Plant diversity along coastal dunes of the Black Sea
(North of Turkey)
Senay Ulu Agır1•Hamdi Guray Kutbay1• Burak Surmen2
Received: 19 March 2015 / Accepted: 10 December 2015 / Published online: 8 January 2016 Ó Accademia Nazionale dei Lincei 2015
Abstract Coastal dunes are particular ecosystems because they include different vegetation types and have high environmental heterogeneity. Plant biodiversity in coastal dunes vary along the seashore to inland environ-mental gradient. In the present study, plant biodiversity along the coastal dune zonation of Central Black Sea Region of Turkey (upper beach or drift line, embryonic or primary dune, main dune, transitional and fixed dune zone) is evaluated using different diversity indexes (Evenness, Functional diversity, Plant biodiversity and Rarity). Plant biodiversity increases from seashore to the inland. In par-ticular evenness and rarity indexes tend to increase from the drift line to fixed dunes as functional diversity is high in the upper beach and drift line.
Keywords Central Black Sea region Coastal dunes Evenness Functional diversity Plant biodiversity Rarity index Zonation
1 Introduction
Coastal dune ecosystems, characterized by highly hetero-geneous environmental mosaics, host an extremely spe-cialized flora and high environmental heterogeneity (Acosta et al.2005,2009a; Carranza et al.2008; Tomaselli et al.2011; Altay and Ozturk2012). Plant communities in coastal dunes are disposed in a complex arrangement along seashore–inland gradient and this leads to high values of biodiversity (Prisco et al.2012). Coastal zonation includes upper beach or drift line, embryonic dune, main dune, transitional and fixed dune zones along seashore–inland gradient (Monserrat et al. 2012).
However, coastal dune vegetation is severely disturbed all over the world due to anthropogenic hazards mainly coastal urbanization (Honrado et al.2010; Stancheva et al.
2011; Malavasi et al.2014a,b; Drius et al.2013). Coastal dunes are highly dynamic systems and thus prone to changes even under natural conditions and often could drive to an advanced state of degradation, irreversible alteration, or lost. Although many studies have focused on the analysis of coastal plant communities on various parts of the world (Doing 1985; Martı´nez and Psuty2004) little is known regarding patterns of spatial variability in species diversity on the Black Sea coastal dunes.
Different plant communities have been identified along the sea–inland gradient, namely related to the upper beach or drift line, to embryonic dunes, main dune, transitional and fixed dunes. These communities constitute the coastal vegetation zonation and could be found in well preserved dune ecosystems.
Plant biodiversity may also be defined as functional diversity. Plant diversity ranges from low to moderately high, but the plant community is functionally redundant, with taxonomically different species in coastal dune & Burak Surmen
buraksurmen@gmail.com Senay Ulu Agır senay.ulu@hotmail.com Hamdi Guray Kutbay hguray@omu.edu.tr
1 Department of Biology, Faculty of Arts and Sciences, University of Ondokuz Mayıs, 55139 Kurupelit-Samsun, Turkey
2 Department of Biology, Kamil Ozdag Science Faculty, Karamanoglu Mehmetbey University, 70200 Karaman, Turkey
ecosystems. Plant functional types (PFTs) approach has been used to describe the responses of coastal dune plants to environmental stress and disturbance (Gallego-Ferna´n-dez and Martı´nez 2011). A functional approach has also been used by Carboni et al. (2011) for coastal dune plants against environmental stress and disturbance. Plant func-tional diversity may be decreased due to harsh conditions (i.e. salinity) in coastal dunes. However, it may be increased along seashore–inland gradient due to less sub-strate mobility.
The aim of this study is to investigate the plant diversity and evenness in coastal dunes in Central Black Sea Region in the north of Turkey along seashore–inland gradient. We used several entropy based biodiversity indexes that mea-sures specific aspects of the partition of abundance among landscape elements (Acosta et al.2005; Drius et al.2013). Beta diversity was also analyzed in order to describe the turnover along seashore to inland gradient. PFTs were also defined along seashore–inland gradient and plant func-tional diversity and rarity index were also calculated. The objectives of this study were: (1) to define whether plant diversity and evenness changed along seashore–inland gradient and (2) to evaluate the changes in functional diversity using two main components namely functional richness (FR) and functional composition (FC) along sea-shore–inland gradient.
