Plant Ecology 132: 155170, 1997. c 1997 Kluwer Academic Publishers. Printed in Belgium.

155

The halophilous vegetation of the Orumieh lake salt marshes, NW. Iran
Y. Asri1 & M. Ghorbanli2
Research Institute of Forests and Rangelands, P.O.Box 13185-116, Tehran-Iran and 2 Biology Dept., Tarbiat Moalem University, Mofatteh Ave., Tehran-Iran
Received 12 September 1995; accepted in revised form 28 April 1997
1

Key words: Halophytes, Numerical analysis, Ordination, Phytosociology, Saline soils, Salt marsh vegetation

Abstract The halophilous vegetation of the Orumieh lake salt marshes has been studied, using the Braun-Blanquet method. Vegetation types have been dened by physiognomic-oristic system. The following six main groups of communities are recognized: (1) Semi-woody shrub and perennial halophytic communities (Class Halocnemetea strobilacei) including 6 associations and 5 subassociations, (2) Annual halophytic communities (Class Thero-Salicornietea) including 5 associations and 1 subassociation, (3) Salt marsh brushwood communities (Class Tamaricetea) including 4 associations, (4) Rush and herbaceous perennial halophytic communities (Class Juncetea maritimi) including 7 associations and 1 subassociation, (5) Rush and herbaceous perennial halotolerant communities (Class Agrostietea stoloniferae) including 5 associations, (6) Hydrophilous halotolerant communities (Class Phragmitetea) including 3 associations and 2 subassociations. The soil of these communities has been analysed and their habitats are described and discussed. Abbreviations: AFC Analyse Factorielle des Correspondances; CAH Classication Ascendant Hierarchique; PCA Principal Components Analysis. Introduction The coastal salt marshes comprise areas of land bordering the seas and lakes, more or less covered with vegetation and subject to periodic inundation by tide. They have certain qualities, which is related to the proximity to the sea and lake, that distinguishes them from inland salt marshes (Chapman 1977). Littoral salt marshes are essentially fringes of inland deserts, their landward boundary being dened by desert conditions. Ecological factors, such as terrain or climate, can be used to delimit the littoral marshes. When there is a narrow belt along the coast surrounded by a steep barrier of mountains (e.g. part of the studied area), the limits are clear. But in a broad plain that stretches inland from the coast, there may be no distinct physiographic barrier. Therefore, other habitat features including vegetation type have to be used. Vegetation characteristics, related to physiographic attributes reecting both climatic and edaphic factors, provide the best single basis for delimiting littoral salt marshes. These salt marshes may be only a narrow belt within the reach of salt spray. They can be a hundred metres wide or they may extend inland for many kilometers (Zahran & Willis 1992). Iran is the classical country of the great salines and kavirs (Zohary 1973). Halophytic communities of Iran are still among the most poorly known vegetation units. The distribution of halophytic communities has been depicted cartographically by Mobayen & Tregubov (1970), Mobayen (1976, Kavir-e Lut), Freitag (1977, Turan Biosphere Reserve), Kramer (1984, Gulf Region near Bandar-e Rig), Carle & Frey (1977), Frey (1982, Maharlu basin) and Frey et al. (1985, northern peripheral region of Dasht-e kavir). Further physiognomic and ecologic-geographic data on such communities have been given by Kunkel (1977, Persian Gulf area), Ghorbanli & Lambinon (1978, Qom lake), Frey & Probst (1986), Breckle (1982, Turan Biosphere Reserve), Breckle (1983), Assadi (1984, Kavir regions), Akhani (1989, Kavire-

156 logical stations, namely; Miandoab, Maragheh, Tabriz, Sharaf Khaneh, Orumieh and Barandoz-Chay, show that the total annual precipitation is around 324 mm, with the maximum occuring in the winter and spring months. Mean maximum and minimum temperatures are 36.4 C for July and ,13:3 C for January. Methods Sample releves were performed according to the Braun-Blanquet method (Mueller-Dombois & Ellenberg 1974; Westhoff & Van der Maarel 1978). The releve size was determined by establishing a speciesarea curve in each vegetation type. The releves of herbaceous perennial and hydrophilous halotolerant communities were recorded on an area of 0.244 m2 , those of the halophilous communities on 0.2516 m2 . The releve and species groups were dened by the AFC method, using the computer program of Anaphyto (Briane 1991). The most popular ordination technique has been PCA, although the strategy of AFC has become an effective tool for phytosociological clustering and table sorting. The AFC method developed by Benzecri (1969), allows all the points (species and releves) to be represented on the same diagram. The species with the most similar patterns form groups, which are placed in, or near, groups of releves with similar species composition. The ordination is a oristic one, and the interpretation of a oristic axis in terms of environmental factors can only be tentative. Important environmental factors may be reected on two or more oristic axes. The releve and species clusters have been dened by numerical analysis of AFC data, using a CAH technique included in the program. CAH is essentially a clustering procedure based on releve similarity, combined with a procedure for obtaining a diagonal structure of clusters in the table. In the phytosociological table, species and releves are ordered in such a way that the species with a more or less similar distribution pattern over the releves are grouped together and, similarly, releves with a more or less similar species content are placed nex to one another. The names of syntaxa correspond with the codes of phytosociological nomenclature (Barkman et al. 1976). The nex step comprises the replacement of each association data by a column in which for each participating species the presence degree is indicated. Such a table is called synoptic table. After comparison of the synoptic table with those from other types of vegeta-

