Biologia 66/6: 10271043, 2011 Section Botany DOI: 10.

2478/s11756-011-0113-3

Plant communities structure and composition in three coastal wetlands in southern Apulia (Italy)
Valeria Tomaselli1 *, Romeo Di Pietro3 & Saverio Sciandrello2
1 2

C.N.R., Institute of Plant Genetics, via G.Amendola 165/A, I-70126 Bari, Italy; e-mail: valeria.tomaselli@igv.cnr.it Department of Botany, University of Catania, via A. Longo 18, I-95125 Catania, Italy; e-mail: sciandrello.s@hotmail.it 3 Department DATA, University of Rome La Sapienza, via Flaminia 70, I-00196 Rome, Italy; e-mail: romeo.dipietro@uniroma1.it

Abstract: In this paper the results of a study on the composition and the distribution of the plant communities in three coastal areas of southern Apulia are presented. A total of about 180 vegetation relevs were performed following the BraunBlanquet phytosociological method. Vegetation data were analysed using both classication (UPGMA, similarity ratio) and ordination methods (including Non-metric Multidimensional Scaling (NMS) and Detrended Correspondence Analysis (DCA). The relevs are distributed in the following classes: Molinio-Arrhenateretea, Phragmito-Magnocaricetea, Juncetea maritimi, Sarcocornietea fruticosae, Saginetea maritimae, Thero-Salicornietea, Helianthemetea guttati. Detailed information about structure and zoning of the detected plant communities are here provided. Two new associations, belonging to the Alkanno-Maresion nanae alliance (microphytic ephemeral plant communities growing on sandy soils, Helianthemetea guttati class) have been described here, both in the Torre Guaceto site. The area of Le Cesine showed the highest total number of plant communities, while the Saline di Punta della Contessa site revealed the largest number of Sarcocornietea fruticosae plant communities. Key words: Apulia; coastal wetlands; cluster analysis; ordination; salt marshes; vegetation; southern Italy

Introduction Coastal wetlands, with their habitats of salt marsh vegetation, are distributed throughout the Mediterranean area (Ghu 1999). In addition to hosting very rare plant species and communities, the coastal wetlands are important areas for the nesting and the staging of birds. In the Adriatic region, the salt marsh vegetation prevails in the western and northern coasts, the east coast being mostly rocky (Panda et al. 2007). Within the Italian side of the Adriatic Sea, the salt marsh vegetation has been the object of various phytosociological studies (Pignatti 1966; Corbetta 1968; Ghu et al. 1984; Biondi et al. 1989; Ghu & Biondi 1996; Piccoli et al. 1996; Biondi 1999; Poldini et al. 1999; Pirone et al. 2001). Nevertheless, a systematic study on the vegetation of these areas is still lacking, and the only phytosociological data available to date come from a few contributions that are often limited to very small areas (Corbetta 1970; Macchia & Vita 1973; Taetani & Biondi 1989; Mariotti et al. 1992; Beccarisi et al. 2003a,b; Biondi et al. 2006). It is wellknown that coastal wetlands are habitats of great biological importance and high taxonomical richness, but, at the same time, they are extremely vulnerable to human activities and disturbances and, for this rea* Corresponding author

son they require constant protection. Their monitoring and safeguard is one of the priorities for a suitable management of coastal environments. In recent decades, these habitats have been subject to intense human pressure, largely due to tourist activities and the subsequent building and infrastructure proliferation and to the development of intensive agriculture. Moreover, the creation of industrial areas has led to the devastation of highly valuable coastal areas. Having a comprehensive knowledge framework on these areas and their conservation status is an essential startingpoint for the planning of future measures of protection. The present study is a part of an INTERREG project III A Italy-Greece 20002006 (Development of an integrated informative system for the monitoring and the management of Natura 2000 protected areas in Geece and in Italy), aimed at the monitoring and the management of common ecosystems. The study is focused on three of the most signicant coastal humid areas of Apulia region: Torre Guaceto, Saline di Punta della Contessa and Le Cesine, on the Adriatic side of southern Apulia. Composition of the vegetation occurring in these three coastal areas is here analyzed, focusing particularly on plant communities of salt marshes and their zoning.

c 2011 Institute of Botany, Slovak Academy of Sciences

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Fig. 1. Study area.

Fig. 2. Cluster analysis of halophilous and hygrophilous perennial vegetation.

Material and methods Study area (Fig. 1) Torre Guaceto is a Ramsar site, a Marine Protected Area and a National Natural Reserve. The Natural Reserve covers an area of about 1.200 ha. According to the EU Directives Habitat and Bird, this site is a proposed Site of Community Interest (pSCI) (IT9140005) and a Special Protection Area (SPA) (IT9140008). The coastal strip is characterized by both rocky and sandy shores. Owing to drainage works and partial silting in the past century to create new

areas for agriculture, the coastal lagoons occupy only small surfaces. The main risk factors are coastal erosion and re, which periodically aicts the protected area. Saline di Punta della Contessa (LS) is a Regional Natural Park which covers an area of 1960 ha. It is also a pSCI (IT9140003) and a SPA (IT9140003). Most of the protected area is occupied by farmlands, while the wet area lies between the cultivated groves and the sandy coastline. The retrodunal coastal area consists of a system of interconnected ponds and marshes which are characterised by several halophytic communities. The main risk factor is the

Plant communities in coastal wetlands in southern Apulia

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Fig. 3. NMS ordination analysis of halophilous and hygrophilous perennial vegetation Axis 1 explains a part of the variance (r squared 0.384; the total of explained variance is of 0.745) and is related mainly to a soil salinity gradient. The distribution along the axis 2 and the axis 3 does not seem to reveal any particularly dened correlation (r squared values are, respectively, 0.219 and 0.142).

presence of some industrial factories which produce high levels of pollutants in waters and soils. Marine erosion has caused a progressive reduction of the sandbank, leading to the increasing salinity in the retrodunal lagoons. Le Cesine (LC) is a Ramsar site and a National Natural Reserve covering about 350 ha. It is also a pSCI (IT9150032) and SPA (IT9150014). The coastal strip consists almost entirely of sandy shores. The retrodunal humid area is one of the most important in southern Italy and it is formed by two large water pools and various channels, marshes, and wet grasslands. The main risk factors are coastal erosion

and re. In recent decades, marine erosion has caused a progressive reduction of the sandbank, causing a slight but progressive increase in salt rates of the coastal lagoons.

