chanduvi F, Lesch S. 1999. Soil salinity assessment: Methods and interpretation of electrical conductivity measurements. Rome: Food and Agriculture Organization of the United Nations. Rozema J, Bijwaard P, Prast G,. 1985. Ecophysiological adaptations of coastal halophytes from foredunes and salt marshes. Vegatatio 62: 499-521. Salisbury EJ. 1952. Downs and dunes. London: Bell. Seelinger U. 1992. Coastal foredunes of southern Brazil: Physiography, habitats and vegetation. In: Seelinger U, editor. Coastal plant communities of Latin America. New York: Academic Press, pp. 367-381. Stanisci A, Acosta ATR, Di Iorio A, Vergalito M. 2010. Leaf and root trait variability of alien and native species along Adriatic coastal dunes (Italy). Plant Biosyst 144: 47-52. Tutin TG, Heywood VH, Burges NA, Valentine DH, Walters SM, Webb DA, editors. 1964a. Flora Europea 1. Cambridge: Cambridge University Press, p. 77, 97, 105, 343. Tutin TG, Heywood VH, Burges NA, Moore DM, Valentine DH, Walters SM, et al., editors. 1964b. Flora Europea 2. Cambridge: Cambridge University Press, p. 156, 216, 225, 322, 324, 375. Tutin TG, Heywood VH, Burges NA, Moore DM, Valentine DH, Walters SM, et al., editors. 1972. Flora Europea 3. Cambridge: Cambridge University Press, p. 56, 74, 78. Tutin TG, Heywood VH, Burges NA, Moore DM, Valentine DH, Walters SM, et al., editors. 1976. Flora Europea 4. Cambridge: Cambridge University Press, p. 136, 143, 325. Tutin TG, Heywood VH, Burges NA, Moore DM, Valentine DH, Walters SM, et al., editors. 1980. Flora Europea 5. Cambridge: Cambridge University Press, p. 84, 104, 156, 184, 197, 201, 225, 236, 243, 289. van der Maarel E, van der Maarel-Versluys M. 1996. Distribution and conservation status of littoral vascular plant species along the European coasts. J Coastal Coserv 2: 73-92. van der Valk AG. 1974. Environmental factors controlling the distribution of forbs on coastal foredunes in Cape Hatteras National Seashore. Can J Botany 52: 1057-1073. Westhoff V, van der Maarel E. 1973. The Braun-Blanquet approach. In: Whittaker RH, editor. Handbook of vegetation science 5. The Hague: Junk. pp. 616-726.