Coastal salt-marshes plant communities of the
Transcript
Coastal salt-marshes plant communities of the
Biologia 69/1: 53—69, 2014 Section Botany DOI: 10.2478/s11756-013-0283-2 Coastal salt-marshes plant communities of the Salicornietea fruticosae class in Apulia (Italy) Saverio Sciandrello1 & Valeria Tomaselli2 1 Department of Biological, Geological and Environmental Sciences, University of Catania, via A. Longo 18 95125 Catania, Italy; e-mail: [email protected] 2 C.N.R., Institute of Plant Genetics, via G. Amendola 165/A 70126 Bari, Italy; e-mail: [email protected] Abstract: An overview of the salt-marsh herbland and scrub vegetation belonging to the class Salicornietea fruticosae Br.-Bl. et Tx. ex A. Bolòs y Vayreda 1950 in Apulia is presented. Data available from literature have been supplemented with original relevés performed in different locations of the Apulia region. On the basis of a total of 297 relevés, fifteen communities have been defined, according to the traditional phytosociological system based on dominant and/or diagnostic taxa. For comparison purposes, the salt-marsh vegetation has been classified using numerical methods. The results obtained show that most of the clusters correspond to specific associations, and confirm the division into vegetation alliances and orders. Numerical analysis also allowed us to assign the proper allocation of some associations and plant communities drawn from literature. Five alliances, with plant communities characterized by specific ecological features, have been discriminated: Sarcocornion alpini and Arthrocnemion glauci (lower marshes), Salicornion fruticosae (middle marshes), Inulion crithmoidis and Suaedion brevofoliae (upper marshes). In addition, during the field work, a population of Halocnemum strobilaceum (Arthrocnemo-Halocnemetum strobilacei), new record for the Apulia region, has been found. Key words: Halocnemum strobilaceum communities; Saliconietea fruticosae; cluster analysis; salt marshes; ordination; vegetation; Apulia Introduction Salicornietea fruticosae Br.-Bl. et Tx. ex A. Bolòs y Vayreda 1950 (Mucina 1997) includes halophilous shrubs plant communities of both coastal and inland areas, developing on halomorphic soils of salt marshes, estuaries, coastal lagoons. It spreads in a bioclimatic range from the infra- to the supramediterranean / arid to dry bioclimatic belts in the Mediterranean and Atlantic European regions (Rivas-Martınez et al. 2011). This type of vegetation is characterized mainly by succulent Chenopodiaceae with a chamaephytic to nanophanerophytic habit, such as Sarcocornia alpini, Sarcocornia fruticosa, Arthrocnemum macrostachyum, Halocnemum strobilaceum, Halimione portulacoides, etc. Traditionally, three alliances have been distinguished in the Italian region: Salicornion fruticosae (halophilous communities regularly inundated), Arthrocnemion glauci (halophilous communities irregularly inundated) and Suaedion brevifoliae (halo-nitrophilous communities rarely inundated). Three more syntaxa have been later ascribed to the Salicornietalia: Sarcocornion alpinii (halophilous communities regularly inundated and for long time), Inulion crithomides (halopholius communities only occasionally inundated), c 2013 Institute of Botany, Slovak Academy of Sciences Limonion ferulacei (thermo-halophilous plant communities flooded for short periods). In Italy the salt-marsh vegetation belonging to the Sarcocornietea fruticosae has been extensively studied, albeit the resulting syntaxonomic schemes show some inconsistencies (Pignatti 1966; Corbetta 1970, 1976; Valsecchi 1964,1972; Brullo & Di Martino 1974; Corbetta & Lorenzoni 1974; Brullo & Furnari 1976; Caneva et al. 1981; Géhu & Biondi 1994; Brullo et al. 1988; Pirone 1988, 1995; Taffetani & Biondi 1989; Corbetta et al. 1989, 2006; Mariotti et al. 1992; Piccoli et al. 1994; Ferrari et al. 1995; Caniglia et al. 1997; Biondi 1999; Poldini et al. 1999; Filigheddu et al. 2000; Pirone et al. 2001; Viciani & Lombardi 2001; Frondoni & Iberite 2002; Beccarisi et al. 2003; Andreucci 2004; Biondi & Bagella 2005; Biondi et al. 2006; Maiorca et al. 2006; Sciandrello 2007; Giusso et al. 2008; Biondi & Casavecchia 2010; Cutini et al. 2010; Minissale & Sciandrello 2010; Tomaselli et al. 2009a, 2009b, 2011). The syntaxonomical units at the community level within the Salicornietea fruticosae, need more clarifications and deepening. In this study, we provide a synthesis of the Salicornietea fruticosae for the Apulia region, taking into account floristic composition structure, distribution and ecological features. Unauthenticated Download Date | 3/16/17 6:12 AM 54 S. Sciandrello & V. Tomaselli Fig. 1. Study area and sample sites. Material and methods Study area The following study sites (Fig. 1) have been considered: A) Adriatic side: 1. Mouth of Fortore river “Foce del Fortore” (Fg), 2/3. Lesina and Varano lakes (Fg), 4. Sfinale and Gusmay swamps (Fg), 5/6. Frattarolo swamp “Palude Frattarolo” and Salso lake “Lago Salso” (Fg), 7. Saltworks and salt marshes “Saline di Margherita di Savoia” (Fg), 8. Torre Canne (Br), 9. Pantanagianni (Br), 10. Torre Guaceto (Br), 11. Salt marshes “Saline di Punta della Contessa” (Br), 12. Le Cesine (Le); B) Ionian side: 13. Swamps “Bacini di Ugento” (Le), 14. Swamp “Palude del Capitano” (Le), 15. Swamp “Palude la Vela”, Mar Piccolo (Ta). The annual average temperature in Apulia is about 15 ◦C, although in coastal areas it ranges between 16 and 17 ◦C (Blasi et al. 2007). Annual precipitations are between 590 and 640 mm. According to Rivas Martinez et al. (2004) the area is characterized by the Mediterranean pluviseasonal oceanic bioclimate, with thermotypes ranging between the termo-Mediterranean and meso-Mediterranean, and ombrotypes mainly between dry and subhumid, with the humid type limited to a few mountain areas. Recent studies revealed a general trend in aridity and then a growing vulnerability to desertification (Blasi et al. 2007). The landscape of coastal areas is characterized by a large extent of cultivated areas, often subject to intensive agriculture. So, most of the coastal wetlands are nested in this matrix of cultivated fields and are subjected to different polluting agents and to intense fragmentation. Vegetation analysis The collection of vegetation data was carried out following the Zürich-Montpellier phytosociological method (BraunBlanquet 1964). Phytosociological relevés (132 unpublished relevés, Table 1) have been performed in the period April 2007-September 2011 and geographical coordinates have been recorded by using GPS. The size of plots varies from a maximum of 200 m2 to 50 m2 , depending on the vegetation type and the micro-topography. All available literature data related to the class Salicornietea fruticosae in the Apulia region have been collected (Géhu et al. 1984; Corbetta et al. 1989; Taffetani & Biondi 1989, Mariotti et al. 1992; Biondi et al. 2006; Corbetta 2006, Biondi & Casavecchia 2010) and then together with new unpublished vegetation relevés, have been combined into a single matrix of 297 relevés × 98 species. In order to compare and verify the syntaxa traditionally including the salt-marsh scrub communities, all relevés were classified by numerical methods. Similarity analysis of the relevés was carried out by using the SYN-TAX 2000 software (Podani 2001). Original BraunBlanquet sampling scale has been transformed into the ordinal scale according to Van der Maarel (1979). A cluster analysis based on the Bray-Curtis dissimilarity index and the UPGMA linkage method was performed. The ordination of the data-set was carried out by using the PC-ORD 4.34 software. In the ordination analyses we ran the Nonmetric Multidimensional Scaling (NMS), based on the Euclidean distance, achieving a consistent ordination diagram. 