Vegetation sampling and analysis The vegetation analysis was carried out following the z urichmontpellier phytosociological method (Braun-Blanquet 1964). Phytosociological relevs (180 relevs) have been performed in the period April 2007October 2009 and mapped using GPS. The size of plots varies from a maximum of 100

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Fig. 4. DCA of the data-set of sub-halophilous helophytic communities and grasslands. The r squared values of axes 1 and 2 are, respectively, 0.324 and 0.204 and are related to an increasing soil salinity and to a decreasing ooding period.

m2 to 5 m2 , depending on vegetation type and microtopography. The nomenclature of plant species follows Conti et al. (2005). A specic data base has been implemented with coordinates, list of plant species, cover values, geographical and topographic features of each phytosociological relev. The phytosociological relevs have been arranged in two different matrix: halophilous-higrophilous perennial vegetation (112 plots 154 species) and therophytic vegetation (both xerophilous and hygrophilous, 96 plots 180 species). Rare species were not excluded from the analysis. For each dataset, similarity analyses of the relevs were carried out using the SYN-TAX 2000 software (Podani 2001). Original Braun-Blanquet sampling scale has been transformed into the ordinal scale according to Van der Maarel (1979). A hierarchic classication method (UPGMA) was performed. Dissimilarity of the relevs was measured using the Simi-

larity ratio. Clusters of relevs were classied into syntaxa according to Rivas-Martinez et al. (2001, 2002). We referred to Perez Prieto & Font (2005) for the Helianthemetea guttati class. The ordination of the data-sets was performed using the PC-ORD 4.34 software. In the ordination analyses we ran both Nonmetric Multidimensional Scaling (NMS) and Detrended Correspondence Analysis (DCA), based on the Euclidean distance. The two methods gave consistent ordination diagrams. NMS is a non-parametric ordination technique based on the ranked compositional dissimilarities among sites, and therefore has advantages over parametric ordination techniques in which the underlying assumptions are rarely satised in eld community data (Clarke 1993). The autopilot routine in the NMS program of PC-ORD showed that a three axes NMS was appropriate, though we have chosen to

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Fig. 5. DCA of the data-set of the halophilous perennial vegetation. Along the axis 1 (r squared, 0.437) a good dened ecological gradient related to the soil salinity and along the axis 2 a gradient of increasing granulometry (sand percentage).

display only two axes where there is a greater variance. Detrended Correspondence Analysis (DCA) (Hill & Gauch 1980) was used to establish ecological patterns in the scattergram and to generate hypotheses about vegetation-environment relationships. The data set of the halo-hygrophilous perennial vegetation has been divided into two separate arrays in order to highlight the ecological pattern: in the rst one only the halophilous, scrubby and helophytic communities (63 plots 66 species) were included, while the second one included the rest of the plant communities (non-halophilous helophytic communities and grasslands, 49 plots 99 species).

Results Halophilous and hygrophilous perennial vegetation Cluster analysis identies two main clusters, A and B (Fig. 2), separating respectively the halophilous (A) from the sub-halophilous (B) communities. In cluster A, three sub-clusters are identied: A1, A2, A3. Cluster A1 includes the chamaephytic halophilous vegetation characterized by succulent chenopodiaceous dwarf shrubs (Sarcocornietea fruticosae class, Sarco-

cornietalia fruticosae order). Cluster A2 includes the helophytic grasslands of salt marshes (Juncetea maritimi class), while cluster A3 includes the helophytic subhalophilous vegetation of Limonio-Juncetum gerardi (Juncetea maritimi) and Scirpetum maritimae (Phragmito-Magnocaricetea). Cluster B includes the helophytic communities of Phragmito-Magnocaricetea (sub-cluster B1), and the mesophilous Molinio-Arrhenateretea communities (sub-cluster B2). The NMS (Fig. 3) shows a fair signicance only along the axis 1 and it is related mainly to a soil salinity gradient. The group of Phragmition (in red) and Paspalo-Polypogonion viridis (in yellow), which occupy the left part of the diagram, are clearly separate from the halophilous communities of Sarcocornietea fruticosae and of Juncetea maritimi (in black, in the right part of the graph). The DCA analysis claries the ecological gradients of the two main groups of relevs. The DCA of the data-set of nonhalophilous/sub-halophilous communities allows identifying fairly good dened ecological gradients along axes 1 and 2 (Fig. 4) related, respectively, to increasing soil salinity and a decreasing ood period. In this case, the

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Fig. 6. Cluster analysis of therophytic vegetation.

halophilous group of Scirpion communities (in the right part of the graph) is well segregated from the Phragmition and the Paspalo-Polypogonion viridis communities (in the left part). The DCA of the data-set of the halophilous communities allows identifying a welldened ecological gradient along the axis 1 related to decreasing soil salinity (Fig. 5): the Sarcocornietea fruticosae communities are arranged in the graph, from left to right, from the Arthrocnemum macrostachyum groups (higher level of soil salinity) to the Limbardia crithmoides groups (medium level of soil salinity). In the right part, the Juncetea maritimi relevs fall in two main groups: the Juncion maritimi sandy communities (top right) and the Plantaginion crassifoliae comm. (bottom right), according to a gradient of increasing granulometry recognizable along axis 2. Therophytic vegetation The cluster analysis forms two main groups, A (temporary inundated soils) and B (dry and pioneer soils) (Fig. 6). Cluster A is divided into two subclusters, A1 and A2. Cluster A1 includes the ephemeral halophilous Saginetea maritimae communities which developes in temporary inundated soils. A1 is further divided into A1.1 and A1.2. The sub cluster A1.1 includes the Parapholis liformis halo-subnitrophilous communities and sub cluster A1.2 groups microphytic halophilous communities characterized by Isolepis cernua. Cluster A2 groups the glasswort pioneer halophilous communities of temporary inundated salt marshes, belonging to Thero-Salicornietea class. Cluster A2 is divided into two sub clusters: A2.1, including the halo-

subnitrophilous communities (Thero-Suaedetalia) and A2.2 including the halophytic communities (TheroSalicornietalia). The Thero-Suaedetalia group includes the communities dominated by Suaeda maritima and Cressa cretica, while the Thero-Salicornietalia group includes Salicornia patula and Bassia hirsuta communities. Cluster B is divided in two sub-clusters. Subcluster B1 includes the Malcolmietalia therophytic plant communities and is divided into two further sub-clusters: B1.1 (Anthemis tomentosa and Rostraria litorea communities) and B1.2 (Plantago albicans communities and Romulea rolii communities). Anthemis tomentosa and Rostraria litorea communities are here described as two new associations, respectively AlkannoPlantaginetum albicantis and Rostrario litoreae-Anthemidetum tomentosae. Sub-cluster B2 includes those therophytic communities characterized by the dominance of Ornithopus compressus (Helianthemietalia order). The NMS have not shown any signicant dierences among the three axes: only the axis 3 seems to be related to a coarse gradient of salinity (Fig. 7). The DCA analysis allowed identifying some welldened ecological gradients which follow Axis 1 and which are related to both an increasing soil salinity and moisture and to a decreasing soil content in sand. Fig. 8 shows how the identied communities are arranged, from left to right, along a gradient which ranges from the psammophilous communities of the dunes (Malcolmietalia) on the left side of the diagram, to the halophilous communi-

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Fig. 7. NMS ordination analysis of therophytic vegetation doesnt show signicant dierence among the three axes. The r squared values of axes 1, 2 and 3 are, respectively, 0.286, 0.276 and 0.293, so it is not easy to identify the precise ecological correlations.

ties of salt marshes (Saginetea maritimae and TheroSalicornietea. Discussion Plant communities of salt marshes are spatially arranged on the basis of their microecological characteristics, especially soil salinity and moisture (Boisset 1985; Rogel et al. 2000; Molina et al. 2003). Soil salinity gradually increases with soil elevation, reaching a maximum just above mean high sea level (MHSL) (Sanchez et al. 1996; Adam 1990; Bockelmann et al. 2002). Above the MHSL, the salinity tends to decrease due to progressively less frequent ooding (Adam 1990). Nevertheless, this relationship between soil salinity and elevation only partially explains the vegetation zoning, since physiological responses of plants to salinity are heavily species dependent and respond to a combination of biotic and abiotic factors (Silvestri et al. 2004).

