Original scientific article	UDC 574.3:581.9(252.6)
Received: 2006-11-09
VEGETATION OF TALL RUSH SALTMARSHES (JUNCETEA MARITIMAE) AND SALTMARSH SCRUBS (ARTHROCNEMETEA FRUTICOSAE) ON THE SLOVENIAN SEACOAST
Mitja KALIGARIČ
University of Maribor, Faculty of Natural Sciences and Mathematics, Department of Biology, SI-2000 Maribor, Koroška 160
and
University of Primorska, Science and Research Centre of Koper, Institute for Biodiversity Studies, SI-6000 Koper, Caribaldijeva 18
E-mail: mitja.kaligaric@uni-mb.si
Sonja ŠKORNIK
University of Maribor, Faculty of Natural Sciences and Mathematics, Department of Biology, SI-2000 Maribor, Koroška 160
ABSTRACT
Halophyte vegetation of Slovenian sedimentary seacoast was studied according to Braun-Blanquet method. 140 collected relevés were analysed by cluster analysis and five main clusters were separated. Relevés from the first, fourth and fifth clusters were further elaborated in this study. The Juncus maritimus-dominated tall rush saltmarshes of the class Juncetea maritimi were classified within two associations: Limonio-Puccinellietum represented hygro-philous stands, exposed to tide, while the association Juncetum maritimi-acuti was represented with more closed stands with higher species richness, thriving on rather drier sites. Within the saltmarsh scrubs of the class Arthrocne-metea fruticosi, 3 associations, following the declining moisture/salinity gradient were recognized: Puccinellio-Arthrocnemetum, Puccinellio-Halimionetum and Limonio-Artemisietum.
Key words: phytosociology, halophyte vegetation, classification, North Adriatic
VEGETAZIONE A GIUNCO Dl ACQUE SALMASTRE (JUNCETEA MARITIMAE) E PRATERIE ALOFILE (ARTHROCNEMETEA FRUTICOSAE) SULLA COSTA SLOVENA
SINTESI
La vegetazione alofila della costa marina sedimentaria in Slovenia è stata studiata secondo il metodo di Braun-Blanquet. Centoquaranta rilievi sono stati analizzati con l'aiuto della analisi numeriche e cinque grandi gruppi sono stati separati. I rilievi appartenenti al primo, quarto e quinto gruppo sono stati ulteriormente elaborati durante lo studio. La vegetazione di acque salmastre dominata da Juncus maritimus, della classe Juncetea maritime, è stata classi-ficata in due associazioni: Limonio-Puccinellietum con banchi igrofili, esposti alla marea, e Juncetum maritimi-acuti con banchi piu ristretti con una diversita di specie piu alta, sprosperanti in siti piuttosto secchi. AU'interno delle pra-terie alofile della classe Arthrocnemetea fruticosi, 3 associationi sono state riconosciute seguendo il gradiente umi-dita/salinita declinante: Puccinellio-Arthrocnemetum, Puccinellio-Halimionetum e Limonio-Artemisietum.
Parole chiave: fitosociologia, vegetazione alofila, classificazione, Adriatico settentrionale
INTRODUCTION
The Slovenian short seacoast is under heavy pressure of urbanization, leisure activities and industry (Port of Koper). Its special feature is the geological bedrock, being the calcareous sandstone - Eocene flysch substrate. This substrate almost perfectly matches the territory of Slovenia; only a small part extends to the Italian territory. Flysch substrate results in dense hydrological system above ground due to its impenetrable properties. Three larger streams/rivers have their mouths in the Gulf of Trieste: the Rizana, Strunjanski potok and Dragonja. Alluvial deposits in the mouths resulted in salt-marshes, where different vegetation types probably occurred before human influence. After the Roman period and afterwards these alluvial coastal regions have been either conversed to salt pans either drained (Darovec, 1992). The present situation is only a transitional stage in abandonment and creation of new habitats on the seacoast. The dynamics have always been fast and enabled assemblages of different vegetation types on the seacoast. Abiotic conditions - shallow coast and large tide area -are in favour of assemblage of the various types of halo-phyte vegetation, probably similar to the natural types, developed before the human interference. The halo-phyte vegetation is interesting and important from the conservational perspective, as it is threatened by all factors mentioned above. Many coastal habitats with different types of halophyte vegetation are listed in Annexes of the EU Directive on the conservation of natural habitats and of wild fauna and flora (Council Directive 92/43/EEC, 1992).
Syntaxonomical classifications of the Slovenian halophyte vegetation are based on the relevés taken by M. Kaligaric in the years 1984-87 and in 1998 and 1999, published so far only for annual pioneer vegetation (Salicornietea) and Spartina maritima-dominated swards (Spartinetea) (Kaligaric & Skornik, 2006). Some of those relevés have been taken in consideration also within the last complete revision of the North Adriatic halophyte vegetation (Poldini et al., 1999). This revision considered the global revision of Mediterranean halo-phyte vegetation by Géhu et al. (1984), Rivas-Marttnez (1990), Géhu & Biondi (1995, 1996), Mucina (1997) and Géhu (1999). Recent studies of halophyte vegetation, to the exclusion of Tyrrhenian district (Latium), have been made by Iberite & Frondoni (1997) and Frondoni & Iberite (2002). Both consider the already revised syntax-onomy.
The first phytosociological assessment of the North Adriatic halophyte vegetation was made much earlier by Béguinot (1941). The most profound research was carried out by Pignatti (1966) and followed by Fornaciari (1968). The Pignatti's syntaxonomical scheme (e.g. macroassociation "Limonietum venetum") was adopted also for Slovenian halophyte vegetation in some short contri-
butions about Secoveljske soline (Kaligaric & Tratnik, 1981; Kaligaric & Wraber, 1988), Strunjan (Sajna & Kaligaric, 2005) and Skocjanski zatok (Kaligaric, 1997, 1998). The Pignatti's syntaxonomical units are also used for the threat status of halophyte flora and vegetation of the Slovenian seacoast (Kaligaric, 1996).
The elaboration of collected relevés from the Slovenian seacoast with classification methods and determination of major groups of vegetation (classes and orders) on the basis of classification methods and characteristic species has been carried out already by Kaligaric & Skornik (2006). The aim of this study was elaboration of the Juncus maritimus-dominated tall rush saltmarshes of the class Juncetea maritimi and the saltmarsh scrubs of the Arthrocnemetea fruticosi class.