2 Materials and methods
2.1 Study areaThe studied coastal dunes covers a 149 km area and includes coastal dunes in western and eastern part of Central Black Sea Region of Turkey between Alacam (41°2902700N and 36°3301200E) and Terme (41°1505200N and
36°5705600E) provinces (Fig.1). Seven localities were
chosen and these localities include characteristic dune zones [upper beach or drift line (A), embryonic or primary dune (B), main dune (C), transitional (D) and fixed dune (E) zones]. The main characteristics of these localities were shown (Table1; Fig.2).
2.2 Vegetation sampling and analysis
Seven vegetation plots were chosen from each locality and each zone. Plot size was determined by minimal area method. 4 9 4 m2 plots were collected from each of the communities of the vegetation zonation: upper beach or drift line, embryonic dune, main dune, transition and fixed dune zones from homogenous places on April–September 2012 (Braun-Blanquet 1964). After registering the com-plete list of the vascular plants occurring inside each plot, cover data of species (in % following Braun-Blanquet1964
scale) were also registered. For the analysis, the Braun-Blanquet scale was transformed on % values as proposed by van der Maarel (1979). The cover-abundance symbols of the Braun-Blanquet scale (r, ?, 1, 2, 3, 4 and 5) were replaced by values according to van der Maarel: 1, 2, 3, 5, 7, 8, and 9, respectively (Yalcin et al.2004).
The Shannon diversity index has been widely used to determine alpha diversity in ecological studies (Magurran
2004). However, Shannon diversity index may fails to highlight major changes in heavily disturbed ecosystems like coastal dunes (De Luca et al. 2011). To solve this problem a new index called Hdune0 index was developed by Grunewald and Schubert (2007). It has been found that Hdune0 was more useful in extreme habitats such as coastal dunes (Attorre et al.2013). H0duneindex was calculated the following formula;
Hdune0 ¼ XPidune In Pð iduneÞ
Fig. 1 Studied localities in coastal dunes in Central Black Sea Region of Turkey
Table 1 Main properties of studied localities Locality Dune zones P/ A M/ AL Characteristic species Main threatening factors for biological diversity Cobanyatag ı (41 °21 005 00N–36 °97 020 00E) Upper beach/drift line P M Salsola ruthenica Three zones were mixed due to wave action Embryonic/primary P M S. ruthenica and Xanthium strumarium subsp. cavanillesii Main P M X. strumarium subsp. cavanillesii and Eryngium maritimum Transitional P A L Pancratium maritimum , Daucus broteri , Cyperus capitatus Fixed dune P A L Eleagnus rhamnoides subsp. rhamnoides Sindel (41 °36 065 00N– 36 °70 010 00E) Upper beach/drift line P A L E. maritimum , Achillea maritima subsp . maritima P. pinaster subsp. pinaster plantations and agricultural activities Embryonic/primary P A L P. maritimum and A. maritima subsp . maritima, X. strumarium subsp. cavanillesii Main P A L Euphorbia paralias Transitional P A L P. maritimum and E.paralias Fixed dune P A L Elymus elongatus subsp. elongatus Hurriyet (41 °19 092 00N– 36 °72 066 00E) Upper beach/drift line P A L S. ruthenica Stabilized dune zone was lost due to tourism activities Embryonic/primary P A L P. maritimum and E. paralias Main P M Sophora alopecuroides var. alopecuroides , E. rhamnoides subsp. rhamnoides Transitional P M S. alopecuroides var. alopecuroides , E. rhamnoides subsp. rhamnoides Fixed dune A – – Costal (41 °15 040 00N– 36 °31 010 00E) Upper beach/drift line P M P. maritimum Three zones were mixed due to wave action and dense tourism activities