Perennial salt-marsh communities, characterized by succulent chenopodiaceous dwarf shrubs, are included in Sarcocornietea fruticosae class and Sarcocornietalia order, with several alliances: Sarcocornion alpini, salt-marshes submerged by tides for long periods (Puccinellio-Sarcocornietum alpini, Sarcocornia alpini comm.); Sarcocornion fruticosae, saltworks and wet salt-marshes submerged for short periods (JuncoSarcocornietum fruticosae); Arthrocnemion glauci, occurring on salt-marshes with very-salted soils (Arthrocnemo-Juncetum subulati) (Rivas-Martinez et al. 2001); Inulion crithmoidis (Agropyro scirpei-Inuletum crithmoidis and Limonio-Artemisietum coerulescentis), typical of the higher levels occasionally reached by tides and on well-drained and relatively salted soils (Brullo & Furnari 1976; Frondoni & Iberite 2002). In Table 1, the synoptic table of the six Sarcocornietea fruticosae plant communities detected in the three study sites is reported.

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Fig. 8. DCA ordination analysis of therophytic vegetation, Axis 1 (r squared 0.345), is related to an increasing soil salinity and soil moisture and to a decreasing soil granulometry (% of sand).

Table 1. Synoptic table of Sarcocornietea fruticosae communities. Vegetation unit Nr. of relevs Char. and Di. Ass. Puccinellia festuciformis (Host) Parl. Sarcocornia fruticosa (L.) A.J. Scott Arthrocnemum macrostachyum (Moric.) Moris Elytrigia scirpea (C. Presl.) Holub Limonium narbonense Mill. Artemisia caerulescens L. Char. Sarcocornietea fruticosae Sarcocornia alpini (Lag.) Rivas Martinez Halimione portulacoides (L.) Aellen Juncus subulatus Forssk. Limbarda crithmoides (L.) Dumort. Aeluropus littoralis (Gouan) Parl. Limonium virgatum (Willd.) Fourr. Triglochin bulbosum L. subsp. barrelieri (Loisel.) Rouy 1 10 2 4 3 4 5 4 Percentage Frequency 80 100 80 20 60 . 75 75 100 . 25 . 5 4 6 7

100 40 50 20 90 .

. . . 75 100 .

. 50 . 100 100 .

86 71 29 100 100 100

100 100 80 20 . . .

100 . 25 50 50 . .

80 100 60 40 . . .

25 100 100 . . . .

75 100 50 100 75 50 25

100 57 100 . . .

Vegetation units: 1: Puccinellio-Sarcocornietum alpini, 2: Sarcocornia perennis communities, 3: Junco-Sarcocornietum fruticosae, 4: Arthrocnemo-Juncetum subulati, 5: Agropyro-Inuletum crithmoidis, 6: Limonio narbonensis-Artemisietum coerulescentis

Sarcocornia alpini occurs in all the study areas although in Saline di Punta della Contessa only,

it is particularly widespread together with Puccinellia festuciformis subsp. festuciformis in the form of

Plant communities in coastal wetlands in southern Apulia the association Puccinellio-Sarcocornietum alpini (Table 1, columns 1 and 2). This community is typical of substrates that are ooded for long periods (Frondoni & Iberite 2002) and grows on the lower belt of salt marshes. When both moisture and salinity degree increase, the Sarcocornia alpini communities are replaced by Sarcocornia fruticosa and Artrochnemum macrostachyum communities. In particular, on slightly more elevated sites, which are still submerged in winter, Junco-Sarcocornietum fruticosae occurs (Table 1, column 3). This association is well-adapted to colonize compact salty-clayey soils. In those areas which are ooded for only short periods in winter Arthrocnemum macrostachyum populations become dominant in the form of the association Junco subulati-Arthrocnemetum glauci (Brullo & Furnari 1976; Table 1, column 4) nom. inv. prop. (=Arthrocnemo-Juncetum subulati Brullo & Furnari 1976), developing on soils which generally show the highest degree of salt content among the Sarcocornietea communities. On those stands that are only rarely ooded, Limbardia crithmoidis communities prevail. The Inulion crithmoidis communities show relatively low salt values. Agropyro scirpei-Inuletum crithmoidis grows on those high and peripheral sites which are not subject to submersion in winter (Table 1, column 5). LimonioArtemisietum coerulescentis (Table 1, column 6) is an endemic Illyrian-Apennine association that develops outside of the tidal zone, on low salty substrates which are only sporadically submerged and which are completely dry in spring and summer. It can be considered a spatial transition between the true coastal halophilous vegetation of Sarcocornietea fruticosaeand the vegetation of Juncetea maritimi (Panda et al. 2007). Juncetea maritimae class (Juncetalia maritimi order) includes perennial grasslands and wet communities of salt marshes (Juncion maritimi), as well as the coastal halophilous vegetation typical of well-drained sandy soils (Plantaginion crassifoliae). The synoptic table of the four Juncetea maritimi plant communities detected in the three study sites is shown in Table 2. Spartina versicolor forms thick grasslands which usually tend to occupy the transition belt between the dunes and the alluvial plain (Frondoni & Iberite 2002) on coarse sandy soils and can be referred to Spartino-Juncetum maritimi (Table 2, column 1). The temporary inundated sands characterizing the retrodunal areas are populated by perennial grasslands dominated by Schoenus nigricans and Plantago crassifolia, belonging to Schoeno-Plantaginetum crassifoliae (Table 2, column 2). Generally this plant community develops in the external parts of the brackish slush and behaves as a transition belt between the halophilous vegetation of Sarcocornietea fruticosae and the communities of Juncetea maritimi. In the reserve of Torre Guaceto, also the Plantagini crassifoliae-Caricetum extensae (Table 2, column 3), which was described by Ghu & Biondi (1988) for the southern Apulia, and the LimonioJuncetum gerardi (Table 2, column 4) have been found.