MATERIAL AND METHODS Study area
The Slovenian seacoast consists of flysch cliffs with scarce halophyte vegetation and sedimentary coast, mainly converted to salt pans or dried. In some parts, natural patches of coast still exist, while in other parts the sedimentary coast is even artificially enlarged (like soil deposits, etc.). In some parts (Secoveljske soline), salt pans were abandoned and halophyte vegetation spread out. Locations (Fig. 1) were relevés have been sampled are: Secoveljske soline (mouth of the Dragonja river, Fontanigge, Lera, San Giorggio channel), Strunjan-ske soline (Stjuza lagoon coast, salt pans), Skocjanski zatok, mud deposits and coastal grasslands near Sv. Katarina/Ankaran (Kaligaric & Skornik, 2006).
Fig. 1: Map of Slovenian seacoast with locations of the collected relevés of halophilous vegetation. Sl. 1: Karta slovenske morske obale z lokalitetatmi popisov halofitne vegetacije.
Sampling methods
Using the standard Braun-Blanquet procedure (Braun-Blanquet, 1964; Westhoff & van der Maarel, 1973; Dierschke, 1994), relevés were taken during the years 1984-87 and 1998-1999. In the initial procedure, all the relevés were classified using the SYN-TAX 2000 software package (Podani, 2001). A hierarchical classification algorithm (Complete linkage, Eucledian distance) based on quantitative data was employed. From the resulting dendrogram (Fig. 2), three clusters were elaborated in previous study (Kaligaric & Skornik, 2006); all the other clusters were elaborated within this study. Division to classes and orders were based on main clusters, further subdivision to associations was carried out with the help of characteristic species (their frequencies and dominance) of the syntaxa.
Taxonomic nomenclature followed Martincic et al. (1999) except for taxon Halimione portulacoides, syn-taxonomic nomenclature followed Poldini et al. (1999).
RESULTS AND DISCCUSION Classification of the relevés
The 140 relevés were classified in 5 main clusters according to species composition and abundance, considering all the species of equal importance (Fig. 2). The most diverse was the first main cluster (No.1), which could be further divided into 7 sub-clusters (1A-1G). The first one (A, relevés 1-4) was represented by floristically poor Salsola soda dominated stands on dry trampled muddy soils with stones, where Elytrigia atherica and Atriplex prostrata indicate relatively dry conditions and lower salinity (Kaligaric & Skornik, 2006). This group of relevés was classified within the class Cakiletea mariti-mae (Kaligaric & Skornik, 2006). The third sub-cluster (1C, relevés 15-20) was characterized by dominance of grass Spartina maritima, and it was also elaborated in previous study (Kaligaric & Skornik, 2006). Other five sub-clusters (1B, 1D, 1F, 1G and part of 1 E) were Juncus maritimus-dominated saltmarshes, relatively rich in various halophyte hygrophilous species. Among them were also halophytes such as Phragmites communis and characteristic species of the class Juncetea maritimae, Carex extensa and Plantago cornuti, which are not present in other halophilous vegetation types. These relevés (relevés 5-14 and 21-44) were classified within the class Juncetea maritimi (Kaligaric & Skornik, 2006).
The second main cluster (No. 2, relevés 45-67) and the third main cluster (No. 3, relevés 68-86), characterized by Salicornia europaea-dominated stands on mud-
flats or salt pans, were classified within the class Thero-Salicornietea and elaborated in the study by Kaligaric & Skornik (2006).
The fourth main cluster (No. 4, relevés 87-137) had many sub-clusters on a lower level of dissimilarity. The main characteristics were scarce presence of pioneer annual halophytes (Salicornia and Suaeda) and strong presence and dominance of halophilous scrubs: Ar-throcnemum macrostachyum, Halimione portulacoides, Limonium angustifolium, Artemisia caerulescens, but also Puccinellia palustris, Inula crithmoides, and Aster tripolium. This perennial halophilous vegetation of relatively dry soil was spread predominately on abandoned salt pans or higher levels (above the tidal area). It was classified within the class Arthrocnemetea fruticosi (Kaligaric & Skornik, 2006).
The fifth main cluster with only 3 relevés (No. 5, relevés 138-140) represented Juncus maritimus-domi-nated salt grasslands near Sv. Katarina (Ankaran), which was classified within the class Juncetea maritimi (Kaligaric & Skornik, 2006), despite the presence of some non-halophytic species and its particular species richness.
Within this study, the clusters representing classes Arthocnemetea, Juncetea and Cakiletea will be further elaborated.
The syntaxa were determined mainly on the basis of cluster analysis (Fig. 2), but in some floristically very impoverished relevés the dominance of characteristic species was weighted over the automatic classification given by the expert system.
Vegetation of saltmarsh scrubs
Syntaxonomical classification of halophilous vegetation in the Mediterranean was very variable - from one class (Salicornietea) up to seven classes (Poldini et al., 1999). On the basis of different studies across the Mediterranean, European coastal halophyte vegetation is classified in three classes. The vegetation of perennial halophytes - mainly succulent chamaephytes and nano-phanerophytes, which form more or less densely vegetated saltmarsh scrubs, is classified as Arthrocnemetea fruticosi. The distribution of this class of vegetation is not limited to the Atlantic and Mediterranean coasts, but extends to the continental halophilous vegetation of North Africa and Asia. The order Arthrocnemetalia fruticosi has a Mediterranean and Atlantic distribution. Alliance Arthrocnemion fruticosi is the Arthrocnemum fruti-cosum-dominated vegetation within the area mentioned above.
Fig. 2: Dendrogram: the result of hierarchical classification analysis for 104 relevés of halophilous vegetation of the Slovenian seacoast.
Legend: black circles - ass. Salsoletum sodae; pink circles - ass. Juncetum maritime-acuti; blue circles - ass. Limonio narbonensis-Puccinellietum palustris; orange circles - ass. Puccinelio festuciformis-Arthrocnemetum perennis; green circles - ass. Limonio narbonensis-Artemisietum coerulescentis; violet circles - ass. Puccinellio festuciformis-Arthrocnemetum fruticosi.