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 satisfied in field community data (Clarke 1993). Clusters of relevés were classified and attributed to syntaxa according to Rivas-Martinez et al. (2001, 2004, 2011), Brullo et al. (2002) and Pandža et al. (2007). The Latin names of marshland communities, with names of authors and year of first valid description, are given at least once in the text. Nomenclature of the taxa cited in the text is in accordance with Flora Europaea (Tutin et al. 1964–1993), Med-Checklist (Greuter et al. 1984–1989, 2009) and Conti et al. (2005). Results and discussion Multivariate analysis Cluster analysis identified two main clusters, A and B (Fig. 2), separating respectively the halophilous (A) from the halo-nitrophilous (B) communities. Cluster A splits in two main sub-clusters: A1 and A2. Unauthenticated Download Date | 3/16/17 6:12 AM Table 1. Unpublished relevés performed on plant communities belonging to the Salicornietea fruticosae class during the period April 2007–September 2011 in coastal sites of the Apulia region. The Salicornietea fruticosae class in Apulia (Italy) 55 Unauthenticated Download Date | 3/16/17 6:12 AM 56 S. Sciandrello & V. Tomaselli Fig. 2. Cluster analysis. Cluster A1 includes the hypersaline communities of Arthrocnemion glauci [sub-cluster A11, with two subclusters A111 (Arthrocnemum macrostachyum communities) and A112 (Halocnemum strobilaceum communities)], and Sarcocornion alpini [sub-cluster A12, with two sub-clusters A121 (Sarcocornia alpini communities) and A122 (Aeluropus littoralis communities)]. Cluster A2 includes the halophilous communities of Salicornion fruticosae (sub-cluster A21) and Inulion crithmoidis (sub-cluster A22, grouping the communities of Inula crithmoides with the communities of Artemisia coerulescens). Cluster B includes the halo-subnitrophilous communities of Suaedion brevifoliae. Ordination diagram The NMS (Fig. 3) shows the main significance along the axis 1, related mainly to the flooding time and to the soil salinity gradient. The groups of Inulion crithmoidis and Salicornion fruticosae, which occupy the left part of the diagram, are clearly parted from the most halophilous communities of Sarcocornion alpini and Arthrocnemion glauci, placed in the right part of the graph. Syntaxonomic survey The plant communities identified are listed in the syntaxonomic list below and their description including floristic composition, structure, ecology and distribution, is provided in the following sections. For the nomenclature of salt-marsh vegetation syntaxa and synonyms, the basic principles of the European Vegetation Checklist, under development by Mucina et al. expected in 2013 (pers. comm.). SALICORNIETEA FRUTICOSAE Br.-Bl. et Tx. ex A. Bolòs y Vayreda 1950 (Sarcocornietea fruticosae Br.-Bl. & R.Tx. ex A. & O. Bolòs 1950) SALICORNIETALIA FRUTICOSAE BR.-BL. 1933 (Sarcocornietalia fruticosae Br.-Bl.1933) Arthrocnemion glauci Rivas-Mart. et Costa 1984 (Arthrocnemion macrostachyi Rivas-Martínez & Costa 1984) (cluster A11) Puccinellio convolutae-Arthrocnemetum macrostachyi (Br.-Bl. (1928) 1933) Géhu ex Géhu, Costa, Scoppola, Biondi, Marchiori, Peris, Géhu-Franck, Caniglia, Veri 1984 subass. typicum subass. limonietosum bellidifolii subass. nov. Unauthenticated Download Date | 3/16/17 6:12 AM The Salicornietea fruticosae class in Apulia (Italy) 57 Fig. 3. NMS ordination analysis. 1. Sarcocornion alpini; 2. Salicornion fruticosae; 3. Arthrocnemion glauci; 4. Inulion crithmoidis; 5 Suaedion brevifoliae. Arthrocnemo-Juncetum subulati Brullo & Furnari 1976 Arthrocnemo-Halocnemetum strobilacei Oberd.1952 Sarcocornion alpini (Rivas-Martínez et al. 1990) Brullo, Giusso del Galdo, Minissale, Siracusa & Spampinato 2002 (Cluster A12) Puccinellio festuciformis-Sarcocornietum alpini Castroviejo & Cirujano 1980 Sarcocornietum alpini Br.-Bl. 1933 corr. RivasMartínez, Lousã, T.E. Díaz, Fernández-González & J.C. Costa 1990 Halimiono portulacoidis-Sarcocornietum alpini Rivas-Martínez & Costa 1984 Puccinellio festuciformis-Aeluropetum littoralis (Corb, 1968) Géhu & Costa 1984 in Géhu, Costa, Scoppola, Biondi, Marchiori, Peris, Géhu- Franck, Caniglia & Veri 1984 Salicornion fruticosae Br.-Bl. 1933 (Sarcocornion fruticosae Br.-Bl. 1933 em. Brullo & Furnari 1988) (Cluster A21) Puccinellio festuciformis-Sarcocornietum fruticosae (Br.-Bl. 1928) J.M. Géhu 1976 subass. typicum subass. juncetosum maritimi Géhu 1976 Junco subulati-Sarcocornietum fruticosae Brullo & Furnari 1990 Unauthenticated Download Date | 3/16/17 6:12 AM 58 Inulion crithmoidis Brullo & Furnari 1988 (Cluster A22) Agropyro scirpei-Inuletum crithmoidis Brullo in Brullo et al.1988 Limonio narbonensis-Artemisietum coerulescentis Horvatic (1933) 1934 Suaedion brevifoliae Br.-Bl. et O. de Bolòs 1958 (Suaedion verae (Rivas-Martínez, Lousã, T.E. Díaz, Fernández-González & J.C. Costa 1990) Rivas-Martínez, Fernández-González & Loidi 1999) (Cluster B) Halimiono portulacoidis-Suaedetum verae Molinier & Tallon 1970 corr. Géhu in Géhu & al. 1984 Suaedetum verae Br.-Bl. ex O. Bolòs & Molinier 1958 Class Salicornietea fruticosae Br.-Bl. et Tx. ex A. Bolòs y Vayreda 1950 The Salicornietea fruticosae includes scrub vegetation communities settling in coastal salt marshes and dominated by succulent plants belonging to the family of Chenopodiaceae. These communities grow on soils periodically inundated by salt or brackish waters and form more or less regular belts around the water bodies. The class spreads in the Mediterranean and Atlantic European regions, in infra-supramediterranean arid-subhumid and thermo-mesotemperate subhumidhumid bioclimatic belts. As regards the structure, these plant communities are characterized mainly by chamaephytic and nanophanerophytic life forms. Among the diagnostic taxa, we mention: Sarcocornia alpini, Sarcocornia fruticosa, Arthrocnemum macrostachyum, Halocnemum strobilaceum, Suaeda vera, Halimione portulacoides, etc. In the Apulia region, this class is represented by the Salicornietalia fruticosae Br.-Bl.1933 order with the following five alliances: I) Alliance Arthrocnemion glauci Rivas-Mart. et Costa 1984 (Table 4) The Arthrocnemion macrostachyi includes halo-xerophilous plant communities growing in arid environments with salty clay soils subject to a more or less long period of submersion. These communities tolerate large variations in salinity of the water table throughout the year. Puccinellio convolutae-Arthrocnemetum macrostachyi (Br.-Bl. (1928) 1933) Géhu ex Géhu, Costa, Scoppola, Biondi, Marchiori, Peris, Géhu-Franck, Caniglia, Veri 1984 Syn.: Puccinellio convolutae-Arthrocnemetum glauci (Br.-Bl. 1928) 1933 Géhu 1984; Arthrocnemum macrostachyum communities. Diagnostic taxa: Puccinellia festuciformis (Host) Parl. subsp. convoluta (Hornem.) W.E. Hughes Structure and ecology: This association grows mainly on clayey soils subject to more o less long S. Sciandrello & V. Tomaselli periods of flooding. The physiognomy of this vegetation is given by the dense bushes of Arthrocnemum macrostachyum and the diagnostic species Puccinellia festuciformis subsp. convoluta. In the most raised parts, where soils are subject to a moderate period of submersion, the association comes in contact with different types of vegetation belonging to Salicornion fruticosae o Inulion crithmoidis. Furthermore, this vegetation supports long drought periods and finds its optimal conditions even on hypersaline soils (Poldini et al. 1999; Andreucci 2004). In areas subject to anthropic pressuresArthrocnemum macrostachyum tends to spread forming almost monospecific communities (Biondi et al. 2006), sporadically in association with other halophytes such as Limonium narbonense, Limbarda crithmoides, Halimione portulacoides, Sarcocornia fruticosa, etc. Distribution: widespread in the Mediterranean area; distributed along the coasts of the Italian peninsula and Sardinia, not reported in Sicily. In Apulia this association has been found in: Salinella lake (Taranto), Mar Piccolo (Taranto), Saline Margherita di Savoia (Foggia), mouth of Carapelle river “Foce Carapelle” (Foggia), mouth of Candelaro river “Foce Candelaro” (Foggia), Tricase Porto, Torre Colimena, Vecchia Salina, Riva degli Angeli, Bacino Frigole, Palude del Capitano (Lecce); Torre Canne and Lido Morelli, Polignano a Mare, Pantanagianni (Brindisi); S. Pietro ”Punta la Forca” (Taranto). Relevés from literature: Géhu et al. 1984, Table 17; Corbetta et al. 1989, Table 15, rel. 1-3; Biondi et al. 2006, Table 8, rel. 1-18; Corbetta et al. 2006, Table 4, rel. 1,3,5,7,9; Biondi & Casavecchia 2010, Table 13, rel. 1–5. Our unpublished relevés: Sciandrello & Tomaselli 2011, Table1, rel. 59, 62, 64, 67–74, 76, 78–83. Puccinellio convolutae-Arthrocnemetum macrostachyi (Br.