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Table 2. Synoptic table of Juncetea maritimi communities. Vegetation unit Nr. of relevs Char. and Di. Ass. Spartina versicolor Fabre Schoenus nigricans L. Carex extensa Gooden. Juncus gerardii Loisel. 1 2 3 4 15 7 3 4 Percentage Frequency 100 33 7 . . 100 57 . . 100 100 . . . . 100

Char. Juncion maritimi and Juncetea maritimi Juncus acutus L. 73 100 Juncus maritimus Lam. 80 100 Centaurium spicatum (L.) Fritsch 7 43 Carex hispida Willd. 20 . Lotus corniculatus L. . . subsp. preslii (Ten.) Fourn. Sonchus maritimus L. 13 14 Scirpoides holoschoenus (L.) Sojk 7 . Tripolium pannonicum (Jacq.) Dobrocz. . 14 . Samolus valerandi L. . . Juncus subulatus Forssk. 7 . Char. Plantaginion crassifoliae Plantago crassifolia Forssk. Cladium mariscus (L.) Pohl Centaurium tenuiorum (Homanns. Link) Fritsch Ipomea sagittata Poir.

100 100 67 33 100 . 67 33 33 .

75 . . . . . . . . .

7 27 7 7

100 . . .

100 . . .

. . . .

Vegetation units: 1: Spartino-Juncetum maritimi, 2: SchoenoPlantaginetum crassifoliae, 3: Plantagini crassifoliae-Caricetum extensae, 4: Limonio-Juncetum gerardi

These two associations occur in small and scattered patches. All the swampy helophytic communities dominated by perennial graminoids, typical of fresh and brackish waters, belong to the Phragmito-Magnocaricetea. In this vegetation type the following environments were identied: brackish water communities (Scirpetalia compacti, Scirpion compacti: Scirpetum compacti, Scirpetum compacto-littoralis) and fresh water communities (Phragmitetalia, Phragmition: Phragmitetum communis, Iridetum pseudacori, Soncho-Cladietum marisci, Sparganietum erecti). In Table 3, the synoptic table of the ve Phragmito-Magocaricetea plant communities detected in the three study sites is reported. Scirpetum compacti is dominated by Bolboschoenus maritimus (Table 3, column 1), and grows on salt clay ooded soils (dry in summer), along the banks of the coastal lagoons. Scirpetum compacto-littoralis, which is dominated by Schoenoplectus litoralis (Table 3, column 2), forms tall helophyte communities growing on brackish mud ooded for most of the year. Phragmitetum communis occurs in the damp stations submerged for most of the year. Physionomically, it is characterized by Phragmites australis which forms monophytic populations (Table 3, column 3) widely distributed in the three study areas, Phragmites australis being an invasive species which spreads easily after cutting or re (often causing the detriment of other natural

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Table 3. Synoptic table of Phragmito-Magnocaricetea communities. Vegetation unit Nr. of relevs Char. and Di. Ass. Bolboschoenus maritimus (L.) Palla Schoenoplectus litoralis (Schrad.) Palla Phragmites australis (Cav.) Trin. ex Steud. Iris pseudacorus L. Sonchus maritimus L. Char. Phragmito-Magocaricetea Cladium mariscus (L.) Pohl Mentha acquatica L. Cirsium creticum (Lam.) dUrv. subsp. triunfetti (Lacaita) Werner Lythrum salicaria L. Ipomea sagittata Poir. Samolus valerandi L. Carex hispida Willd. Euphorbia hirsuta L. Galium palustre L. subsp. elongatum (C. Presl) Lange Typha angustifolia L. Alisma lanceolatum With. Schoenoplectus tabernaemontani (C.C. Gmel.) Palla Lycopus auropaeus L. Eupatorium cannabinum L. Typha latifolia L. Persicaria decipiens (R.Br.) K.L. Wilson 1 8

V. Tomaselli et al.

2 3 4 3 5 3 Percentage Frequency 100 100 100 . . . . 100 20 . . . 33 100 .

5 14

100 . 25 . .

. . 79 7 86

. . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . .

20 20 40 20 . 40 . . . . . . . . 20 20

100 . 33 . . . 67 . . . 67 67 33 33 . .

100 64 36 50 57 36 29 21 21 14 . . . . . .

Vegetation units: 1: Scirpetum compacti, 2: Scirpetum compacto-litoralis, 3: Phragmitetum communis, 4: Iridetum pseudoacori, 5: Soncho maritimi-Cladietum marisci

Phragmitetea communities, such as Soncho-Cladietum marisci). Especially in Torre Guaceto it covers a wide surface formerly occupied by coastal lagoons and subsequently buried and used for agricultural purposes and nally abandoned. It is likely that the permanence of Phragmites australis could be favoured by re (that periodically occurs in this area). On soils submerged for the most of the year by oligoaline waters, swampy plant communities dominated by Cladium mariscus occur (Table 3, column 5). In the reserve of Le Cesine, Soncho-Cladietum marisci covers a wide surface, while in Torre Guaceto it occurs only in a small surface probably due to competition with Phragmites australis. Iridetum pseudoacori grows mainly along some drainage ditches at the boundary between wetlands and cultivated areas (Table 3, column 4) and forms a typical mosaic with Sparganietum erecti, which requires owing fresh waters. The distribution of Phragmitetea communities is mainly inuenced by the ooding period length. Scirpion communities occur in those areas where the water salinity shows signicant values. The wet meadows developed on deep and moist soils are here represented by dierent types of Mediterranean summer green grasslands growing on damp, meso-eutrophic and base-rich soils (Holoschoenetalia vulgaris, Molinio-Holoschoenion: Schoeno nigricantisErianthetum ravennae) and on eutrophic soils ooded for a large part of the year by fresh waters (PaspaloHeleochloetalia, Paspalo-Polygonion viridis: Loto tenuis-Paspaletum paspaloidis, Hydrocotile vulgaris com-

munities). The synoptic table of the three MolinioArrhenatheretea plant communities detected in the three study sites is shown in Table 4. The SchoenoErianthetum ravennae (Table 4, column 2) often forms a mosaic with the oligohaline communities of Phragmitetea class. The plant communities dominated by Paspalum distichum (Table 4, column 1), developing in late summer-autumn in correspondence of hollows within cultivated areas, are here referred to the Loto-Paspaletum paspaloidis. The presence of hygrophilous species of Phragmito-Magnocaricetea and Isoeto-Nanojuncetea is to be related to the period of ooding, decreasing from the middle part towards the outlying belts. Hydrocotile vulgaris communities (Table 4, column 3) occur on stagnant and oligotrophic waters (Rivas Martinez et al. 2001, 2002) and often form mosaic with the Soncho-Cladietum marisci. Saginetea maritimae class includes ephemeral, halo-subnitrophilous communities occurring on salt marshes during springtime. The synoptic table is in Table 5. In the study areas we have found two dierent associations, Parapholidetum liformis and IsolepidoSaginetum maritimae (Frankenietalia pulverulentae). Parapholidetum liformis (Table 5, column 1) generally colonises sandy soils rich in organic matter, completely dry in summer (Brullo et al. 1994). This ephemeral vegetation occurs in spatial contact with Sarcocornietea fruticosae communities. Isolepido-Saginetum maritimae (Table 5, column 2) usually grows in the form of small-size patches on sandy-silty soils which are ooded

Plant communities in coastal wetlands in southern Apulia

Table 4. Synoptic table of Molinio-Arrhenatheretea communities. Vegetation unit Nr. of relevs Char. and Di. Ass. Lotus tenuis Waldst.& Kit. ex Willd. Erianthus ravennae (L.) P. Beauv. Schoenus nigricans L. Hydrocotyle vulgaris L. Char. Paspalo-Polypogonion viridis Paspalum distichum L. Lythrum junceum Banks & Solander 1 2 3 11 1 3 Percentage Frequency 55 . . . . 100 67 . . . 33 100

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Table 5. Synoptic table of Saginetea maritimae communities. Vegetation unit Nr. of relevs Char. and Di. Ass. Parapholis liformis (Roth) C.E. Hubb. Isolepis cernua (Vahl) Roem. & Schult. 1 2 15 4 Percentage Frequency 100 . 75 100

100 27

33 .