Sl. 2: Dendrogram: rezultat hierarhične klasifikacije 104 popisov halofitne vegetacije ob slovenski obali. Legenda: črni krogci - as. Salsolo kali-Cakiletum maritimae; rožnati krogci - as. Juncetum maritime-acuti; modri krogci - as. Limonio narbonensis-Puccinellietum palustris; oranžni krogci - as. Puccinelio festuciformis-Arthro-cnemetum perennis; zeleni krogci - as. Limonio narbonensis-Artemisietum coerulescentis; vijolični krogci - as. Puccinellio festuciformis-Arthrocnemetum fruticosi.
Association Limonio narbonensis-Artemisietum co-erulescentis Horvatic (1933) 1934 corr. Gehu et Biondi 1996 (Tab. 1)
This is the driest association in the humidity/salinity gradient of perennial scrubs, with really dense vegetation cover, and highest species-richness in this series of syntaxa. The presence of halotolerant species, which penetrate from marginal areas outside salty soils, is also characteristic. It was described from Dalmatia by Horvatic in 1934 and registered many times in NE Italy (e.g. Poldini et al., 1999). Halophytes with less extreme adaptations are characteristic of these stands, which could be explained with less extreme site conditions. The summer dry period is also characteristic.
On the Slovenian seacoast, this association occurs predominately in abandoned salt pans, on elevated sites, banks, and closed muddy surfaces with only temporary inundation.
Association Pucclnelllo festuciformis-Halimionetum portulacoidis Géhu, Biondi, Géhu Franck et Costa 1992
(Tab. 2)
This association was described by Ferrari et al. (1985) from Italy. It is characterized by dominance of Halimione portulacoides. Its position along the humidity/salinity gradient is the middle one, between the three associations of the order. It covers smaller surfaces, on better drained soils that the previous association. It develops on margins of abandoned salt pans; bottoms of banks, the presence of rougher granulation of soils (e.g. smaller parts of stony banks in salt pans) seem to promote the dominance of Halimione portulacoides. The vegetation is not as dense as in the previous association. This association had been previously at least partly classified as a subassociation (halimionetosum) of the macroassociation "Limonietum venetum", described by Pig-natti (1952). This macroassociation was later divided into different units by Géhu et al. (1984) due to scientific and technical (nomenclatural) arguments.
Relevé number	87	89	99	112	115	116	117	118	119	120	121	122	
Original relevé number	54	65	62	55	56	57	58	59	60	64	61	63	Fr. (%)
Diagnostic species of the association													
Cl Limonium angustifolium	1	2	+	2	1	+		3	2	3	2	2	92
Cl Artemisia coerulescens	2	1	2		3	3	3	2	2	1	2	3	92
Diagnostic species of higher syntaxonomic units													
O Halimione portulacoides	4	2	3	2	3	1	+	+	+	1	+	+	100
Cl, A Inula crithmoides	+	+	1	3	+		+	+	1	1	+	1	92
Cl, O Sarcocornia fruticosa	1	1	3	2								+	42
Others													
Aster tripolium	+		1		+	+	+	+			2		58
Elytrigia atherica	+		+				+	+		1	+		50
Suaeda maritima	1				+	+	+	1				+	50
Puccinellia palustris	2								+		1	2	42
Elytrigia elongata			+							+	+		25
Melilotus albus		+								+			17
Dactylis glomerata			+							+			17
Arthrocnemum macrostachyum									+				8
Aster lynosiris										+			8
Atriplex prostrata			+										8
Lotus corniculatus										+			8
Phragmites australis	+												8
Salicornia europaea												+	8
Tab. 1: Analytical table of the association Limonio narbonensis-Artemisietum coerulescentis Horvatic (1933) 1934 corr. Géhu et Biondi 1996.
Legend: Cl - Arthrocnemetea fruticosi Br.-Bl. et R. Tx. 1943 corr. O. Bolos 1967; O - Arthrocnemetalia fruticosi Br.-Bl. 1931 corr. O. Bolos 1967; A - Arthrocnemion fruticosi Br.-Bl. 1931 corr O. Bolos 1967.
Tab. 1: Analitična tabela asociacije Limonio narbonensis-Artemisietum coerulescentis Horvatic (1933) 1934 corr. Géhu et Biondi 1996.
Legenda: Cl - Arthrocnemetea fruticosi Br.-Bl. et R. Tx. 1943 corr. O. Bolos 1967; O - Arthrocnemetalia fruticosi Br.-Bl. 1931 corr. O. Bolos 1967; A - Arthrocnemion fruticosi Br.-Bl. 1931 corr O. Bolos 1967.
Tab. 2: Analytical table of the association Puccinelio festuciformis-Halimionetum portulacoidis Géhu, Biondi. Géhu Franck et Costa 1992.
Legend: Cl - Arthrocnemetea fruticosi Br.-Bl. et R. Tx. 1943 corr. O. Bolos 1967; O - Arthrocnemetalia fruticosi Br.-Bl. 1931 corr. O. Bolos 1967; A - Arthrocnemion fruticosi Br.-Bl. 1931 corr O. Bolos 1967.
Tab. 2: Analitična tabela asociacije Puccinelio festuciformis-Halimionetum portulacoidis Géhu, Biondi. Géhu Franck et Costa 1992.
Legenda: Cl - Arthrocnemetea fruticosi Br.-Bl. et R. Tx. 1943 corr. O. Bolos 1967; O - Arthrocnemetalia fruticosi Br.-Bl. 1931 corr. O. Bolos 1967; A - Arthrocnemion fruticosi Br.-Bl. 1931 corr O. Bolos 1967.