-Bl. 1928) Géhu ex Géhu, Costa, Scoppola, Biondi, Marchiori, Peris, Géhu-Franck, Caniglia & Veri 1984 limonietosum bellidifolii subass. nov. (holotypus: rel. 58, Table 2) Diagnostic taxa: Limonium bellidifolium (Gouan) Dumort. Structure and ecology: This vegetation type replaces the previous one in the most low-lying parts of salt marshes, enduring long periods of submersion and high salt concentration. The presence of Limonium bellidifolium, a thermophilous species widespread in northern Mediterranean, characterizing the subassociation has to be pointed out. From a structural point of view, the vegetation is characterized by the presence of several halophilous species, such as Puccinellia festuciformis subsp. convoluta, Arthrocnemum macrostachyum, Sarcocornia alpini, Halimione portulacoides, Limbarda crithmoides, Triglochin bulbosum subsp. barrelieri, Limonium narbonense, etc. (Table 2) Distribution: widespread in the Mediterranean area; distributed along the coasts of the Italian peninsula and Sardinia, not reported in Sicily. In Apulia this associUnauthenticated Download Date | 3/16/17 6:12 AM The Salicornietea fruticosae class in Apulia (Italy) 59 Table 2. Puccinellio convolutae-Arthrocnemetum macrostachyi (Br.-Bl. 1928) Géhu ex Géhu et al. 1984 limonietosum bellidifolii subass. nov. ation has been found in: Saline Margherita di Savoia (Foggia). Relevés from literature: Corbetta et al. 1989, Table 15, rel. 1-3; Corbetta et al. 2006, Table 4, rel. 2,4,6,8,10. Our unpublished relevés: Sciandrello & Tomaselli 2011, Table1, 58, 60-61, 63, 65-66, 75, 77-78, 84-85. Arthrocnemo-Juncetum subulati Brullo & Furnari 1976 Diagnostic taxa: Juncus subulatus Forssk. Structure and ecology: This association finds its optimum in the most interior of brackish and salt marshes, on muddy-sandy soils subjected to long flooding periods (Brullo & Furnari 1976). In fact the plant community is characterized by Juncus subulatus which is linked to edaphic moisture, even during the summer period. The structure of these plant communities is shaped by a gradient closely related to edaphic moisture. In the most humid part Juncus subulatus presents the hightest coverage, while in the less humid and generally more raised parts Arthrocnemum macrostachyum tends to become the dominant species (Tomaselli et al. 2011). This vegetation replaces (vicariant) the Puccinellio convolutaeArthrocnemetum macrostachyi in the most arid areas of southern Mediterranean territories. Distribution: widely distributed in the Southern Mediterranean areas. In Apulia has been found in Saline Punta della Contessa (Brindisi), Torre Canne (Brindisi), Saline Margherita di Savoia (Foggia). Relevés from literature: Biondi & Casavecchia 2010, Table 13, rel.5. Our unpublished relevés: Tomaselli & Sciandrello 2007, Table1, rel. 53-57. Arthrocnemo-Halocnemetum strobilacei Oberd. 1952 (Table 3) Syn.: Salicornio-Halocnemetum strobilacei Wolff 1968 Diagnostic taxa: Halocnemum strobilaceum (Pall.) M. Bieb. Structure and ecology: Markedly thermo-xerophilous vegetation, that is suited to high salt concentraUnauthenticated Download Date | 3/16/17 6:12 AM 60 S. Sciandrello & V. Tomaselli Fig. 4. Transects of halophilous vegetation in some sample sites. 1. “Saline della Contessa”; 2.“Torre Canne”; 3. “Palude Frattarolo”; 4. “Saline Margherita di Savoia, Almadannata”; 5. “Foce Carapelle”. A: Salsolo-Cakiletum maritimae; B: Cypero-Elymetum farcti; C: Spartino-Juncetum maritimi; D: Puccinellio-Sarcocornietum alpini; E: Suaedo-Salicornietum patulae; F: Puccinellio-Sarcocornietum fruticosae; G: Limonio-Atriplicetum coerulescentis; H: Entheromorpho-Ruppietum maritimae; I: Medicagini-Ammophiletum australis; L: Pistacio-Juniperetum macrocarpae; M: Schoeno-Plantaginetum crassifoliae; N: Agropyro scirpei-Inuletum crithmoidis; O: Inulo-Juncetum maritimi; P: Salsolo sodae-Suaedetum splendentis; Q: Halimiono-Sarcocornietum alpini; R: Halimiono portulacoidisUnauthenticated Suaedetum verae; S: Salicornietum emerici; U: Puccinellio convolutae-Arthrocnemetum galuci subass. limonietosum bellidifolii; T: Puccinellio convolutae-Arthrocnemetum glauci; V: Arthrocnemo-Halocnemetum strobilacei. Download Date | 3/16/17 6:12 AM The Salicornietea fruticosae class in Apulia (Italy) 61 Table 3. Arthrocnemo-Halocnemetum strobilacei Oberd. 1952. Fig. 5. Halocnemum strobilaceum communities in their natural environments near the mouth of Carapelle river. tion in the soil (El-Sheikh et al. 2006). It occupies the central portion of the salt marsh, subject to desiccation from the end of the spring to the early autumn. In summer these surfaces are usually covered with salt deposits. Guide species of the association is Halocnemum strobilaceum which is accompanied by Arthrocnemum macrostachyum and a few other halophytes. In the slightly raised parts surrounding this plant community, subject to a shorter period of submersion, it takes usually contact with the Puccinellio convolutaeArthrocnemetum macrostachyi. Distribution: widespread in southern Mediterranean, southwestern and western Central Asia. In Apulia Halocnemum strobilaceum has been found for the first time during the field surveys for the present research, in the site Foce del Carapelle (Capitanata, Fg) (Fig. 5,6). Notes: Halocnemum strobilaceum is a rare halophyte. It had never been reported for the Apulia region. In Italy this species is extremely localized, in the western part of Sicily (Saline di Trapani and Isola Lunga dello Fig. 6. Halocnemum strobilaceum, flowering branch system. Stagnone (Brullo & Di Martino 1974), in the southern part of Sardinia (Stagni di Cagliari and Isola di S. Antioco) (De Marco et al. 1980; De Martis & Serri 2009), in Emilia-Romagna (Sacca di Bellocchio, Valli di Comacchio, Ravenna) (Piccoli et al. 1994) and Tuscany (Foce dell’ Ombrone, Maremma Natural Park, Grosseto) (Arrigoni et al. 1985; Andreucci 2004). The species is included in the red book of plants of Italy with the IUCN categories “critically endangered” (CR) (Conti et al., 1997). In addition, the plant communities with Halocnemum strobilaceum and Arthrocnemum macrostachyum, are included in the Appendix I of the Habitats Directive, with the code 1420 “Mediterranean and thermo-Atlantic halophilous scrubs” (Salicornietea fruticosae). Our unpublished relevés: Sciandrello & Tomaselli 2011, Table 1, Rel. 86-96. II) Alliance Sarcocornion alpini (Rivas-Martínez et al. 1990) Brullo, Giusso del Galdo, Minissale, Siracusa & Spampinato 2002 (Table 4) Unauthenticated Download Date | 3/16/17 6:12 AM 62 S. Sciandrello & V. Tomaselli Table 4. Synoptic table of Salicornietea fruticosae communities: 1. Puccinellio convoluti-Sarcocornietum alpini; 2. Sarcocornietum alpini; 3. Halimiono portulacoides-Sarcocornietum alpini; 4. Puccinellio festuciformis-Aeluropetum litoralis; 5. Junco subulatiSarcocornietum fruticosae; 6. Puccinellio-Sarcocornietum fruticosae; 7. Puccinellio festuciformis-Arthrocnemetum fruticosi juncetosum maritimi; 8. Arthrocnemo-Juncetum subulati; 9. Puccinellio convolutae-Arthrocnemetum glauci; 10. Puccinellio convolutaeArthrocnemetum glauci limonietosum bellidifolii; 11. Arthrocnemo-Halocnemetum strobilacei; 12. Limonio-Artemisietum coerulescentis; 13. Agropyro-Inuletum crithmoidis; 14. Halimiono-Suaedetum verae; 15. Suadetum verae. Unauthenticated Download Date | 3/16/17 6:12 AM The Salicornietea fruticosae class in Apulia (Italy) In the Sarcocornion alpini alliance fall halophilous dwarf scrub (with chamephytes) communities, typical of those parts of the salt marshes that are subjected to long flooding periods and with salty soils. Puccinellio festuciformis-Sarcocornietum alpini Castroviejo & Cirujano 1980 Syn.: Salicornietum radicantis Br.