33 .

Char. Molinio-Arrhenatheretea and Holoschoenetalia vulgaris Juncus articulatus L. 100 . 100 Lippia nodiora (L.) Michx. 73 . 67 Pulicaria dysenterica (L.) Bernh. 45 33 100 Plantago major L. 73 . . Rumex conglomeratus Murray 73 . . Cirsium creticum (Lam.) dUrv. . 100 100 subsp. triunfetti (Lacaita) Werner Scirpoides holoschoenus (L.) Sojk 18 67 33 Dorycnium rectum (L.) Ser. . 67 67 Epilobium tetragonum L. 36 . . Kickxia commutata 27 . . (Bernh. ex Rchb.) Fritsch Trifolium fragiferum L. 27 . . Verbena ocinalis L. 27 . . Orchis palustris Jacq. . . 67 Festuca arundinacea Schreb. 18 . . Agrostis stolonifera L. 9 . . Trifolium repens L. 9 . . Potentilla reptans L. . . 33 Vegetation units: 1: Loto tenuis-Paspaletum paspaloidis, 2: Schoeno nigricantis-Erianthetum ravennae, 3: Hydrocotyle vulgaris communities

Char. Saginetea maritimae and Frankenion pulverulentae Juncus hybridus Brot. 60 100 Polypogon maritimus Willd. 73 . Spergularia salina J. & C. Presl 73 . Plantago coronopus L. 40 . Polypogon monspeliensis (L.) Desf. 33 25 Centaurium tenuiorum 20 75 (Homanns. & Link) Fritsch Sagina maritima G. Don . 100 Polypogon subspathaceus Req. . 100 Anagallis arvensis L. subsp. . 75 parviora (Homanns. & Link) Arcang. Gaudinia fragilis (L.) P.Beauv. . 50 Hordeum marinum Huds 13 . Hainardia cylindrica (Willd.) Greuter 13 . Parapholis incurva (L.) C.E. Hubb. 13 . Vegetation units: 1: Parapholidetum liformis, 2: Isolepido cernuae-Saginetum maritimae

Table 6. Synoptic table of Thero-Salicornietea communities. Vegetation unit Nr. of relevs Char. and Di. Ass. Suaeda maritima (L.) Dumort. Cressa cretica L. Char. Thero-Salicornietea Salicornia patula Duval-Jouve Bassia hirsuta (L.) Asch. Salicornia emerici Duval-Jouve Salsola soda L. Trasg. Saginetea maritimae Atriplex prostrata Boucher ex DC Puccinellia convoluta (Hornem.) Hayek Spergularia salina J. & C. Presl Parapholis liformis (Roth) C.E. Hubb. Hordeum marinum Huds Crypsis aculeata (L.) Aiton 1 2 3 4 9 5 6 3 Percentage Frequency 89 . 100 . 100 17 100 100

in winter. Usually it forms a small-scale mosaic with the Juncetea maritimi class communities. The Thero-Salicornietea class, including the pioneer communities rich in glasswort and other succulent plants of salt marshes, usually appearing during the late summer-autumn period, is represented by the two orders Thero-Suadetalia and Thero-Salicornietalia. The synoptic table of the three Thero-Salicornietea plant communities detected in the three study sites is shown in Table 6. Suaedetum maritimae (Table 6, column 3) and Cressetum creticae (Table 6, column 4) belong to Thero-Suadetalia order, which includes halonitrophilous therophytic communities growing usually at the edges of cultivated areas. Suaedo-Salicornietum patulae and Suadeo maritimae-Bassietum hirsutae belong to Thero-Salicornietalia order. SuaedoSalicornietum patulae (Table 6, column 2) occurs in the more depressed areas, in the form of narrow stripes surrounding the water bodies. Suadeo maritimaeBassietum hirsutae (Table 6, column 1) tends to occupy the adjacent external belt, often in close proximity of the perennial halophyte communities of Sarcocornietea fruticosae (Ghu 1992).

100 100 78 11

100 60 . 60

50 17 100 33

67 . . .

22 . . . . .

. . . . . .

83 50 33 33 33 .

33 100 67 33 . 33

Vegetation units: 1: Suaedo maritimae-Bassietum hirtae, 2: Suaedo maritimae-Salicornietum patulae, 3: Suaedetum maritimae, 4: Cressetum creticae

Spring and early summer microphytic ephemeral plant communities belong to the Helianthemetea guttati class. The vegetation growing on sandy soils is referred to the Malcomietalia order and to the AlkannoMalkolmion alliance. The synoptic table is in Table 7. Plantago albicans communities (Table 7, column 1 and Table 8, rel. 18) occur in the retrodunal area of the protected area of Torre Guaceto, and form a mo-

1038
Table 7. Synoptic table of Helianthemetae guttatae communities. Vegetation unit Nr. of relevs Char. and Di. Ass. Plantago albicans L. Romulea rollii Parl. Rostraria litorea (All.) Holub Ornithopus compressus L. 1 2 3 4 8 3 9 7 Percentage Frequency 100 . . . . 100 . . . . 100 . . . . 100