Relevé number	SS	9O	91	92	9S	94	95	96	97	9S	1OO	1O1	ioa	103	1O4	1O5	1O6	1OS	
Original relevé number	72	7S	75	76	SS	77	79	SO	7S	Sa	71	66	67	68	74	69	Si	7O	Fr. (%)
Diagnostic species of the association																			
O Halimione portulacoides	S	S	4	4	S	S	a	a	S	s	a	4	S	S	S	a	a	a	1OO
Puccinellia palustris	a	+	+	+	+	+	+	+		i	+	+	+	1	+	+		+	S9
Diagnostic species of higher syntaxonomic units																			
Cl, O Sarcocornia fruticosa	a	1	+	1	1	+	+	+	+		a	S	s	a	a	a	a	s	94
Cl Limonium angustifolium	+	1	1	+	+	+	+	i	+		+	1	+	+	+	i	i	+	94
Cl Artemisia coerulescens	i		+	+		+					i							+	SS
A Inula crithmoides			+			+					+							+	aa
Others																			
Aster tripolium	+					+		+	+	+	+			+		+		+	5O
Elytrigia atherica	+	+	+	+			+			i	+				+	+			5O
Suaeda maritima					+	+	+	+	a	+	+		1	+					5O
Arthrocnemum macrostachyum	i				+					i									17
'uncus maritimus									+										6
Tab. 3: Analytical table of the association Puccinellio festuciformis-Arthrocnemetum fruticosi (Br.-Bl. 1928) Géhu 1976.
Legend: Cl - Arthrocnemetea fruticosi Br.-Bl. et R. Tx. 1943 corr. O. Bolos 1967; O - Arthrocnemetalia fruticosi Br.-Bl. 1931 corr. O. Bolos 1967; A - Arthrocnemion fruticosi Br.-Bl. 1931 corr O. Bolos 1967. Tab. 3: Analitična tabela asociacije Puccinellio festuciformis-Arthrocnemetum fruticosi (Br.-Bl. 1928) Géhu 1976. Legenda: Cl - Arthrocnemetea fruticosi Br.-Bl. et R. Tx. 1943 corr. O. Bolos 1967; O - Arthrocnemetalia fruticosi Br.-Bl. 1931 corr. O. Bolos 1967; A - Arthrocnemion fruticosi Br.-Bl. 1931 corr O. Bolos 1967.
Relevé number	2S	SS	S4	S5	S6	S7	1O7	1O9	11O	111	11S	12S	124	125	126	127	12S	129	1SO	1S1	132	1SS	1S4	1S5	1S6	137	
Original relevé number	9O	91	92	9S	94	95	1O7	1O1	1OS	1O4	S9	S4	SS	S5	1O5	1O9	S6	S7	96	9S	99	1O6	97	1OO	1O2	103	Fr. (%)
Diagnostic species of the association																											
Cl, O Sarcocornia fruticosa	+	a	1	1	1	1	S	S	s	S	S	4	S	4	S	4	S	S	4	4	4	5	S	S	a	S	1OO
Puccinellia palustris	+	+							+		1	+	1	1	1	1	1		1				+				46
Diagnostic species of higher syntaxonomic units																											
Cl Limonium narbonense	4	4	4	S	S	S	1	+	+	1	4	a	a	S	S	S	S	S				+	+	+	+	1	SS
O Halimione portulacoides	+	+	+	1	1	1	a	1	1	1	1		+											+	+	+	5S
A Inula crithmoides			+	+	+						1		+	+													as
Cl Artemisia caerulescens				+	+						+																12
Others																											
Aster tripolium	+			+	+	+						+	+	+	+		1	+	+		+						46
Suaeda maritima		1		+	+	+				+			+			+						+				+	S5
Salicornia europaea									+	1					+	+					+						19
Phragmites australis		+													+			1			+	+					19
Elytrigia atherica																			+								4
Puccinellio festuciformis-Arthrocnemetum fruticosi (Br.-Bl. 1928) Gehu 1976 (Tab. 3)
This association, formerly considered as one of the most widespread subassociations ("arthrocnemetosum fruticosi") of the "Limonietum venetum", is extremely humid, for very long time inundated along the humid-
ity/salinity gradient of the three associations of the order. It has also the lower cover values. Water - predominately rainwater - stands in wintertime for longer periods, with no possibility to flow off. In the summer time it may dry up completely, especially within the abandoned salt pans of Secoveljske soline, where one of the most abundant halophyte associations can be found. It is
characterized by dominance of Sarcocornia fruticosa and weak presence/cover of Halimione portulacoides. The rest of the floristic inventory is very similar to the previous association. Outside the Secoveljske soline, patches of this association do not reach larger surfaces (a few square metres at the most). The association occurs on elevated sites in Strunjanske soline and Skocjanski zatok.
Vegetation of ¡uncus maritimus tall rush saltmarshes
The vegetation of coastal saltmarsh vegetation of the class Juncetea maritimi is characterized by the dominance of Juncus maritimus and other helophytes among halophytic species. These habitats are also supported by brackish water. The classis is further classified into the order Juncetalia maritimi and alliance Juncion maritimi, which is present on the North Adriatic coast with two suballiances: the (a) Juncenion maritimi is always characterized by dominance of Juncus maritimus and it is relatively rich with species of different syntaxa, from therophytes to scrubs, marshy halotolerant species to extreme succulents. Indicator species beside J. maritimus
are J. acutus, which is currently not present on the Slovenian seacoast, Plantago cornuti, Sonchus maritimus and Elytrigia elongata. The suballiance Puccinellienion festuciformis is characterized by the strong presence of Puccinellia palustris (=P. festuciformis) and Aster tri-polium. These stands are wet and frequently inundated.
Association Juncetum maritimi-acuti Horvatic 1934 (suballiance Juncenion maritimi) (Tab. 4)
Juncus maritimus is absolutely dominant perennial in this association, which also forms main physiognomic aspect of these habitats, being in the permanent contact with sea or brackish water. One group of the relevés is more hygrophilous and characterized by Limonium an-gustifolium and Pucinellia palustris. It covers smaller surfaces at the shores of the channels or lagoons, sometimes at closed habitats in depressions with accumulating water. Another group of relevés is characterised by the natural saltmarsh at Sv. Katarina (Ankaran), where the conditions are drier and more species occur, including the rare ones, like Linum maritimum, Cen-taurium spicatum and Elytrigia elongata.