-Bl. (1931) 1952; Puccinellio festuciformis-Arthrocnemetum perennis (Br.-Bl. 1931) Géhu 1976; Puccinellio festuciformisSarcocornietum perennis (Br.-Bl. 1931) Géhu 1976; Sarcocornietum deflexae (Braun-Blanquet 1931) Lahonderè, Gèhu et Paradis 1992. Diagnostic taxa: Puccinellia festuciformis (Host) Parl. Structure and ecology: This association usually colonizes the central part of the salt marshes, in correspondence of the sections subject to long periods of submersion (Frondoni & Iberite 2002). Physiognomically it is characterized by Sarcocornia alpini, a shrubby chamaephyte with a reptant habit, which settles down quite well with salty soils where it tends to form dense populations (Brullo et al. 2000; Maiorca et. al. 2007; Tomaselli et al. 2009b). Only few species are associated with Sarcocornia alpini, including Puccinellia festuciformis, a markedly thermophilous species widespread mainly in the northern Mediterranean. In the most raised part of the marsh, subject to a shorter period of submersion, this vegetation takes contact with plant communities belonging to the Arthrocnemion glauci or, more rarely, with those of the Salicornion fruticosae. It can be considered as the (north) vicariant of Aeluropo lagopoides-Sarcocornietum alpini, the latter distributed mainly in the southern Mediterranean area (Brullo & Furnari 1990). Distribution: widely distributed in the Mediterranean area; occurs in Italy and in Sardinia, but not in Sicily. In Apulia this association has been found inSaline Punta della Contessa (Brindisi), Torre Canne (Brindisi), Saline Margerita di Savoia (Foggia), Lago di Lesina (Foggia), Lago Salinella (Taranto), Palude Frattarolo (Foggia), Foce Carapelle (Foggia), Zapponeta (Foggia). Notes: in recent papers, Sarcocornia alpini communities have been assigned to the Sarcocornietum deflexae (Braun-Blanquet 1931) Lahonderè, Gèhu et Paradis 1992 (Filigheddu et al. 2000; Biondi & Bagella 2005; Corbetta 2006, Biondi & Casavecchia 2010), but according to Frondoni & Iberite (2002) we prefer to assign these communities to the association Puccinellio festuciformis-Sarcocornietum alpini. Relevés from literature: Corbetta et al. 1989, Table 16, rel. 1-6,8-16; Corbetta et al. 2006, Table 2, rel. 5,7. Our unpublished relevés: Tomaselli & Sciandrello 2007, Table 1, rel.1-10; Sciandrello & Tomaselli 2011, Table 1, rel.15-28, 31-45. Sarcocornietum alpini Br.-Bl. 1933 corr. RivasMartínez, Lousã, T.E. Díaz, Fernández-González & J.C. 63 Costa 1990 Syn.: Salicornietum radicantis Br.-Bl. (1931) 1952; Sarcocornietum deflexae (Braun-Blanquet 1931) Lahonderè, Gèhu et Paradis 1992 (Corbetta et al., 2006; Biondi & Casavecchia, 2010) Diagnostic taxa: Sarcocornia alpini (Lag.) Castrov. Structure and ecology: This association, often in contact with the previous one, usually colonizes the central part of the salt marshes, in correspondence of the sections subject to very long periods of submersion. Physiognomically it is characterized by Sarcocornia alpini, which tends to form dense and almost monophytic populations (Biondi & Casavecchia 2010). In the most raised part of the marsh, subject to a shorter period of submersion, this vegetation takes contact with plant communities belonging to the Arthrocnemion glauci or to the Salicornion fruticosae alliances. Distribution: widely distributed in western and southern Mediterranean area; In Apulia this association has been found in: Saline Punta della Contessa (Brindisi), Torre Guaceto (Brindisi), Le Cesine (Lecce), Saline Margerita di Savoia (Foggia). Relevés from literature: Corbetta et al. 1989, Table 16, rel. 7,17,24; Corbetta et al., 2006, Table 2, rel. 1-4, 10; Biondi & Casavecchia, 2010, Table 12, rel. 1-3. Our unpublished relevés: Tomaselli & Sciandrello. 2007, Table 1, rel. 11-14. Halimiono portulacoidis-Sarcocornietum alpini Rivas-Martínez & Costa 1984 Syn.: Halimione portulacoides comm. (Corbetta et al., 1989) Diagnostic taxa: Halimione portulacoides (L.) Aellen Structure and ecology: This association usually grows in those parts of the salt marshes subject to moderate periods of submersion with soils rich in organic component and well drained (Álvarez-Rogel et al. 2007). The structure is given by a layer of Halimione portulacoides mixed with scattered bushes of Sarcocornia alpini. This halo-nitrophilous vegetation in the most raised parts, subject to a shorter period of submersion, takes contact with plant communities of Suaedion brevifoliae. If these areas are subject to prolonged flooding periods, Sarcocornia alpini tends to predominate, developing almost monophytic communities. Distribution: Mediterranean area. In Apulia: Lago Salinella (Taranto), Saline Punta della Contessa (Brindisi), Saline Margherita di Savoia (Foggia). Relevés from literature: Géhu et al. 1984, Table 15; Corbetta et al., 1989, Table 16, rel. 22-26, Table 17, rel. 1-2; Corbetta et al., 2006, Table 2, rel. 6,8,9. Puccinellio festuciformis-Aeluropetum litoralis (Corbetta 1968) Géhu et Costa 1984 Syn.: Aeluropus litoralis and Agropyron elongatum comm. in Corbetta 1968 Diagnostic taxa: Puccinellia festuciformis (Host) Parl., Aeluropus littoralis (Gouan) Parl. Structure and ecology: The association tends to localize in small sandy depressions subject to moderate Unauthenticated Download Date | 3/16/17 6:12 AM 64 periods of submersion, covering limited surfaces and in contact, in correspondence of the slightly raised parts (outer edges), with the Puccinellio-Sarcocornietum fruticosae. The association is characterized by the dominance of Aeluropus littoralis, rhizomatous species forming dense thickets, and by the presence of Puccinellia festuciformis subsp. convoluta. Géhu et al. (1984) assigned this association to the Juncetea maritimi class, but in our opinion, considering the presence and dominance of several salt-tolerant species, such as Sarcocornia alpini, Arthrocnemum macrostachyum, Limbarda crithmoides, Limonium narbonense etc., it should be appropriate to assign this vegetation to the Salicornietea fruticosae class (see cluster analysis). Distribution: localized mainly along the Adriatic coast of the Italian peninsula, not reported in Sicily and Sardinia. In Apulia: Foce Fiume Bifero, Foce Fiume Candelaro (Foggia), Lago Salinella (Taranto) and Saline Margherita di Savoia (Foggia). Relevés from literature: Gehu et al. 1984, Table 21; Corbetta et al. 1989, Table 18, rel. 1-10; Taffetani & Biondi 1989, Table 15, rel. 1-4; Corbetta et al., 2006, Table 6, rel.1-3. Our unpublished relevés: Sciandrello & Tomaselli 2011, Table 1, rel. 29-30. III) Alliance Salicornion fruticosae Br.-Bl. 1933 (Table 4) The Salicornion fruticosae alliance, includes the shrub communities related to salty soils subject to not too prolonged submersion. Puccinellio festuciformis-Sarcocornietum fruticosae (Br.