V. Tomaselli et al. lia region. Plantago albicans communities observed in our study site are lacking in Anchusa hybrida, character species of Anchuso hybridae-Plantaginetum albicantis. So, here we propose a new association, AlkannoPlantaginetum albicantis (holotypus: ril. 4, tab. 8) which exhibits a peculiar specic character component when compared to that of Anchuso-Plantaginetum. On the small-size surfaces of the retrodunal areas of Saline di Punta della Contessa the associationVulpioRomuletum rollii has been found. This vegetation type is characterised by the presence of Romulea rollii (Table 7, column 2). In Torre Guaceto, on coarse soils along the rocky coast, a special therophytic vegetation type characterized by the dominance of Anthemis tomentosa and Rostraria litorea (Table 7, column 3 and Table 8, rel. 917) forms a belt above the Limonietum apuli. For this community a new association named Rostrario litoreae-Anthemidetum tomentosae (holotypus: ril. 16, tab. 8) is here proposed. This new association exhibits some similarities in oristic composition with the Medicagini littoralis-Anthemidetum tomentosae (Ghu et al. 1992) association described for the coast of Rhodes island (Euphorbion peplis R.Tx. 1950 alliance, Cakiletea maritimae class). Nevertheless, the two plant communities dier for the ecological (edaphic) characteristics. Ephemeral plant communities grow in the inland areas of Le Cesine, on red soils, characterised by the dominance of Ornithopus pinnatus (Table 7, column 4) and of a group of species belonging to the Helianthemion guttati alliance and to the Helianthemetea guttati class (Rivas-Martnez et al. 2002, 2001; Perez Prieto & Font 2005). The vegetation of the coastal dunes and of rocky shores, belonging to Ammophiletea and Crithmo-Limonietea classes, and the hydrophytic vegetation of Potametea and Ruppietea classes have been observed in the eld but have not been submitted to multivariate analysis. Hence, these plant communities are reported only in the syntaxonomical scheme. Here follows a brief description. The sandy coastline is characterized by the typical zonation of the dune banks and represented by the annual psammophilous vegetation (Cakiletea maritimae: Salsolo-Cakiletum maritimae) and by dunecolonising vegetation (Ammophiletea: Cypero capitatiAgropyertum juncei, Medicagini-Ammophiletum australis). A typical rocky shores occurs in Torre Guaceto only and it is characterized by coastal chasmophytic communities (Crithmo-Limonietea: Crithmo-Limonietum apuli) and by perennial pioneer grasslands (Artemisietea vulgaris, Elytrigion athericae: Elytrigietum athericae). The submerged rooted hydrophytes communities of the deep waters (Rodwell 1995) belong to Potametea (Potametum pectinati, Potametum colorati) and Ruppietea (Entheromorpho-Ruppietum maritimae, Ruppietum spiralis). These plant communities are well developed in Le Cesine site. This study revealed a very high richness and diversity of dierent plant communities in the three sites. The entire syntaxonomic listing is reported in Ta-

Char. Alkanno-Malcolmion and Malcolmetalia Medicago littoralis Loisel. 100 Cerastium diusum Pers. 100 Alkanna tinctoria Tausch 100 Aira cupaniana Guss. . Romulea rollii Parl. 75 Silene niceensis All. 63 Pseudorlaya pumila (L.) Grande 63 Corynephorus divaricatus (Pourr.) . Breister. Brassica tournefortii Gouan 50 Lotus parviorus Desf. . Vulpia fasciculata (Forssk.) Fritsch . Cutandia divaricata (Desf.) Benth. . Erodium laciniatum (Cav.) Willd. . Herniaria hirsuta L. . Char. Helianthemetea guttatae Filago eriocephala Guss. Polycarpon tetraphyllum (L.)L. subsp. diphyllum (Cav.) B` olos & FontQuer Euphorbia exigua L. Anthemis tomentosa L. Hippocrepis ciliata Willd. Arenaria leptoclados (Rchb.) Guss. Trifolium glomeratum L. Linaria pelisseriana (L.) Mill. Trifolium arvense L. Ornithopus pinnatus (Mill.) Druce Tuberaria guttata (L.) Fourr. Vulpia ciliata Dumort. Trifolium nigrescens Viv. Sedum rubens L. Trifolium campestre Schreb. Trifolium scabrum L. Trisetaria segetum (Savi) Soldano Tolpsis umbellata Bertol. Rumex bucephalophorus L. Trifolium angustifolium L. Trifolium bocconei Savi Tuberaria praecox Grosser Filago gallica L.

100 . . . . 33 33 . . . 67 33 33 .

100 67 . . . . . . . . . . . .

86 57 14 100 . . . 71 . 43 . . . 14

63 . 100 . 50 100 . . . . . . . . . . . . 25 . . . .

. 100 . . . . . . . . . . . . . . . . . . . . .

89 78 44 100 . . . . . . . . . 56 . . . . . . . . .

. . . . 71 . 100 100 100 86 71 71 71 . 71 71 71 43 . 29 29 29 29

Vegetation units: 1: Alkanno-Plantaginetum albicantis (Tab. 9, rel. 1-8), 2: Vulpio-Romuletum rollii, 3: Rostrario-Anthemidetum tomentosae (Tab. 9, rel. 9-17), 4: Ornithopus pinnatus and Tuberaria guttata communities

saic with the Thymus capitatus garrigues. This plant community exhibits ecological and structural features which are similar to those shown by Anchuso hybridaePlantaginetum albicantis, association described by Corbetta et al. (1989) for the Ionian side of the Apu-

Plant communities in coastal wetlands in southern Apulia

Table 8. Alkanno-Plantaginetum albicantis and Rostrario-Anthemidetum tomentosae. Alkanno-Plantaginetum albicantis ass. nova (18) Rostrario-Anthemidetum tomentosae ass. nova (917) Relev. Nr. Area (m2 ) Cover (%) Slope ( ) Face Di. Ass. Plantago albicans L. Rostraria litorea (All.) Holub Char. Alkanno-Malcolmion e Malcolmetalia Medicago littoralis Loisel. Cerastium diusum Pers. Alkanna tinctoria Tausch Romulea rollii Parl. Silene niceensis All. Pseudorlaya pumila (L.) Grande Brassica tournefortii Gouan Char. Tuberarietea guttatae Filago eriocephala Guss. Euphorbia exigua L. Anthemis tomentosa L. Arenaria leptoclados (Rchb.) Guss. Polycarpon tetraphyllum (L.) L. subsp. diphyllum (Cav.) O. B` olos & Font Quer Sedum rubens L. Hippocrepis ciliata Willd. Rumex bucephalophorus L. Trasg. Saginetea maritimae Catapodium pauciorum (Merino) Brullo, Giusso, Minissale & Spamp. Plantago coronopus L. Frankenia pulverulenta L. Catapodium balearicum (Willk.) H.Scholz Parapholis incurva (L.) C.E. Hubb. Sagina maritima G. Don Spergularia salina J.& C. Presl 4 . 3 . 4 . 4 . 3 . 3 . 4 . 4 . . 1 . 1 . 2 . 1 . 3 . 1 . 2 . 3 1 4 85 . . 2 4 80 . . 3 4 85 . . 4 3 90 . . 5 3 90 . . 6 5 95 . . 7 10 90 . . 8 10 90 . . 9 4 70 . . 10 5 60 . . 11 5 65 . . 12 5 70 . . 13 8 90 . . 14 5 85 5 NE 15 10 95 5 NE 16 15 95 3 N

1039

17 20 90 . .

. +

2 1 + 1 . + .

2 1 + 1 . . .

1 + 1 2 1 + +

2 2 1 1 + + +

1 1 + 2 . + .

1 1 1 1 + . 1

1 2 + . + . .

1 1 1 . 1 + +

1 1 . . . . .

2 1 . . . . .

2 2 . . . . .

3 + . . . . .

2 + . . . . .

1 . . . . . .

1 + . . . . .

2 . . . . . .