Releve number	5	6	7	8	9	10	11	12	13	14	38	39	40	41	42	43	114	138	139	140	
Original releve number	123	136	129	130	131	138	133	139	140	132	121	122	125	126	127	124	128	134	137	135	Fr. (%)
Diagnostic species of the association																					
Cl, O -uncus maritimus |3|3|3|4|4|4|3|3|4|5|4|3|3|2|2|2|1|4|3|4|100																					
Diagnostic species of higher syntaxonomic units																					
Cl Limonium angu stifolium	2		+	+	+	+		1	1		2	3	2	5	4	3	4		+	+	80
Cl, O Puccinellia palustris	2	2	+	+				+			+	+		+	+	2	+				55
Cl, O Aster tripolium		1			+	+	+		+	+				+	1			+		2	50
O Carex extensa							+						+				+	1	+		25
A Plantago cornuti															+			3	3	3	20
A Sonchus maritimus																			+	+	10
A Elytrigia elongata						+															5
Others																					
Phragmites australis		+			+		+	+	+	2				+		2		1	+	2	55
Suaeda maritima			+				+		+	+	+		1			1					35
Inula crithmoides											+	+	+			+	2			1	30
Halimione portulacoides		+				+						+	+	+			1				30
Artemisia caerulescens							+										+	+	+	1	25
Elytrigia atherica												+						+	2	2	20
Salicornia europaea			+							+			+		+						20
Sarcocornia fruticosa	+						+										+				15
Bolboschoenus maritimus							+	+											+		15
Centaureum spicatum						+														+	10
Linum maritimum																			+	2	10
Holoschoenus maritimus																			+		5
Dittrichia viscosa																			+		5
Tab. 4: Analytical table of the association Juncetum maritimi-acuti Horvatic 1934.
Legend: Cl - Juncetea maritimi Br.-Bl. 1952 em. Beeftink 1965; O - Juncetalia maritimi Br.-Bl. 1931; A - Juncenion maritimi Gehu et Biondi 1995.
Tab. 4: Analitična tabela asociacije Juncetum maritimi-acuti Horvatic 1934
Legenda: Cl - Juncetea maritimi Br.-Bl. 1952 em. Beeftink 1965; O - Juncetalia maritimi Br.-Bl. 1931; A - Juncenion maritimi Gehu et Biondi 1995.
Tab. 5: Analytical table of the association Limonio narbonensis-Puccinellietum palustris (Pignatti 1966) Gehu et Scopp. 1984 in Gehu et al. 1984.
Legend: Cl - Juncetea maritimi Br.-Bl. 1952 em. Beeftink 1965; O - Juncetalia maritimi Br.-Bl. 1931; subA - Pucci-nellienion festuciformis (Gehu et Scopp. 1984 in Gehu, Scoppola, Caniglia, Marchiori et Gehu Franck 1984) Gehu et Biondi 1995.
Tab. 5: Analitična tabela asociacije Limonio narbonensis-Puccinellietum palustris (Pignatti 1966) Gehu et Scopp. 1984 in Gehu et al. 1984.
Legenda: Cl - Juncetea maritimi Br.-Bl. 1952 em. Beeftink 1965; O - Juncetalia maritimi Br.-Bl. 1931; subA - Pucci-nellienion festuciformis (Gehu et Scopp. 1984 in Gehu, Scoppola, Caniglia, Marchiori et Gehu Franck 1984) Gehu et Biondi 1995.
ReIevé number	22	23	24	25	26	27	29	30	3i	32	44	
OriginaI reIevé number	113	111	ii2	110	118	i20	ii7	ii9	ii5	ii6	ii4	Fr. (%)
Diagnostic species of the association												
CI Limoniumangustifolium	+	+	+	+	2	2	5	4	4	2	2	i00
CI, O, subA Puccinellia palustris	3	5	4	3	4	3	+	i	i	i		9i
Diagnostic species of higher syntaxonomic units												
CI, O Aster tripolium	+			+	+	+	+	+			4	64
CI, O Juncus maritimus					i	+	+		+	+		45
O Carex extensa											+	9
Others												
Phragmites australis					+	+		+			+	36
Suaeda maritima				i					i	+	+	36
Salicornia europaea	+						+			+		27
Halimione portulacoides			i									9
Spartina maritima					+							9
Association Limonio narbonensis-Puccinellietum palustris (Pignatti 1966) Géhu et Scopp. 1984 in Géhu et al. 1984 (suballiance Puccinellienion festuciformis) (Tab.
5)
This association is characterized by combined domination of Puccinellia palustris and Limonium angustifo-lium. It appears in belts along the channel banks, lagoon margins, with high level of salt or brackish water. These stands are exposed to tidal oscillations and strong nutrient flow. -uncus maritimus is present, but not in all relevés. This association is in contact with syntaxa of the class Spartinetea on one side and syntaxa of the class Arthrocnemetea on the other side. The morphology of the dominant Limonium angustifolium species is different from drier sites, inhabited by Arthrocnemetea vegetation: here Limonium is taller and bigger. This species reaches its ecological optimum in two quite different habitats, classified in two different phytosociological classes.
CONCLUSIONS
The elaborated associations are presented in synoptic table (Tab. 6), where some Salsola soda-dominated stands are included and classified within the class Cak-iletea maritime, provisionally treated in the rank of association Salsoletum sodae, due to the presence of only two Cakiletea characteristic species - Salsola soda and
Atriplex prostrata. This anthropogenic stands with very small surfaces on the stony seashore or at the tops of the muddy dikes within some salt basins are very poor in species composition and therefore very hard to classify. This classification in the synoptic table is provisional only and it will not be further discussed in this treatise.
We could summarize that the two classes Juncetea maritimae and Arthrocnemetea fruticosae indeed differ ecologically, but further delimitation of the lower syntaxa (associations, namely) is based mostly on frequency and coverage of the species, e.g. Arthrocnemum vs. Halimione vs. Artemisia within the Arthrocnemetea clas-sis, and Limonium/Puccinellia vs. Juncus and typical Juncetea inventory. Characteristic species for the two classes of halophilous vegetation are distributed more or less across all five (six) associations. It is very difficult to draw conclusions only on the basis of species presence and cover. The "understanding" of vegetation assemblage is beyond the species combinations and numerical classifications: it should be accompanied with ecological data (salinity, water potential, soil properties, nutrients etc) and functional plant traits (Kaligaric & Skornik, 2006). For a better understanding of the classification of respective vegetation types, Poldini et al. (1999) incorporated structural data (life form and growth form of plants) into his treatise. For causal understanding, where pure classification is only a starting point, an eco-physiological and morphological approach is needed.