-Bl. 1928) J.M. Géhu 1976 typicum Syn.: Puccinellio festuciformis-Arthrocnemetum fruticosi (Br. Bl. 1928) Géhu 1976; Sarcocornia fruticosa communities; Puccinellio festuciformis-Arthrocnemetum fruticosi juncetosum maritimi (Taffetani & Biondi 1989); Limonium virgatum and Sarcocornia fruticosa communities (Mariotti 1992). Diagnostic taxa: Puccinellia festuciformins (Host) Parl. subsp. convoluta (Hornem.) W.E. Hughes Structure and ecology: This association is localized in stands subject to moderate periods of submersion. Usually it forms closed structures, covers limited areas and is adjacent, in the most raised parts, to the Agropyro scirpei-Inuletum crithmoidis. This vegetation is characterized by the dominance of Sarcocornia fruticosa which forms dense populations. Puccinellia festuciformis subsp. convoluta differentiates the association. Corbetta et al. (1989) recognized three subassociations: sarcocornietosum alpini Corbetta & Pirone 1989, colonizing soils subject to prolonged drying in summer; arthrocnemetosum glauci Gehu 1976, which colonizes the highest and arid stands; hordeetosum maritimi Corbetta & Pirone 1989, which represents an aspect of transition to the halo-nitrophilous and therophytic communities of Saginetea maritimae. As regards the study area, we recognized the typical form (subass. typicum) and the subass. junceto- S. Sciandrello & V. Tomaselli sum maritimi Géhu 1976. In areas subject to anthropic pressures Sarcocornia fruticosa tends to spread forming almost monospecific communities, rarely accompanied by other halophytes such as Limonium virgatum, Limbarda crithmoides, Arthrocnemum macrostachyum, Halimione portulacoides, etc. Distribution: this association is distributed in WMediterranean areas. In Apulia it has been found in Lago Salinella and Mar Piccolo (Taranto), Saline Margherita di Savoia (Foggia), Laguna di Lesina (Foggia), Foce F. Biferno (Foggia), Saline di Punta della Contessa (Brindisi), Torre Guaceto (Brindisi). Relevés from literature: Gehu et al. 1984, Table16; Corbetta et al., 1989, Table 13, rel. 1-12; Taffetani & Biondi 1989, Table 10, rel. 1-2; Corbetta 2006, Table 3, rel. 5; Mariotti et al. 1992, Table 5, rel. 29-31; Corbetta et al., 2006, Table 3, rel. 1-4; Biondi & Casavecchia 2010, Table 11, rel. 1-5. Puccinellio festuciformis-Sarcocornietum fruticosae (Br.-Bl. 1928) J.M. Géhu 1976 subass. juncetosum maritimi Géhu 1976 Syn.: Puccinellio festuciformis-Arthrocnemetum fruticosi (Br.-Bl. 1928) J.M. Géhu 1976 subass. juncetosum maritimi Géhu 1976. Diagnostic taxa: Puccinellia festuciformis (Host) Parl., Juncus maritimus Lam. Structure and ecology: Replaces the typical form in the most depressed areas, subject to long flooding periods and characterized by weakly salty soils (Taffetani & Biondi 1989). This vegetation is characterized by the dominance of Juncus maritimus, which normally characterizes the communities belonging to the Juncetea maritimi class. Distribution: localized mainly along the Adriatic coast of the Italian peninsula. In Apulia: Foce del F. Biferno (Foggia). Relevés from literature: Taffetani & Biondi 1989, Table 10. Junco subulati-Sarcocornietum fruticosae Brullo & Furnari 1990 Diagnostic taxa: Juncus subulatus Forssk. Structure and ecology: The association is localized in the low-lying parts of the marsh, subject to prolonged periods of flooding, and mantaining moist soils during the summer (Brullo et al. 2000). It tends to cover extended surfaces areas and is adjacent, in the most raised parts, to the Agropyro scirpei-Inuletum crithmoidis. This vegetation is dominated by Sarcocornia fruticosa, that form dense populations. Juncus subulatus well characterizes the association from the other Mediterranean and Atlantic vegetation types with Sarcocornia fruticosa. Distribution: in S-Mediterranean areas; in Apulia it has been found in Saline Punta della Contessa (Brindisi) and Palude la Vela (Taranto). Our unpublished relevés: Sciandrello & Tomaselli 2007, Table 1, Rel. 46-50; Tomaselli 2010, Table 1, rel. 51-52. Unauthenticated Download Date | 3/16/17 6:12 AM The Salicornietea fruticosae class in Apulia (Italy) IV) Alliance Inulion crithmoidis Brullo & Furnari 1988 (Table 4) The Inulion crithmoidis alliance groups halophilic communities with chamaephytic and hemicriptophytic habit, growing of soils subject to sporadic flooding periods. Limonio-Artemisietum coerulescentis Horvatic (1933) 1934 Syn.: Staticeto-Artemisietum coerulescentis Horvatič 1933; Puccinellio festuciformis-Arthrocnemetum fruticosi juncetosum maritimi (Taffetani & Biondi, 1989). Diagnostic taxa: Limonium narbonense Mill., Artemisia coerulescens L. Structure and ecology: This vegetation grows in peripheral stations of salt marshes, on silty-clay soils, moist for most of the year. Physiognomically, it can be recognized by the dominance of Artemisia coerulescens that is associated with Limonium narbonense and many other halophytes, as Puccinellia festuciformis subsp. convoluta, Halimione portulacoides, Elytrigia scirpea, ect. (Pignatti 1966; Poldini et al. 1999; Tomaselli et al. 2011). This association spatially settles between the communities of Sarcocornion alpini (o Artrocnemion glauci) and those of Inulion crithmoidis. Distribution: in Italy, along the Adriatic coasts. In Apulia it has been found in: Saline Punta della Contessa (Brindisi), Saline Margherita di Savoia (Foggia), Foce del Fiume Biferno (Foggia). Relevés from literature: Taffetani & Biondi 1989, Table 11, rel. 1-8. Our unpublished relevés: Sciandrello & Tomaselli 2007, Table 1, rel. 97-103; Sciandrello & Tomaselli 2011, Table 1, rel. 104-105. Agropyro scirpei-Inuletum crithmoidis Brullo in Brullo et al.1988 Syn.: Agropyro elongati-Inuletum crithmoidis BraunBlanquet (1931) 1952. Diagnostic taxa: Limbarda crithmoides (L.) Dumort. (=Inula crithmoides L.), Elytrigia scirpea (Willd.) Pax. Structure and ecology: This association is one of the most mature in the brackish marsh environments. It occurs in the highest and peripheral parts, generally not subject to flooding. Physiognomically it is distinguished by the dominance of Limbarda crithmoides with Elytrigia scirpea and many other halophytes. On more mature soils, the association is replaced by shrub communities belonging to Oleo-Ceratonion. In northern Mediterranean areas, this vegetation is replaced by Agropyro elongati-Inuletum crithmoidis Braun-Blanquet (1931) 1952. Distribution: this association is distributed in SMediterranean areas. In Apulia it has been found in: Torre Canne (Brindisi), Saline Punta della Contessa (Brindisi), Le Cesine (Lecce), Saline Margherita di Savoia (Foggia), Foce Carapelle (Foggia), Foce Candelaro (Foggia). Relevés from literature: Biondi & Casavecchia 2010, Table 14, rel.1-10. 65 Our unpublished relevés: Tomaselli et al., 2011, Table 1, rel. 106-114; Sciandrello & Tomaselli 2007, Table 1, rel. 115-116. V) Alliance Suaedion brevifoliae Br.-Bl. et O. de Bolòs 1958 (Table 4) The Suaedion brevifoliae, groups of chamaephytic succulent halo-nitrophilous communities, are located in the higher parts of the salt marshes, only occasionally subject to flooding. Halimiono portulacoidis-Suaedetum verae Molinier & Tallon 1970 corr. Géhu in Géhu & al. 1984 Syn.: Suaedetum fruticosae Br.Bl. 1952; ObionetoSuaedetum fruticosae Mol. et Tall. (1969) 1970. Diagnostic taxa: Halimione portulacoides (L.) Aellen, Suaeda vera J.F. Gmel. Structure and ecology: Halo-nitrophilous shrub vegetation, which develops in the most elevated parts of salt marshes, with high levels of organic substance, on silty and clayey soils, only exceptionally subject to submersion (Brullo et al. 2000; Frondoni & Iberite 2002). It tends to form dense bushes dominated by Suaeda vera and Halimione portulacoides. Distribution: widely distributed in Mediterranean areas. In Apulia it has been found in Lago Salinella (Taranto), Lago Salso (Foggia), Palude la Vela (Taranto), Foce del F. Carapelle (Foggia), Saline Margherita di Savoia (Foggia), Palude Frattarolo (Foggia). Relevés from literature: Géhu et al. 1984, Table 19; Corbetta et al., 1989, Table 14, rel. 1-2; Corbetta et al., 2006, Table 5, rel. 10, 20-21, 24-25. Our unpublished relevés: Tomaselli 2010, Table 1, rel. 117-122; Sciandrello & Tomaselli 2011, Table 1, rel. 123-132. Suaedetum verae Br.