1 . . . . . .

. + . 1 . . . .

. 1 . 1 . . . .

. + . 1 . . . .

1 + . 2 . . + .

+ 1 . + . . . +

+ 1 . 2 . . 1 +

1 + . 1 . . + .

+ + . + . . + .

+ + 2 . + . . .

1 + 1 . . + . .

+ . + . 1 . . .

1 + 1 . 1 + . .

1 . 4 . . + . .

+ . 4 . + . . .

1 . 4 . 1 1 . .

+ + 4 . + + . .

. . 4 . + . . .

. . . . . . .

. . . . . . .

. . . . . . .

. . . . . . .

. . . . . . .

. . . . . . .

. . . . . . .

. . . . . . .

+ 1 + 1 . + .

1 1 1 + + 1 +

1 + 1 3 1 2 +

+ 2 1 1 + 2 +

2 + + 1 1 + .

1 1 1 2 2 + .

1 + + + 2 + .

+ 2 + 1 1 + .

1 . 1 + 2 . .

Rel. 1-8, 11-03-2008, Torre Guaceto Rel. 9, 11-03-2008, Torre Guaceto Rel. 10-16, 19-05-2008, Torre Guaceto Rel. 17, 22-05-2008, Torre Guaceto

ble 9. Several plant communities were described elsewhere but were not known for the Adriatic site of southern Italy and the Apulia region, such as Ruppietum spiralis, Sparganietum erecti, Agropyro scirpeiInuletum crithmoidis, Limonio narbonensis-Juncetum gerardi, Parapholidetum liformis, Cressetum creticae, Elytrigietum athericae (Marchiori & Albano 2007). Two new associations have been described here (Alkanno tinctoriae-Plantaginetum albicantis, Rostrario litoreaeAnthemidetum tomentosae). In Table 10 the list of the plant communities is reported, with the indication of the presence in the three study sites. The area of Le Cesine presents the main number of plant communities and a richness especially in hydrophytic and helophytic vegetation types, an indication also of the good quality of pond environments. The site of Saline di Punta della Contessa revealed the largest number of

Sarcocornietea fruticosae communities, due to the phenomenon of salinization of ponds occurring in the last decades. These highly specic vegetation types identied in the studied area are severely threatened by human activities, in particular the development of tourism, which have led to the transformation of coastal habitats into beaches. The plant communities which normally characterize the transition belt between the halophilous vegetation of Sarcocornietea fruticosae and the helophytic communities of Juncetea maritimi are particularly endangered at present. Typical examples are the associations Limonio-Artemisietum coerulescentis and Agropyro scirpei-Inuletum crithmoidis, which are situated on the borderline between the high and low tide, or Limonio narbonensis-Juncetum gerardi and Plantagini crassifoliae-Caricetum extensae which have been

1040
Table 9. Syntaxonomical scheme of the phytosociological units cited in the text.

V. Tomaselli et al.

POTAMETEA Klika in Klika & Novak 1941 POTAMETALIA Koch 1926 POTAMION (Koch 1926) Libbert 1931 Potametum pectinati Cartensen 1955 NYMPHAEION ALBAE Oberd. 1957 Potametum colorati Allorge 1921 RUPPIETEA J.Tx.1960 RUPPIETALIA J.Tx.1960 RUPPION MARITIMAE Br.-Bl. ex Westho in Bennema, Sissingh & Westho 1943 Ruppietum spiralis Hocquette 1927 corr. Iversen 1934 Enteromorpho intestinalidis-Ruppietum maritimae Westho ex R.Tx. & B ockelmann 1957 PHRAGMITO-MAGNOCARICETEA Klika in Klika & Novk 1941 PHRAGMITETALIA Koch 1926 PHRAGMITION Koch 1926 Phragmitetum communis (Koch 1926) Schmale 1939 Iridetum pseudoacori Krzywanski 1974 Soncho-Cladietum marisci (Br.-Bl.& O.Bol` os 1958) Cirujano 1980 Sparganietum erecti Philippi 1973 SCIRPETALIA COMPACTI Hejny in Holub et al. 1967 corr. Rivas et al. 1980 SCIRPION COMPACTI Dahl & Hadac 1941 corr. Rivas-Martnez et al.1980 Scirpetum compacti Van Langendonck 1931 corr. Bueno & F. Prieto in Bueno 1997 Scirpetum compacto-littoralis (Br.-Bl. in Br.-Bl. et al. 1952) O. Bol` os 1962 corr. Rivas CRITHMO-LIMONIETEA Br.-Bl. in Br- Bl., Roussine & N` egre 1952 CRITHMO-LIMONIETALIA Molinier 1934 CRITHMO-LIMONION MOLINIER 1934 Crithmo-Limonietum apuli Bartolo, Brullo, Signorello 1989 Elytrigietum athericae Corillion 1953 corr. Bueno 1997 AMMOPHILETEA Br.-Bl. & R.Tx. ex Westho et al. 1946 AMMOPHILETALIA Br.-Bl. 1933 AMMOPHILION AUSTRALIS Br.-Bl.1921 em. G` ehu, Rivas-Martnez & R.Tx. in Rivas-Martnez et al. 1980 Medicagini marinae-Ammophiletum australis Br.-Bl. 1921 corr. Prieto & Diaz 1991 AGROPYRENION FARCTI Rivas-Martnez, Costa, Castroviejo & Valdes Bermajo 1980 Cypero capitati-Agropyretum juncei K uhnholtz-Lordat (1923) Br.-Bl. 1933 CAKILETEA MARITIMAE R.Tx. & Preising in Br.-Bl. & R.Tx.1952 CAKILETALIA INTEGRIFOLIAE R.Tx. ex Oberd. 1949 corr. Rivas-Martnez, Costa & Loidi 1992 CAKILION MARITIMAE Pignatti 1953 Salsolo-Cakiletum maritimae Costa & Mansanet 1981 corr. Rivas-Martnez et al. 1992 JUNCETEA MARITIMI Br.Bl. in Br.-Bl., Roussine & N` egre 1952 JUNCETALIA MARITIMI Br.Bl. ex Horvatic 1934 JUNCION MARITIMI Br.Bl. ex Horvatic 1934 Spartino-Juncetum maritimi O. Bol` os 1962 Limonio narbonensis-Juncetum gerardi G` ehu et Biondi 1994 PLANTAGINION CRASSIFOLIAE Br.-Bl.(1931) 1952 Schoeno-Plantaginetum crassifoliae Br.-Bl. in Br.-Bl., Roussine & Negre 1952 Plantagini crassifoliae-Caricetum extensae G` ehu & Biondi. 1988 SARCOCORNIETEA FRUTICOSAE Br.-Bl. & R.Tx. ex A. & O. Bol` os 1950 SARCOCORNIETALIA FRUTICOSAE Br.-Bl.1933 SARCOCORNION ALPINI (Rivas-Martnez et al. 1990) Brullo, Giusso del Galdo, Minissale, Siracusa & Spampinato 2002 (Sarcocornienion alpini Rivas-Martnez, Lousa, T.E. Diaz, Fernandez-Gonzalez & J.C. Costa 1990) Puccinellio-Sarcocornietum alpini Castroviejo & Cirujano 1980 Sarcocornia alpini comm. ARTHROCNEMION GLAUCI Rivas-Martnez in Rivas-Martnez et al. 1980 Junco subulati-Arthrocnemetum glauci Brullo & Furnari 1976 nom. inv. propos. (Arthrocnemo-Juncetum subulati Brullo & Furnari in Not. Fitosoc. 11:1314, 1976) SARCOCORNION FRUTICOSAE Br.-Bl. 1933 em. Brullo & Furnari 1988 Junco-Sarcocornietum fruticosae Brullo, Guarino, Ronsisvalle 2000 INULION CRITHMOIDIS Brullo & Furnari 1988 Agropyro scirpei-Inuletum crithmoidis Brullo in Brullo et al.1988 Limonio-Artemisietum coerulescentis Horvatic (1933)1934 MOLINIO-ARRHENATHERETEA R.Tx.1937 HOLOSCHOENETALIA VULGARIS Br.-Bl. ex Tchou 1948 MOLINIO-HOLOSCHOENION VULGARIS Br.-Bl. ex Tchou 1948 Schoeno nigricantis-Erianthetum ravennae Pignatti 1953 PASPALO-HELEOCHLOETALIA Br.-Bl. in Br.-Bl. et al. 1952 PASPALO-POLYPOGONION VIRIDIS Br.-Bl. in Br.-Bl. et al.1952 Loto tenuis-Pasapletum paspaloidis Biondi, Casavecchia & Radetic 2002 Hydrocotile vulgaris comm. SAGINETEA MARITIMAE Westho, Van Leeuwen & Adriani 1962 FRANKENIETALIA PULVERULENTAE Rivas-Martnez ex Castroviejo & Porta 1976 FRANKENION PULVERULENTAE Rivas-Martnez ex Castroviejo & Porta 1976 Parapholidetum liformis Brullo, Scelsi & Siracusa 1994 Isolepido-Saginetum maritimae Brullo 1988