Tab. 6: Synoptic table of the associations of the classes Arthrocnemetea fruticosi, Juncetea maritimi and Cakiletea maritimae. Values in the table correspond to the relative frequencies (in percentage) of the species in presented group of relevés.
Legend: Li-Ar: Limonio narbonensis-Artemisietum coerulescentis; Pu-Ha: Puccinellio festuciformis-Halimionetum portulacoidis; Pu-Ar: Puccinellio festuciformis-Arthrocnemetum fruticosi; Ju ma: Juncetum maritimi-acuti; Li-Pu: Limonio narbonensis-Puccinellietum palustris; Sals: Salsoletum sodae; Di1-5: diagnostic species of ass.; Cl1: Arthrocnemetea fruticosi; Cl2: Juncetea maritimi; 01: Arthrocnemetalia fruticosi; 02: Juncetalia maritimi; subA2: Puc-cinellienion festuciformis.
Tab. 6: Sinoptična tabela asociacij razredov Arthrocnemetea fruticosi, Juncetea maritimi in Cakiletea maritimae. Vrednosti v tabeli ustrezajo frekvencam (v odstotkih) pojavljanja vrst v predstavljenih skupinah popisov. Legenda: Li-Ar: Limonio narbonensis-Artemisietum coerulescentis; Pu-Ha: Puccinellio festuciformis-Halimionetum portulacoidis; Pu-Ar: Puccinellio festuciformis-Arthrocnemetum fruticosi; Ju ma: Juncetum maritimi-acuti; Li-Pu: Limonio narbonensis-Puccinellietum palustris; Sals: Salsoletum sodae; Di1-5: diagnostične vrste asociacij; Cl1: Arthrocnemetea fruticosi; Cl2: Juncetea maritimi; 01: Arthrocnemetalia fruticosi; 02: Juncetalia maritimi; subA2: Puc-cinellienion festuciformis.
	Association	Li-Ar	Pu-Ha	Pu-Ar	Ju ma	Li-Pu	Sals
	Number of relevés	12	18	26	2O	11	4
Diagnostic species of the associations							
Di1,Di5(Cl1, CI2)	Limonium angustifolium	92	94	88	8O	1OO	
Dil, (Cll)	Artemisia caerulescens	92	SS	12	25		
Di2(O1)	Halimione portulacoides	1OO	1OO	58	SO	9	
Di2,Di3,Di5(Cl2, O2, subA2)	Puccinellia palustris		89	46	55	91	
Di3(Cl1, Ol)	Sarcocornia fruticosa	42	94	1OO	15		
Di4(Cl2, O2)	Juncus maritimus		6		1OO	45	
Cll Arthrocnemetea fruticosi Br.-Bl. et R. Tx. 1943 corr O. Bolos 1967							
Inula crithmoides 92 22 2S SO . .							
CI2 Juncetea maritimi Br.-Bl. 1952 em. Beeftink 1965							
	Aster tripolium	58	5O	46	5O	64	
	Carex extensa				25	9	
	Plantago cornuti				2O		
	Sonchus maritimus				1O		
	Elytrigia elongata	25			5		
CI3 Cakiletea maritimae R. Tx. et Prsg. 195O							
	Salsola soda						1OO
	Atriplex prostrata						
Thero-Salicornietea Pignatti ex Tx. In Tx. Et Oberdorfer 1958 corr. Tx.1974							
	Salicornia europaea	8		19	2O	27	
	Suaeda maritima	5O	5O	S5	S6		
Spartinetea maritimae (R. Tx. 1961) Beeft., Géhu, Ohba et R. Tx. 1971							
Spartina maritima .... 9 .							
Others							
	Puccinellia palustris	42					
	Melilotus albus	17					
	Dactylis glomerata	17					
	Aster lynosiris	8					
	Atriplex latifolia	8					
	Lotus corniculatus	8					
	Arthrocnemum macrostachyum	8	17				
	Elytrigia atherica	5O	5O	4	2O		66
	Phragmites australis	8		19	55	S6	
	Bolboschoenus maritimus				15		
	Centaurium spicatum				1O		
	Linum maritimum				1O		
	Holoschoenus maritimus				5		
	Dittrichia viscosa				5		
ACKNOWLEDGMENTS
The research for this paper was funded by the program group "Biodiversity" (P1-0078), the research project "Biodiversity gradient in relation to biogeo-graphic, altitudinal and ecological factors" (J1-6577) and research project "Mapping of Secoveljske soline and cultivation of autochtonous grassworts" (L1-7001-006), the first two headed by B. Krystufek, the last one by J. Necimer and M. Regvar, all granted from the Slovenian Research Agency.
APPENDIX 1
Synoptic table of studied vegetation types:
Arthrocnemetea fruticosi Br.-Bl. et R. Tx. 1943 corr. O. Bolos 1967 (=Sarcocornietea fruticosae Br.-Bl. et R. Tx. 1943 corr. Castroviejo et Cirujano 1980)
Arthrocnemetalia fruticosi Br.-Bl. 1931 corr. O. Bolos 1967 (=Sarcocornietea fruticosi Br.-Bl. 1931 corr. Castroviejo et Cirujano 1980) Artrhrocnemion fruticosi Br.-Bl. 1931 corr. O. Boloo 1967
•	Limonio narbonensis-Artemisietum coerulescentis Horvatic (1933) 1934 corr. Géhu et Biondi 1996
•	Puccinellio festuciformis-Halimionetum portula-coidis Géhu, Biondi, Géhu Franck et Costa 1992
•	Puccinellio festuciformis-Arthrocnemetum fruticosi (Br.-Bl. 1928) Géhu 1976
Juncetea maritimi Br.-Bl. 1952 em. Beeftink 1965 juncetalia maritimi Br.-Bl. 1931 juncion maritimi Br.-Bl. 1931
Puccinellenion festuciformis (Géhu et Scopp. 1984 in Géhu, Scoppola, Caniglia, Marchiori et Géhu Franck 1984) Géhu et Biondi 1995
•	Limonio narbonensis-Puccinellietum palustris (Pignatti 1966) Géhu et Scopp. 1984 in Géhu et al. 1984
•	juncenion maritimi Géhu et Biondi 1995
•	juncetum maritimi-acuti Horvatic 1934
Cakiletea maritimae R. Tx. et Prsg. 1950 Euphorbietalia peplis R. Tx. 1950 Thero-Suaedion splendentis Br.-Bl. 1931 • Salsoletum sodae Pignatti 1953
APPENDIX 2
Localities of relevés:
Table 1: 87, 89: Sečovlje - Fontanigge, MTB: 0547/2; 99: Ankaran, MTB: 0448/1; 112, 115, 116-119: Sečovlje - Fontanigge, MTB: 0547/2; 120: Ankaran, MTB: 0448/1; 121, 122: Sečovlje - Fontanigge, MTB: 0547/2. Table 2: 88, 90: Sečovlje - Fontanigge, MTB: 0547/2; 91: Koper-Škocjanski zatok, MTB: 0448/3 and 0448/4; 92-97: Strunjan, MTB: 0447/4; 98, 100, 101: Sečovlje -Fontanigge, MTB: 0547/2; 102, 103: Strunjan, MTB: 0447/4; 104-106: Sečovlje - Fontanigge, MTB: 0547/2; 108: Koper - Škocjanski zatok, MTB: 0448/3 and 0448/4.