in transitional dune zone and stabilized dune zone was lost Embryonic/primary P M E. maritimum and Cionura erecta Main P M E. maritimum and Cionura erecta Transitional P A L D. broteri and E. rhamnoides subsp. rhamnoides Fixed dune A – – Cernek (41 °45 000 00N– 36 °07 030 00E) Upper beach/drift line P A L S. ruthenica and E. paralias Inundation was occurred from seashore to primary dune zone. Many of the zones were narrowed due to sand extraction Embryonic/primary P A L P. maritimum and E. paralias Main P A L E. maritimum Transitional P A L S. alopecuroides var. alopecuroides Fixed dune P A L Euphorbia peplis and E. rhamnoides subsp. rhamnoides Sahilkent (41 °68 061 00 N– 35 °85 017 00E) Upper beach/drift line P A L Calystegia soldanella Sand extraction Embryonic/primary P A L A. maritima subsp . maritima , P. maritimum and Ammophila arenaria subsp. arundinacea Main P A L Jurinea kilaea and C. capitatus Transitional P A L E. peplis and D. broteri Fixed dune P A L E. rhamnoides subsp. rhamnoides and E. peplis
where Pidune is the coverage percentage and of the ith
species relative to the plot size.
Hdunemax0 is maximum value of Hdune0 for combination of
total cover and number of species (Grunewald and
Schu-bert 2007). Hdunemax0 index was calculated the following
formula; Hdunemax0 ¼ s Pp idune s In Pp idune s
s the number of species in a plot.
Edune is ratio between H0dune and Hdunemax0 (Grunewald and Schubert2007) Edunewas calculated as:
Edune ¼ Hdune0 =H 0 dunemax
Species diversity and evenness were also calculated by the Simpson and Berger–Parker indexes. Simpson diversity and evenness was calculated the following formulas (Smith and Wilson1996):
D¼ 1X
s
i¼1
p2i
where s is the total number of species and piis the relative cover of ith species
E1=D¼ ð1=DÞ=S
where E1/D, evenness value according to Simpson’s index; D, Simpson’s diversity index; S, number of species; pi, relative cover of the ith species.
Berger–Parker index (d) was calculated the following formula (Berger and Parker1970);
d¼ Nmax=N
where d Berger–Parker index; N is total number of species in a plot, and, Nmaxis the number of individuals of the most abundant species. Reciprocal of d values were used (Magurran2004).
Rarity index was also calculated using the following formula;
R¼ P
jIjkðN FjÞ=N
Sk
where Ijkis the incidence of species j in plot k, Fjis total number of plots containing species j, N is the total number of plots and Sk is the number of species in the plot k (Colwell and Coddington1994).
Plant diversity and evenness in studied swamp forests were calculated by Biodiversity 2.0 version software (McAleece et al. 1997). Rarefaction method has been widely used for estimating standardized species richness (Santoro et al.2012). Rarefaction curves were drawn using Biodiversity 2.0 version software (McAleece et al. 1997; Hsieh and Li1998). Table 1 continued Locality Dune zones P/ A M/ AL Characteristic species Main threatening factors for biological diversity Doyran (41 °63 000 00N– 35 °69 050 00E) Upper beach/drift line P A L E. maritimum and E. paralias Dense grazing especially in transitional and stabilized dune zones Embryonic/primary P A L Juncus littoralis, P. maritimum Main P A L E. maritimum and Cynodon dactylon var. dactylon Transitional P A L P. maritimum, D. broterii , J. kilaea Fixed dune P A L E. rhamnoides subsp. rhamnoides and J. kilea P present, A absent, M mixed, AL alone
EðSnÞ ¼ Xs i¼1 1 N Ni n . N n
where n is the expected number of species, Ni is the number of species belonging to ith species, and N is the total number of species.