-Bl. ex O. Bolòs & Molinier 1958 Syn.: Suaedetum fruticosae Br.Bl. 1952; Halimiono portulacoidis-Suadetum verae (Corbetta et al., 1989; Corbetta et al. 2006) Diagnostic taxa: Suaeda vera J.F. Gmel. Structure and ecology: Halo-nitrophilous shrub vegetation, which develops in the most elevated parts of salt marshes, with high levels of organic substance and anthropic pressures, on silty and clayey soils, only exceptionally subject to submersion. It tends to form dense bushes dominated by Suaeda vera. This vegetation replaces the Halimiono portulacoidis-Suaedetum verae on clay soils and especially in more xeric (dry) zones. In Italy, Suaeda vera communities have been usually ascribed to Halimiono portulacoidis-Suaedetum verae, although even in absence of Halimione portulacoides in the surveyed plant communities (Frondoni & Iberite 2002; Corbetta 2006; Maiorca et al. 2007). On the basis of our in field surveys, the two associations, with distinct ecological requirements, have to be well differentiated and the Suaeda vera (almost monophytic) communities, growing on soils with high organic matter and in disturbed sites, have to be referred to the Suaedetum verae. Unauthenticated Download Date | 3/16/17 6:12 AM 66 Distribution: widely distributed in the Mediterranean region (Trinajstić 2009). In Apulia it has been found in Saline Margherita di Savoia (Foggia), Lago Salinella (Taranto). Relevés from literature: Corbetta et al., 1989, Table 14, rel. 3; Corbetta et al. 2006, Table 5, rel. 1-9,1119, 22-23, 26. Conclusion The Salicornietea fruticosae class is represented in Apulia by well-differentiated phytosociological units, each of them characterized by specific ecological features (Table 4). Our results show that the surveyed alliances comprising halophytic plant communities are distributed mainly in 3 spatially differentiated zones, following the flooding time and the soil salinity (Fig. 4). The lower zone (regularly flooded) is characterized by Sarcocornion alpini and Arthrocnemion glauci communities, the middle zone (irregularly flooded) by Salicornion fruticosae, and the upper zone (rarely flooded) is colonized by Inulion crithmoidis and Suaedion brevifoliae communites. In addition to the phytosociological interest, it is worth noting that all the surveyed associations could be assigned to the Annex I of the Dir. 92/43/EEC habitat type “1420: Mediterranean and thermo-Atlantic halophilous scrubs (Sarcocornietea fruticosi)”. Numerical analysis of the vegetation data allowed us to properly assign certain associations and/or plant communities drawn from the literature data, thus solving inconsistencies emerged from comparison with previous contributions and providing a comprehensive picture of the Sarcocornietea fruticosae class in Apulia. Moreover the phytosociological study allowed to highlight two new interesting plant communities for Apulia: Arthrocnemo-Halocnemetum strobilacei and Puccinellio convolutae-Arthrocnemetum macrostachyi limonietosum bellidifolii. Coastal marshland habitat types are among the most endangered environments in Europe, especially in the Mediterranean region, where they are undergoing rapid anthropic development with a decreasing surface area, fragmentation and isolation (Valdemoro et al. 2007; Levin et al. 2009). Watercourse engineering, draining of the marshes, abandonment of mowing, overgrowth by woody species, spreading of neophytes are the main decline phenomena affecting these habitats. These fragile environments deserve protection and detailed study and monitoring activities to enable effective conservation measures. Field observations allowed us to undertake a global assessment of the conservation status of these salttolerant communities. In the study area, due to the intensive agricultural activities and the presence of a network of infrastructures linked to specific production sites, these habitat types have undergone reduction and fragmentation of their distribution area. So, in most cases, they do not form well-structured vegetation belts, but rather a mosaic of fragmented patches (Tomaselli et al. 2012). The most threatened communities of the class S. Sciandrello & V. Tomaselli are: Agropyro scirpei-Inuletum crithmoidis, LimonioArtemisietum coerulescentis and Halimiono-Suaedetum verae, normally spreading in the upper part of the salt marshes and therefore subjected to more intensive anthropogenic pressures. In the distribution area of the surveyed plant communities, the occurrence of several rare species, such as Bassia hirsuta (L.) Asch., Suaeda splendens (Pourr.) Gren. & Godr., Damasonium alisma Mill. subsp. bourgaei (Coss.) Maire, Limonium avei (De Not.) Brullo & Erben, Limonium bellidifolium (Gouvan) Dumort, Cornucopiae cucullatum L. (Sciandrello & Tomaselli 2011), has been recorded. Finally, a significant anthropogenic threat recorded in the salt marshes of Apulia is related to the presence of alien species, such as Sonchus bulbosus (L.) Kilian & Greuter., that represents a serious threat to the integrity of halophilous community in Italy (Cutini et al. 2010). Acknowledgements The authors would like to thank Dr. Giuseppe Silletti and Dr. Ruggiero Matera, Corpo Forestale dello Stato for logistical support, Dr. Giuseppe Albanese, Oasi Lago Salso (Fg), for logistical support and useful information, and two anonymous referees for helpful comments on the previous version of this paper. References Andreucci F. 2004. La vegetazione della laguna di Orbetello (Toscana, Grosseto). Fitosociologia 41: 31–49. Álvarez-Rogel J., Jiménez-Càrceles F.J., Roca M.J. & Ortiz R. 2007. Changes in soils and vegetation in a Mediterranean coastal salt marsh imparte by human activies. Estuar Coast Shelf S 73: 510–526. Arrigoni P.V., Nardi E. & Raffaelli M. 1985. La vegetazione del parco naturale della Maremma (Toscana). Arti Grafiche Giorgi & Gambi, Firenze, 39 pp. Cutini M., Agostinelli E., Acosta T.R.A. & Molina J.A. 2010. Coastal salt-marsh zonation in Tyrrhenian central Italy and ist relationship with other Mediterranean wetlands. Plant Biosyst 144: 1–11. Beccarisi L., Medagli P., Minonne F., Zuccarello V. & Marchiori S. 2003. Descrizione ed analisi fitostorica della vegetazione del sito di Torre Guaceto. Thalassia Salentina 27: 99–116. Biondi E. 1999. Diversità fitocenotica degli ambienti costieri italiani. Atti XIII Convegno G. Gadio, suppl. Boll. Museo Civ. Sc. Nat. Di Venezia 49: 39–105. Arsenale Ed. Biondi E. & Bagella S. 2005. Vegetazione e paesaggio vegetale dell’arcipelago di La Maddalena (Sardegna nord-orientale). Fitosociologia 42(2) Suppl. 1: 3–99. Biondi E. & Casavecchia S. 2010. The halophilous retro-dune grasslands of the italian adriatic coastline. Braun-Blanquetia 46: 11–127. Biondi E., Casavecchia S. & Guerra V. 2006. Analysis of vegetation diversity in relation to the geomorphogical characteristics in the Salento coasts (Apulia-Italy). Fitosociologia 43: 25–38. Blasi C., Michetti L., Del Moro M.A., Testa O. & Teodonio L. 2007. Climate change and desertification vulnerability in Southern Italy. Phytocoenologia 37: 495–521. Braun-Blanquet J. 1964. Pflanzensoziologie. Grundzüge der Vegetationskunde. 