Table 3: 28, 33-36: Sečovlje - Fontanigge, MTB: 0547/2; 37: Strunjan, MTB: 0447/4; 107, 109: Sečovlje -Fontanigge, MTB: 0547/2; 110: Strunjan, MTB: 0447/4; 111, 113, 123-127: Sečovlje - Fontanigge, MTB: 0547/2; 128: Koper - Škocjanski zatok, MTB: 0448/3 and 0448/4; 129, 130-133: Koper - Škocjanski zatok, MTB: 0448/3 and 0448/4; 134: Strunjan, MTB: 0447/4; 135: Sečovlje - Fontanigge, MTB: 0547/2; 136: Strunjan, MTB: 0447/4; 137: Sečovlje - Fontanigge, MTB: 0547/2. Table 4: 5, 6, 7: Sečovlje - Fontanigge, MTB: 0547/2; 8: Strunjan, MTB: 0447/4; 9: Sečovlje - Fontanigge, MTB: 0547/2; 10, 11: Ankaran, MTB: 0448/1; 12: Strunjan, MTB: 0447/4; 13: Sečovlje - Fontanigge, MTB: 0547/2; 14: Koper - Škocjanski zatok, MTB: 0448/3 and 0448/4; 38: Sečovlje - Fontanigge, MTB: 0547/2; 39: Koper -Škocjanski zatok, MTB: 0448/3 and 0448/4; 40, 41: Sečovlje - Fontanigge, MTB: 0547/2; 42, 43: Koper -Škocjanski zatok, MTB: 0448/3 and 0448/4; 114, 138: Sečovlje - Fontanigge, MTB: 0547/2; 139, 140: Ankaran, MTB: 0448/1.
Table 5: 22-27, 29, 30-32, 44: Sečovlje - Fontanigge, MTB: 0547/2.
VEGETACIJA OBMORSKIH MOČVIRIJ Z OBMORSKIM LOČKOM (JUNCETEA MARITIMAE) IN HALOFITNIH TRAJNIC (ARTHROCNEMETEA FRUTICOSAE) NA SLOVENSKI OBALI
Mitja KALIGARIC
Univerza v Mariboru, Fakulteta za naravoslovje in matematiko, Oddelek za biologijo, SI-2000 Maribor, Koroška 160
in
Univerza na Primorskem, Znanstveno-raziskovalno središče Koper, Inštitut za biodiverzitetne študije, SI-6000 Koper, Garibaldijeva 18
E-mail: mitja.kaligaric@uni-mb.si
Sonja ŠKORNIK
Univerza v Mariboru, Fakulteta za naravoslovje in matematiko, Oddelek za biologijo, SI-2000 Maribor, Koroška 160
POVZETEK
Na kratki slovenski morski obali prevladujejo flišni klifi, kjer je halofitna vegetacija zelo slabo razvita. Zelo raznolika pa je halofitna vegetacija na muljastih sedimentih, tako naravnih kot antropogenih ali tistih v opuščanju (soline). Med leti 1984-87 in 1998-1999 je bilo zbranih 140 popisov halofitne vegetacije po standardni Braun-Blanquetovi metodi. V pričujočem članku smo podrobneje analizirali in predstavili tiste popise, ki zajemajo vegetacijo razredov halofitnih trajnic- grmičkov in polgrmičkov - Arthrocnemetea fruticosae in obmorskega ločkovja -Juncetea maritimi-acuti. Ob pomoči značilnih in dominantnih vrst v popisih smo identificirali 5 združb, in sicer v okviru prvega razreda tri združbe, ki sledijo gradientu vlažnosti/slanosti od najbolj suhih habitatov do najbolj zalitih v tem vrstnem redu: Limonio narbonensis-Artemisietum coerulescentis, Puccinellio festuciformis-Halimionetum portulacoidis in Puccinellio festuciformis-Arthrocnemetum fruticosi. Prva asociacija je najbolj sušna, dvignjena od vod, pojavlja se manj ekstremnih halofitov, značilna je vrsta Artemisia caerulescens. Sestoji so precej sklenjeni in vrstno pestri. V drugi je dominantna vrsta Halimione portulacoides, vegetacija je manj sklenjena, vrstno revnejša in zaseda manjše površine. V tretji asociaciji absolutno prevladuje vrsta Sarcocornia fruticosa, ki je najbolj namočena in ima najmanj sklenjeno vegetacijo. Za morska močvirja z vrsto Juncus maritimus (J. acutus manjka!) sta značilni dve asociaciji. Sestoji asociacije Limonio narbonensis-Puccinellietum palustris so razviti v pasovih ob obalah kanalov, lagun, v slani ali brakični vodi, so precej namočeni oziroma izpostavljeni plimovanju in dotoku hranil. Asociacijo označuje dominantnost vrst Limonium angustifolium ter Puccinellia palustris in je vrstno razmeroma revna. Asociacija Juncetum maritimi-acuti je vrstno pestrejša, označuje pa jo dominantnost obmorskega ločka. Nekaj popisov s prevladujočo vrsto Salsola soda je bilo klasificiranih kot Salsoletum sodae, ki sodi v razred Cakiletea maritimae. Sintezna tabela pokaže razlikovanje asociacij glede na dominantne vrste ter obstoj značilnic 2 (3) razredov halofitne vegetacije, kar je za uvrščanje v višje sintaksone v malovrstnih halofitnih sestojih vedno problematično. Zato se upoštevajo tudi nekatere strukturne značilnosti rastlin (življenjska oblika in oblika rasti), za globlje razumevanje pa je treba poznati tudi okoljske parametre in funkcionalne značilnosti rastlin.