Beta diversity is defined as spatial heterogeneity or pattern diversity was calculated using the Whittaker for-mula (Whittaker 1960; Magurran 2004; Luis-Calabuig et al.2006; Gulsoy and Ozkan2008).
b¼S a 1 Fig. 2 Profile diagrams of studied localities
where S is the total number of species, a is the mean species richness.
Functional richness (FR) and functional diversity FC were also calculated because PFTs approach has been used to describe the responses of coastal dune plants to environ-mental stress and disturbance (Carboni et al.2011; Gallego-Ferna´ndez and Martı´nez2011). FR was defined as the total number of plant functional PFTs per plot. FC is the presence and relative abundance of PFTs per plot (Gallego-Ferna´ndez and Martı´nez2011). Salinity resistance, vegetative repro-duction, growth form and life span were selected to define PFTs. Salinity resistance (S) and vegetative reproduction (V) was evaluated with binary values (S: 1/present, 0/absent; V: 1/present, 0/absent). For growth form (G) triplet scale were used (G: herbaceous:1/shrub/2; tree/3). Life span (L) was also evaluated with binary values (1/annual; 2/perenial) (Gallego-Ferna´ndez and Martı´nez2011).
340 plots 9 4 functional traits matrix was subjected to TWINSPAN and four groups to identify clusters of species with similar traits, which were assumed to represent functional types at the species level (Keddy1992).
Taxonomic nomenclature followed that of Guner et al. (2012).
3 Results
Diversity tends to increase along seashore to inland gra-dient. Hdune0 values, Berger–Parker and Simpson along the coastal zonation in the different studied localities are
reported in Table2. Hdune0 values of transitional dune zones had the highest plant diversity index (Table3). The results of Berger–Parker are coherent with Hdune0 index. The highest Simpson indexes were found in transitional zone. Eduneand E1/Devenness indexes were increased from drift line to fixed dune zones (Table4).
Statistically significant differences were found among dune zones with respect to FR and FC. The highest FC and FR values were found in transitional and fixed dune zones, respectively (Table4).
Rarefaction curves showed that the lowest diversity was found in upper beach or drift line, while main dune and transitional dune zones had the highest diversity (Fig.3).
The highest rarity index values were found in transi-tional and fixed dune zones, respectively (Table5). Beta diversity was followed similar trend and the highest beta diversity was found in transitional and fixed dune zones, respectively (Table6).
Four groups were separated according to TWINSPAN analysis based on functional traits. The first and second separations were formed according to vegetative repro-duction and salinity resistance, respectively. Salinity-re-sistant species were grouped in drift-line. Both drift-line and fixed dune species have vegetative reproduction in the study area. The third and fourth separations were formed according to life span. Drift-line and dune grasslands include many annual species. The number of shrubs and trees were rather low, and the study area includes many herb species (Fig.4).
Table 2 H0
duneindex (adapted from Shannon index), Berger–Parker (d) and Simpson (D) plant biodiversity in coastal dune zones Zones Upper beach or drift line Embryonic or primary dune Main dune Transitional dune Fixed dune H0
dune 1.216 ± 0.059c 1.294 ± 0.081bc 1.494 ± 0.064ab 1.564 ± 0.051ab 1.638 ± 0.025a d 3.329 ± 0.204c 3.816 ± 0.181bc 4.048 ± 0.260bc 4.646 ± 0.269ab 5.222 ± 0.501a D 0.136 ± 0.014a 0.109 ± 0.008ab 0.099 ± 0.009ab 0.082 ± 0.003b 0.068 ± 0.005b Different lowercase letters indicate significant differences among coastal dune zones
Table 3 Edune(adapted from Shannon’s evenness), E1/D(Simpson’s evenness) in coastal dune zones
Zones Upper beach or drift line Embryonic or primary dune Main dune Transitional dune Fixed dune Edune 0.650 ± 0.017a 0.644 ± 0.022a 0.672 ± 0.024a 0.674 ± 0.010a 0.673 ± 0.005a E1/D 9.198 ± 1.183b 10.581 ± 0.822b 12.133 ± 1.319b 13.357 ± 0.524b 18.709 ± 1.925a Different lowercase letters indicate significant differences among coastal dune zones