3. Aufl. – Springer Verl., Wien & New York, 330 pp. Unauthenticated Download Date | 3/16/17 6:12 AM The Salicornietea fruticosae class in Apulia (Italy) Brullo S. & Di Martino A. 1974. Vegetazione dell’Isola Grande dello Stagnone (Marsala). Boll. St. Inform. Giard. Col. Palermo 26: 15–62. Brullo S., Guarino R. & Ronsisvalle G. 2000. La vegetazione del litorale di Manfria, presso Gela (Sicilia), area soggetta a vincolo archeologico. Arch. Geobot. 4: 91–107. Brullo S. & Furnari F. 1976. Le associazioni vegetali degli ambienti palustri costieri della Sicilia. Not. Fitosoc. 11: 1–43. Brullo S., De Santis C., Furnari F., Longhitano N. & Ronsisvalle G.A. 1988. La vegetazione dell’oasi della foce del Simeto (Sicilia orientale). Braun-Blanquetia 2: 165–188. Brullo S. & Furnari F. 1990. La vegetazione costiera della Cirenaica. Boll. Acc. Gioenia Sc. Nat. 21 (334): 31–117. Brullo S., Minissale P., Giusso del Galdo G., Siracusa G. & Spampinato G. 2002. Considerazioni sintassonomiche e fitogeografiche sulla vegetazione della Sicilia. Boll. Acc. Gioenia. Sci. Nat. 35 (361): 325–359. Caneva G., De Marco G. & Mossa L. 1981. Analisi fitosociologica e cartografia della vegetazione (1:25.000) dell’isola di S. Antioco. C.N.R. AQ/1/124, pp. 1–59. Caniglia G., Conti G., Fusco M., Anoè N. & Zanobi A. 1997. Confronto su base vegetazionale tra due barene della Laguna di Venezia. Fitosociologia 34: 111–119. Clarke K.R. 1993. Non parametric multivariate analysis of changes in community structures. Australian J. Ecol. 18: 117–143. Conti F., Manzi A. & Pedrotti F. 1997. Liste rosse Regionali delle Piante d’Italia. WWF-SBI, Camerino. Conti F., Abbate G., Alessandrini A. & Blasi C. (eds) 2005. An annotated checklist of the Italian vascular flora. Palombi, Roma, 420 pp. Corbetta F. 1970. Lineamenti della vegetazione macrofitica dei Laghi di Lesina e Varano. Giorn. Bot. Ital. 104(3): 165–191. Corbetta F. 1976. Lineamenti vegetazionali della Sacca di Bellocchio (Foce del Reno). Ricerche di Biologia della Selvaggina 7: 247–270. Corbetta F. & Lorenzoni G.G. 1974. La vegetazione degli stagni del Golfo di Oristano (Sardegna). Laboratorio di zoologia applicata alla caccia, Bologna: pp. 271–315. Corbetta F., Gratani L., Morioni M. & Pirone G. 1989. Lineamenti vegetazionali e caratterizzazione ecologica delle spiagge dell’Arco Jonico da Taranto alla Foce del Sinni. Coll. Phytosoc. 19: 461–521. Corbetta F., La Monica M., Pirone G., Burri E. & Ivona A. 2006. La vegetazione delle Saline di Margherita di Savoia (Puglia). Micol. e Veget. Medit. 21(2): 141–156. Cutini M., Agostinelli E., Acosta T.R.A. & Molina J.A. 2010. Coastal salt-marsh zonation in Tyrrhenian central Italy and ist relationship with other Mediterranean wetlands. Plant Biosyst. 144: 1–11. De Marco G., Dinelli A. & Mossa L. 1980. Aspetti della vegetazione costiera dell’isola di S. Antioco (Sardegna sudoccidentale). Ann. Bot. 38: 173–191. De Martis G. & Serri G. 2009. L’analisi fitosociologica della vegetazione per il monitoraggio degli habitat nel Parco Naturale Regionale Molentargius-Saline (Sardegna meridionale). Primi risultati. Inf. Bot. Ital. 41: 293–301. El-Sheikh M.A.E., El-Ghareeb R.M. & Testi A. (2006) Diversity of plant communities in coastal salt marshes habitat in Kuwait. Rendiconti Lincei 17: 311–331. Ferrari C., Gerdol R. & Piccoli F. 1995. The halophilous vegetation oft he Po Delta (northern Italy). Vegetatio 61: 5–14. Filigheddu R., Farris E. & Biondi E. 2000. The vegetation of S’Ena Arrubia lagoon (centre-western Sardionia). Fitosociologia 37: 39–59. Frondoni R. & Iberite M. 2002. The halophile vegetation of the sedimentary coast of Lazio (central Tyrrhenian district, Italy). Plant Biosyst. 136: 49–68. Géhu J.M. & Biondi E. 1994. Assai de typologie phytosociologique des habitats et des vègètations halophiles des littoraux sedimentaires pèrimèditerranèens et thermo-atlantiques. Fitosociologia 30: 201–212. 67 Géhu J.M., Costa M., Scoppola A., Biondi E., Marchiori S., Peris J.B., Frank J., Caniglia G. & Veri L. 1984. Essai synsystématique et synchorologique sur les végétations littorales italiennes dans un but conservatoire. Doc.Phytosoc N. S. 8: 393– 474. Giusso del Galdo G., Marcenò C., Musarella C.M. & Sciandrello S. 2008. La vegetazione costiera della R.N.O. “Torre Salsa” (Siculiana – AG). Inf. Bot. Ital. 40: 73–89. Greuter W., Burdet H.M. & Long G. (eds) 1984–1989, 2009. MedChecklist. Vols. 1, 2, 3 & 4. Gèneve: Conserv. Jard. Bot. Géneve. Levin N., Elron E. & Gasith A. 2009. Decline of wetland ecosystems in the coastal plain of Israel during the 20th century: inplications for wetland conservation and management. Landsc. And Urban Plan. 92: 220–232. Maiorca G., Spampinato G., Crisafulli A & Cameriere P. 2007. Flora vascolare e vegetazione della Riserva Naturale Regionale “Foce del Fiume Crati” (Calabria, Italia meridionale). Webbia 62: 121–174. Mariotti M.G., Braggio Morucchio G., Cornara L. & Placereani S. 1992. Studio fitosociologico e palinologico della vegetazione attuale e del passato a Torre Guaceto (Puglia, Italia meridionale). Candollea 47: 31–60. Minissale P. & Sciandrello S. 2010. Flora e vegetazione terrestre della Riserva Naturale di Vendicari (Sicilia sud-orientale). Ente Fauna Siciliana 12: 145–208. Mucina L. 1997. Conspectus of Classes of European Vegetation. Folia Geobot. Phytotax. 32: 117–172. Pandža M., J. Franji’ & Ž. Škvorc 2007. The salt marsh vegetation on the East Adriatic coast. Biologia 62: 24–31. Piccoli F., Merloni N. & Pellizzari M. 1994. The vegetation of the Comacchio Saltern (Northern Adriatic coast, Italy). Ecologia Mediterranea 20(3-4): 85–94. Pignatti S. 1966. La vegetazione alofila della laguna veneta. Mem. Ist. Ven. Sc. Lett. Arti 33: 1–174. Pirone G. 1988. La vegetazione alofila residua alle foci del fiume Saline e del torrente Piomba (Abruzzo-Itralia). Doc. Phytosoc. 11: 447–458. Pirone G. 1995. La vegetazione alofila della costa abruzzese (Adriatico centrale). Fitosociologia 30: 233–256. Pirone G., Corbetta F., Frattaroli A.R. & Ciaschetti G. 2001. Aspetti della vegetazione costiera d’Abruzzo. Biogeographia 22: 169–191. Podani J. 2001. SYN-TAX 2000, computer program for multivariate analysis in ecology and taxonomy. Scientia Publishing, Budapest. Poldini L., Vidali M. & Fabiani M.L. 1999. La vegetazione del litorale sedimentario del Fiuli-Venezia Giulia (NE Italia) con riferimenti alla regione alto-adriatica. Studia Geobotanica 17: 3–68. Rivas-Martínez S. 2004. Bioclimatic Map of Europe: Bioclimates, scale 1:16 mill. Cartographic Service, University of León. Rivas-Martínez S., Fernández-González F., Loidi J., Lousã M. & Penas A., 2001. Syntaxonomical checklist of vascular plant communities of Spain and Portugal to association level. Itin. Geobot. 14: 5–341. Sciandrello S. 2007. La vegetazione alofila di Piana del Signore presso Gela (Sicilia meridionale): proposte di conservazione e gestione del biotopo. Inf. Bot. Ital. 39: 129–141. Sciandrello S. & Tomaselli V. 2011. Distribution and ecology of Cornucopiae cucullatum L. (Poaceae) in the Mediterranean area and evaluations of its conservation status in Italy. Acta Bot. Gallica 158: 401–407. Taffetani F. & Biondi E. 1989. La vegetazione del litorale molisano e pugliese tra le foci dei fiumi Biferno e Fortore (Adriatico centro-meridionale). Coll. Phytosoc. 18: 323–350. Tomaselli V., Di Pietro R. & Sciandrello S. 2011. Distribution, structure and ecology of coastal wetlands plant communities in southern Apulia (Italy). Biologia 66: 1027–1043. Tomaselli V., Tenerelli P. & Sciandrello S. 2012. Mapping and quantifying habitat fragmentation in small coastal areas: a case study of three protected wetlands in Apulia (Italy). Environ. Monit. Assess. 