Ključne besede: fitosociologija, halofitna vegetacija, klasifikacija, severni Jadran
REFERENCES
Beguinot, A. (1941): La vita delle piante vascolari. La laguna di Venezia. Ferrari, Venezia, 3, p. 5, t. 9, 1-369. Braun-Blanquet, J. (1964): Pflanzensoziologie. Grundzüge der Vegetationskunde. 3. Auflage. Springer Verlag, Berlin, Wien, New York, 865 p.
Council Directive 92/43/EEC of 21 May 1992 on the
conservation of natural habitats and of wild fauna and flora. Official Journal L 206, 22/07/1992 P. 0007 -0050.
Darovec, D. (1992): Pregled zgodovine Istre. Knjižnica Annales. 1. Ponatis. Zgodovinsko društvo za južno Primorsko, Koper, 88 str.
Dierschke, H. (1994): Pflanzensoziologie: Grundlage und Methode. Ulmer, Stuttgart, 683 p. Ferrari, C., R. Gerdol & F. Piccoli (1985): The halophi-lous vegetation of the Po Delta (northern Italy ). Vegeta-tio, 61, 5-14.
Fornaciari, G. (1968): Aspetti floristici e fitosociologici della Laguna di Grado e del suo litorale. Atti Accad. Sci. Lett. Arti Udine, 6, 5-199.
Frondoni, R. & M. Iberite (2002): The halophile vegetation of the sedimentary coast of Lazio (central Tyrrhenian District, Italy). Plant Biosystems, 136, 49-67. Géhu, J. M. (1999): Schéma synsystématique des principales classes de végétations littorales sédimentaries européennes avec références à d'autres territories holoarc-tiques. Ann. Bot., 56, 5-52.
Géhu, J. M. & E. Biondi (1995): Essai de typologie phy-tosociologique des habitats et des végétations halophiles des littoraux sedimentaries perimediterraneens halophiles des littoraux sedimentaries périméditerranéens et thermo-atlantiques. Fitosociologia, 30, 201-212. Géhu, J. M. & E. Biondi (1996): Synoptique des associations végétales du littoral adriatique italien. Giorn. Bot. Ital., 130, 257-270.
Géhu, J. M., M. Costa, A. Scoppola, E. Biondi, S. Mar-chiori, J. B. Peris, J. Franck, G. Canglia & L. Veri (1984):
Essai synsystématique et synchorologique sur les végétations littorales italiennes dans un but conservatiore. Doc. Phytosoc. N.S., 8, 393-474.
Iberite, M. & R. Frondoni (1997): Contributo alla cono-scenza della vegetazione del Parco Nazionale del Ci r-ceo: le comunità a salicornie annuali. Fitosociologia, 34, 121-125.
Kaligaric, M. (1996): Threat status of halophyte flora and vegetation. In: Gregori, J. (ed.): Narava Slovenije, stanje in perspective. Zbornik prispevkov o naravni dediščini Slovenije. Društvo ekologov Slovenije, Ljubljana, 113121 str.
Kaligaric, M. (1997): Rastlinstvo Škocjanskega zatoka -deset let kasneje. Falco, 11, 11-12. Kaligaric, M. (1998): Botanični pogled na možne ureditve naravnega rezervata Škocjanski zatok. Annales, Ser. Hist. Nat., 8, 131-142.
Kaligaric, M. & M. Tratnik (1981): Ohranimo Sečovel-jske soline. Proteus, 44, 122-126.
Kaligarič, M. & T. Wraber (1988): Obmorski lan in klasnata tavžentroža v Sloveniji nista izumrla. Proteus, 50,372-373.
Kaligarič, M. & S. Skornik (2006): Halophile vegetation of the Slovenian seacoast: Thero-Salicornietea and Spar-tinetea maritimae. Hacquetia, 5, 25-36. Martinčič, A., T. Wraber, N. Jogan, V. Ravnik, A. Podobnik, B. Turk & B. Vreš (1999): Mala flora Slovenije. Ključ za določanje praprotnic in semenk. 3. izdaja. Tehniška založba Slovenije, Ljubljana. Mucina, L. (1997): Conspectus of classes of European vegetation. Folia Geobot. Phytotax., 32, 117-172. Pignatti, S. (1952): Introduzione allo studio fitosociolo-gico della Pianura Veneta orientale con particolare ri-guardo alla vegetazione litoranea. Arch. Bot., 28, 265329.
Pignatti, S. (1966): La vegetazione alofila della laguna veneta. Mem. 1st. Veneto Sci. Lett. Arti, Cl. Sci. Mat. Nat., 33, 3-147.
Podani, J. (2001): SYN-TAX 2000. Computer Programs for data analysis in ecology and systematics. Budapest. Poldini, L., M. Vidali & M. L. Fabiani (1999): La vegetazione del litorale sedimentario del Friuli-Venezia Giulia (NE Italia) con riferimenti alla regione alto-adriatico. Studia Geobot., 17, 3-68.
Rivas-Martinez, S. (1990): Sintaxonomia de la clase Thero-Salicornietea en Europa occidental. Ecol. Medit., 16,359-364.
Sajna, N. & M. Kaligarič (2005): Vegetation of the Stjuža coastal lagoon in Strunjan landscape park (Slovenia): a draft history, mapping and conservation evaluation. Annales, Ser. Hist. Nat., 15(1), 79-90. Westhoff, V. & E. van der Maarel (1973): The Braun-Blanquet approach. In: Whittaker, R. H. (ed.): Ordination and classification of communities. 2nd Edition. Dr. W. Junk Publishers, The Hague, p. 617-726.