Table 4 FR and FC values of studied localities
Zones Upper beach or drift line Embryonic or primary dune Main dune Transitional dune Fixed dune FC 1.430 ± 0.047c 1.559 ± 0.033bc 1.539 ± 0.034bc 1.717 ± 0.035a 1.667 ± 0.058ab FR 3.552 ± 0.111c 3.824 ± 0.079c 3.873 ± 0.087bc 4.200 ± 0.093ab 4.307 ± 0.128a Different lowercase letters indicate significant differences among coastal dune zones
4 Discussion
As reported on previous research performed on Mediter-ranean coastal dunes (Acosta et al. 2009a, b), species diversity on Black Sea district tends to increase with the distance from shoreline. Berger–Parker index values were increasing tendency from drift line to transitional zone thus dominance of a particular species were decreased (Zagh-loul2008). It has been stated that species diversity was low in drift line because only certain species have adapted to high sand burial and the salt spray due to high wind velocity, and with a higher sediment supply to the foredune (da Silva et al.2008). Acosta et al. (2009b) and Carboni et al. (2009) also stated that only highly specialized
vascular plant species can survive along drift line with because sand burial, high salt content due to salt spray and unstable substrate. Lowest species richness is expected in exposed areas near the shoreline (Nyle´n and Luoto 2014). Primary dune (or embryonic) zone had also low plant biodiversity and this zone is usually characterized by Am-mophila arenaria (L.) Link subsp. arundinacea H. Lindb. Fil. in the study area. Isermann (2011) found that coastal dunes with Ammophila arenaria (L.) Link subsp. arundi-nacea H. Lindb. Fil. had the lowest plant biodiversity. This species has the capacity to survive deep sand burial and the presence of this species is an important factor in impeding the movement of sand due to sea winds (Attorre et al.2013; Agir et al. 2014). Latorre et al. (2013) found that embry-onic dunes had the lowest richness, while high richness values were found with distance from the shoreline. Plant density and diversity gradually increases with distance from the shoreline until a dune ridge vegetation complex has evolved, virtually halting any further inland movement of sediment and resulting in the formation of a large pre-cipitation ridge (Abuodha et al. 2003). Primary dunes are also characterized by Achillea maritima (L.) Ehrend. et Y., (=Otanthus maritimus Hoffmans. et Link). It has been reported that Achillea maritima (L.) Ehrend. et Y., (= Otanthus maritimus Hoffmans. et Link) is a marker of foredune stability because of its sensitivity to the changes in the dune structure and this species improved habitat conditions for rare species (Honrado et al.2010; Agır et al.
2014). This may be attributed to the high disturbance in drift line regarding tourism activities in the study area.
It has been found that plant diversity has increasing tendency in transitional dune zone. Coastal dune species developed special adaptation strategies against relatively harsh conditions and away from shoreline habitat condi-tions allowed the establishment of several plant commu-nities (Cakan et al. 2011; A´ lvarez-Molina et al. 2012; Huseyinova et al.2013). Outside the drift line, coastal dune plant communities tend to be permanent and less exposed Fig. 3 Rarefaction diagrams of
studied localities along seashore inland gradient
Table 5 Rarity index values in studied localities
Zones Rarity index
Upper beach or drift line 0.357 ± 0.033a Embryonic or primary dune 0.383 ± 0.039a
Main dune 0.404 ± 0.042a
Transitional dune 0.446 ± 0.044a
Fixed dune 0.438 ± 0.076a
Different lowercase letters indicate significant differences among coastal dune zones
Table 6 Beta diversity in studied localities
Zones b diversity
Upper beach or drift line 1.259 ± 0.249a Embryonic or primary dune 1.165 ± 0.123a
Main dune 1.193 ± 0.116a
Transitional dune 1.436 ± 0.127a
Fixed dune 1.406 ± 0.265a
Different lowercase letters indicate significant differences among coastal dune zones
to harsh conditions and biological diversity gradually increased (Maun2009; Acosta et al.2007,2009b; Attorre et al.2013).