184: 693–713. Tomaselli V., Sciandrello S., Dibitonto P., Wagensommer R. P., Urbano M., Calabrese I.T., Cimmarusti G., Garziano G. & Unauthenticated Download Date | 3/16/17 6:12 AM 68 Di Pietro R. 2009a. Analisi della biodiversità vegetale e cartografia della vegetazione, degli habitat e dell’uso del suolo della riserva naturale statale di “Torre Guaceto” (Brindisi – Puglia). Quad. Bot. Amb. Appl. 20(2): 57–74. Tomaselli V., Urbano M., Sciandrello S., Wagensommer R. P., Costanzo E., Albano A., Medagli P., Mele C. & Di Pietro R. 2009b. Analisi del paesaggio vegetale ed agricolo della Riserva Naturale Statale del Parco Naturale Regionale “Saline di Punta della Contessa” (Brindisi – Puglia). Cartografia della vegetazione, degli habitat e dell’uso del suolo. Quad. Bot. Amb. Appl. 20(2): 103–125. Trinajstić I. 2009. Suaeda vera Forsk. ex J. F. Gmelin in the plant cover of Croatia. Natura Croatica 18: 387–391. Tutin T.G., Heywood W.H., Burges N.A., Valentine D.H., Walters S.M. & Webb D.A. (eds) 1964 – 1993. Flora Europaea. Vols. 1–5. Cambridge: Cambridge Univ. Press. S. Sciandrello & V. Tomaselli Valdemoro H.I., Sanchez-Arcilla A. & Jiménez J.A. 2007. Coastal dynamics and wetlands stability. The Ebro delta case. Hydrobiologia 577: 17–29. Valsecchi F. 1964. Ricerche sulla vegetazione littorale della Sardegna. IV: La vegetazione dello Stagno di Calich (Sardegna nord-occidentale). Ann. Bot. 28: 157–218. Valsecchi F. 1972. La vegetazione dello stagno di S’Ena Arubbia nel Golfo di Oristano. Boll. Soc. Sarda Sci. Nat. 10: 3–21. Van der Maarel E. 1979. Transformation of cover-abundance values in phytosociology and its effects on community similarità. Vegetatio 39: 97–114. Viciani D. & Lombardi L. 2001. La vegetazione della palude di Orti-Bottagone (Piombino, Toscana meridionale) e la sua importanza botanica ai fini conservazionistici. Parlatorea 5: 101–118. Received January 22, 2013 Accepted May 28, 2013 Appendix 1 Data and localities of relevés (Appendix 1, Table 1) Rel. 1-10, 18.06.2007, Saline Punta della Contessa (Brindisi), Sciandrello & Tomaselli (Puccinellio-Sarcocornietum alpini); Rel. 11-12, 19.06.2007, Torre Guaceto (Brindisi), Sciandrello & Tomaselli (Sarcocornietum alpini); Rel. 13, 20.11.2007, Saline Punta della Contessa (Brindisi), Sciandrello & Tomaselli (Sarcocornietum alpini); Rel. 14, 19.07.2007, Le Cesine (Lecce), Sciandrello & Tomaselli (Sarcocornietum alpini); Rel. 15-17, 18.07.2007, Torre Canne (Brindisi), Sciandrello & Tomaselli (Puccinellio-Sarcocornietum alpini); Rel. 18-26, 15.05.2011, Palude di Frattarolo (Capitanata, Foggia), Sciandrello & Tomaselli, (Puccinellio-Sarcocornietum alpini); Rel. 27-30, 13.05.2011, Foce del Carapelle (Capitanata, Foggia), Sciandrello & Tomaselli (Puccinellio-Sarcocornietum alpini); Rel. 31-35, 14.05.2011, Area umida Azienda ittica Carapelle (Capitanata, Foggia), Sciandrello & Tomaselli, (Puccinellio-Sarcocornietum alpini); Rel. 36-37, 01.09.2011, Zapponeta (Capitanata, Foggia), Sciandrello & Tomaselli (Puccinellio-Sarcocornietum alpini); Rel. 38-42, 02.09.2011, Almadannata (Saline Margherita di Savoia), Sciandrello & Tomaselli, (Puccinellio-Sarcocornietum alpini); Rel. 43-45, 15.05.2011, Palude di Frattarolo (Capitanata, Foggia), Sciandrello & Tomaselli (Puccinellio-Sarcocornietum alpini); Rel. 46-49, 17.07.2007, Saline Punta della Contessa (Brindisi), Sciandrello & Tomaselli (Junco subulati-Sarcocornietum fruticosae); Rel. 50, 18.06.2007, Saline Punta della Contessa (Brindisi), Sciandrello & Tomaselli (Junco subulati-Sarcocornietum fruticosae); Rel. 51-52, 10.06.2010, Palude la Vela (Taranto), Tomaselli (Junco subulati-Sarcocornietum fruticosae); Rel. 53-56, 04.10.2007, Saline Contessa (Brindisi), Sciandrello & Tomaselli (Arthrocnemo-Juncetum subulati); Rel. 57, 02.10.2007, Torre Canne (Brindisi), Sciandrello & Tomaselli (Arthrocnemo-Juncetum subulati); Rel. 58, 12.05.2011, Foce Fiume Candelaro, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi subass. limonietosum bellidifolii); Rel. 59, 12.05.2011, Foce Fiume Candelaro, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi); Rel. 60-61, 12.05.2011, Foce Fiume Candelaro, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi subass. limonietosum bellidifolii); Rel. 62, 13.05.2011, Saline Margherita di Savoia, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi); Rel. 63, 13.05.2011, Saline Margherita di Savoia, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi subass. limonietosum bellidifolii); Rel. 64, 13.05.2011, Saline Margherita di Savoia, Sciandrello & Tomaselli (PuccinellioArthrocnemetum macrostachyi); Rel. 65-66, 13.05.2011, Saline Margherita di Savoia, Sciandrello & Tomaselli (PuccinellioArthrocnemetum macrostachyi subass. limonietosum bellidifolii); Rel. 67, 13.05.2011, Saline Margherita di Savoia, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi); Rel. 68-69, 13.05.2011, Vasca Paradiso, Saline Margherita di Savoia, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi); Rel. 70-73, 13.05.2011, Foce Carapelle, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi); Rel. 74, 14.05.2011, Foce Carapelle, Canale Pertuso, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi); Rel. 75, 14.05.2011, Foce del Candelaro, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi subass. limonietosum bellidifolii); Rel. 76, 13.05.2011, Saline Margherita di Savoia, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi); Rel. 77-78, 13.05.2011, Saline Margherita di Savoia, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi subass. limonietosum bellidifolii); Rel. 79-83, 02.09.2011, Almadannata, Saline Margherita di Savoia, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi); Rel. 84, 02.09.2011, Almadannata, Saline Margherita di Savoia, Sciandrello & Tomaselli (PuccinellioArthrocnemetum macrostachyi subass. limonietosum bellidifolii); Rel. 85, 02.09.2011, Almadannata, Saline Margherita di Savoia, Sciandrello & Tomaselli (Puccinellio-Arthrocnemetum macrostachyi subass. limonietosum bellidifolii; Rel. 8696, Foce Carapelle, Capitanata, Sciandrello & Tomaselli (Arthrocnemo-Halocnemetum strobilacei); Rel. 97-99, 17.07.2007, Saline Punta della Contessa (Brindisi), Sciandrello & Tomaselli (Limonio narbonensis-Artemisietum coerulescentis); Rel. 100-103, 04.10.2007, Saline Punta della Contessa (Brindisi), Sciandrello & Tomaselli (Limonio narbonensis-Artemisietum coerulescentis); Rel. 104-105, 01.09.2011, Saline Margerita di Savoia, Sciandrello & Tomaselli (Limonio narbonenseArtemisietum coerulescens); Rel. 106-107, 20.11.2007, Saline Punta della Contessa (Brindisi), Sciandrello & Tomaselli (Agropyro-Inuletum crithmoidis); Rel. 108, 19.07.2007, Le Cesine (Lecce), Sciandrello & Tomaselli (Agropyro-Inuletum crithmoidis); Rel. 109, 21.05.2008, Le Cesine (Lecce), Sciandrello & Tomaselli (Agropyro-Inuletum crithmoidis); Rel. 110112, 18.07.2007, Torre Canne (Brindisi), Sciandrello & Tomaselli (Agropyro-Inuletum crithmoidis); Rel. 113-114, 02.10.2007, Unauthenticated Download Date | 3/16/17 6:12 AM The Salicornietea fruticosae class in Apulia (Italy) 69 Torre Canne (Brindisi), Sciandrello & Tomaselli (Agropyro-Inuletum crithmoidis); Rel. 115, 13.05.2011, Foce del Carapelle, Sciandrello & Tomaselli (Agropyro-Inuletum crithmoidis); Rel. 116, 14.05.2011, Foce Candelaro, Sciandrello & Tomaselli (Agropyro-Inuletum crithmoidis); Rel. 117-118, 20-06-2010, Lago Salso, Tomaselli (Halimiono-Suaedetum verae); Rel. 119122, 10.06.2010, Palude la Vela (Taranto), Tomaselli. Halimiono-Suaedetum verae; Rel. 123-127, 12.05.2011, Palude di Frattarolo, Sciandrello & Tomaselli (Halimiono-Suaedetum verae); Rel. 128, 13.05.2011, Almadannata, Saline Margherita di Savoia, Sciandrello & Tomaselli (Halimiono-Suaedetum verae); Rel. 129-130, 13.05.2011, Foce Carapelle, Sciandrello & Tomaselli (Halimiono-Suaedetum verae); Rel. 131-132, 03.09.2011, Palude Frattarolo, Sciandrello & Tomaselli (HalimionoSuaedetum verae). Unauthenticated Download Date | 3/16/17 6:12 AM