Plant communities in coastal wetlands in southern Apulia

Table 9. (continued)

1041

THERO-SALICORNIETEA R.Tx. ex Ghu & Ghu-Franck 1984 THERO-SALICORNIETALIA R.Tx. ex Ghu & Ghu-Franck 1984 SALICORNION PATULAE Ghu & Ghu-Franck 1984 Suaedo-Salicornietum patulae Brullo & Furnari ex Ghu & Ghu-Franck 1984 Suaedo maritimae-Bassietum hirtae Br.-Bl. 1928 THERO-SUAEDETALIA Br.-Bl. & O. Bol` os 1958 THERO-SUAEDION Br.-Bl. in Br.-Bl., Roussine & N` egre 1952 Suaedetum maritimae Pignatti 1953 Cressetum creticae Brullo & Furnari 1976 HELIANTHEMETEA GUTTATI (Br.-Bl. in Br.-Bl., Roussine & N` egre 1952) Rivas Goday & Rivas Martnez 1963 em. Rivas Martnez 1978 HELIANTHEMETALIA GUTTATI Br.-Bl. in Br.-Bl., Molinier & Wagner 1940 (Tuberarietalia guttatae Br.-Bl. in Br.-Bl., Molinier & Wagner 1940 nom. mut.) HELIANTHEMION GUTTATI Br.-Bl., in Br.-Bl., Molinier & Wagner 1940 Ornithopus pinnatus and Tuberaria guttata comm. MALCOLMIETALIA Rivas Goday 1958 ALKANNO-MARESION NANAE Rivas Goday ex Rivas Goday & Rivas-Martnez 1963 corr. Diaz-Garretas et al. 2001 (= AlkannoMalcolmion parviorae Rivas Goday 1958) Vulpio-Romuletum rollii Brullo & Scelsi 1998 Alkanno tinctoriae-Plantaginetum albicantis ass. nova Rostrario litoreae-Anthemidetum tomentosae ass. nova

Table 10. Presence of the detected plant communities in the three sites (TG: Torre Guaceto; LS: Saline di Punta della Contessa; LC: Le Cesine). Torre Guaceto Potametum pectinati Potametum colorati Ruppietum spiralis Enteromorpho intestinalidis-Ruppietum maritimae Phragmitetum communis Iridetum pseudoacori Soncho-Cladietum marisci Sparganietum erecti Scirpetum compacti Scirpetum compacto-littoralis Crithmo-Limonietum apuli Elytrigietum athericae Medicagini marinae-Ammophiletum australis Cypero capitati-Agropyretum juncei Salsolo-Cakiletum maritimae Spartino-Juncetum maritimi Limonio narbonensis-Juncetum gerardi Schoeno-Plantaginetum crassifoliae Plantagini crassifoliae-Caricetum extensae Puccinellio-Sarcocornietum alpini Sarcocornia alpini comm. Arthrocnemo-Juncetum subulati Junco-Sarcocornietum fruticosae Agropyro scirpei-Inuletum crithmoidis Limonio-Artemisietum coerulescentis Schoeno nigricantis-Erianthetum ravennae Loto tenuis-Pasapletum paspaloidis Hydrocotile vulgaris comm. Parapholidetum liformis Isolepido-Saginetum maritimae Suaedo-Salicornietum patulae Suaedo maritimae-Bassietum hirtae Suaedetum maritimae Cressetum creticae Ornithopus pinnatus comm. Vulpio-Romuletum rollii Alkanno tinctoriae-Plantaginetum albicantis Rostrario litoreae-Anthemidetum tomentosae X Saline di Punta della Contessa Le Cesine X X X X X X X X X X

X X X X X X X X X X X X

X X

X X X X X

X X X X X X

X X X X X

X X X X X X X X X X

X X

X X X X X X X

X X

restricted to a few scattered patches, due to the reclamation of their potential areas for agricultural use. One of the most evident physiognomic eects of the anthro-

pogenic pressures is the passage from the typical homogeneous and continuous distribution of the salt marsh communities in the form of concentric belts around the

1042 coastal pools to an extremely fragmented and discontinuous mosaic. In order to preserve the remaining high degree of biological diversity (both in terms of species and ecosystems) of these areas, it is necessary to keep up a high level of policy and research monitoring (if possible expanding and diversifying the type of scientic studies). Indeed, it is likely that in addition to the wellknown role of the physical factors in determining the potential heterogeneity pattern that characterises these environments, there is a complicated network of biotic interrelationships established between species and/or communities which is still not completely understood or explained.
Acknowledgements This phytosociological study is a part of the activities of an INTERREG project (III A Italy-Greece 20002006) concerning the environmental monitoring of coastal wetlands belonging to the Natura 2000 network. The authors would like to thank Dr. Massimo Terzi of CNR-IGV for statistical consultation and two anonymous referees for helpful comments on the previous version of this paper. References
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