Evenness was tend to increased from drift line to fixed dunes in other localities in the present study. This can be interpreted by the distribution of communities in transitional and fixed dune zones were more even than the distribution of communities in drift line. This is probably due to a balanced species composition in these zones (Acosta et al.2009a,b). The highest rarity index values were found in transi-tional and fixed dune zones in the study area and this means these zones include more exclusive species (i.e. Euphorbia terracina L., Jurinea kilea Azn.) as compared to the other zones (Acosta et al.2009b). In the study area, inundations occurred. It has been known that inundation has a pro-nounced regulatory effect on the distribution and abun-dance of plant species (Deegan and Harrington 2004).
Acosta et al. (2009b) also stated that drift line showed higher values of rarity index, whereas this zone had the lowest number of plant species. This zone has extreme maritime conditions and only highly specialized vascular plant species can survive here. However, we had not found high rarity index values in drift line probably due to dense tourism activities in this zone.
The highest beta diversity was found in transitional dune zone. This zone subjected to threatening factors such as agricultural and tourism activities and inundation usually occurred and such factors lead to environmental hetero-geneity. Beta diversity is usually driven by environmental heterogeneity especially in local scale (El-Ghareeb et al.
2006; Kallimanis et al.2008; Isermann2011; Rickert et al.
2012) and this heterogeneity leads to distinct local varia-tions i.e. topographical variavaria-tions (Shaltout et al. 1995,
We found that D (transitional) and E (fixed dune) zones were the highest FC. Gallego-Ferna´ndez and Martı´nez (2011) stated that stable dunes had larger relative presence and abundance of functional groups (FC). Functional diversity indices differed significantly between primary dunes and stable dunes and primary dunes had the lowest values in terms of the occurrence of different functional groups per plot (FR). Likewise, species richness and diversity, as well as functional diversity, were lowest on primary dunes.
Vegetative reproduction indicates exposure to stress or disturbance intensity. Similarly, growth form indicated the response to disturbance events and both traits may be used a tolerance mechanism for recurring disturbances (Novo et al. 2004; Gallego-Ferna´ndez and Martı´nez 2011). Vegetative reproduction protects drift-line species (i.e. Euphorbia paralias L.) from high salt concentration, unstable substrate and wave effect, while protects fixed dune-species (i.e. Cyperus capitatus Vandelli, Sophora alopecurioides L. var. alopecurioides) against the negative effects of inundation (Gallego-Ferna´ndez and Martı´nez
2011; Agır et al. 2014). Drift-line and fixed dune zones include many herbaceous species in the study area (i.e. Lagurus ovatus L.), and these zones subject to several disturbance factors (i.e., agriculture and tourism, trampling, construction of houses and roads, waste disposal, and plantation of trees and shrubs) (Agır et al. 2014). It has been stated that annual species are better able to thrive after recurring disturbances than perennial species (Castillo and Moreno-Casasola1998; De Luca et al.2011).
In conclusion, plant diversity and evenness was increased from the seashore to inland gradient in the studied coastal dunes. High beta diversity was found in inland zones in some localities, whereas drift line had the highest beta diversity in some localities most probably due to local differences. Rarity index found to be higher in transitional zone in most of the studied localities. Coastal dune species in Central Black Sea Region are subjected to disturbance factors (i.e. wave action, dense tourism activ-ities, sand extraction, etc.). Extreme physical stress and disturbance factors act shaping community zonation even at very small spatial scales in coastal dune ecosystems (Carboni et al. 2010). Sustainable management pro-grammes in coastal sand dunes should be included the conservation of species poor-habitats containing unique or endangered species elements (Acosta et al.2009b).
Acknowledgments We are grateful to University of Ondokuz Mayıs Research Fund (PYO.FEN.1904.10.010) for supporting this study financially.
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