Figure 1. Location map of the study area.

Meyghan) and, Akhani & Ghorbanli (1993). There have been relatively few investigations using the phytosociological approach on salt marshes of Iran, including oristic survey of salt desert vegetation by Zohary (1963, 1973) and Leonard (1991), littoral salt marsh vegetation by Asri et al. (1995) and Atri et al. (1995).

The study area The study area is salt marshes of Orumieh lake between western and eastern Azarbaijan provinces (Figure 1). The total area is ca. 2000 km2 ; most of which is located in the eastern part of the lake. The average altitude is about 1284 m. The geological substratum mainly consists of Alluvium, Coastal plains and Swamps corresponding to the Quaternary and recent times depostis. Other formations are belong to the Oligo-Miocene and Cretaceous periods (Iran Oil Co. 1960). A remarkable part of the soils belongs to Saline soils series. They are chiey Solonchaks, which fall into the Aridisols category according to the US comprehensive system of soil classication (Dregne 1976). Also, there are LowHumic Gley and Alluvial soils groups, that belong to the Inceptisols and Entisols categories, respectively. The area belongs to the cold-semiarid climatic zone of Embergers system. According to the Gaussans system, eastern and western parts of the lake belong to hot-arid Mediterranean zone and cold steppe zone, respectively. Means of over 25 years from six meteoro-

157 tion from the same region an idea can be formed about the local diagnostic species groups in the table under study. The syntaxonomical research step starts when a vegetation type is to be tted into the hierarchic syntaxon tables. A few samples of the upper layer of the soil were taken from each community. Analysis of soil samples has been carried out following Richards (1954) and Jackson (1960). ical scheme (Table 2) is not in accordance with the scheme of European salt marsh vegetation, proposed by Chapman (1974). According to Chapman (1974) this vegetation type should be assigned in the class Halostachyetea. The synoptic table (Table 1) clearly shows that the Halocnemetum strobilacei association includes the following four subassociations of perennial and annual halophytic plants: Halopeplidetosum pygmaeae, Phragmitetosum stenophyllae, Climacopteretosum crassae and Frankenietosum pulverulentae. These subassociations are distributed along a gradient of decreasing salt in the soil. Also, the synoptic table (Table 1) shows that Kalidietum caspici includes the Psylliostachyetosum leptostachyae subassociation. The syntaxonomical scheme of this class is shown in Table 2. Class Thero-Salicornietea The communities characterized by annual halophytes settled on soils subject to natural or articial disturbances have been assigned to the class TheroSalicornietea. This is in accordance with the classication of European salt marsh vegetation, proposed by Chapman (1974), Ellenberg (1986) and Biondi (1989). The associations belonging to this class are the following (Table 1): Salicornietum europaeae, Suaedetum maritimae, Salsoletum sodae, Petrosimonietum brachiatae and Petrosimonietum glaucae. Our results suggest the syntaxonomical scheme for this class (Table 2). Class Tamaricetea Zohary (1973) described the class Tamaricetea salina for salt marsh brushwood communities of Iran. But he did not give a hierarchical classication for it. The associations belonging to the class Tamaricetea are the following (Table 1): Tamaricetum meyeri, Tamaricetum octandrae, Tamaricetum kotschyi and Tamariceto meyeri-octandrae. In all the associations a substratum of Aeluropus littoralis occurs (Table 1). The syntaxonomical scheme of this class is shown in Table 2. Class Juncetea maritimi Similar the others, e.g. There are vast stands of herbaceous perennial halophytic communities in most parts of Orumieh lake salt marshes. They are accompanied by mosaics or rush plants. This vegetation type according to the syntaxonomical scheme (Table 2)

Results The program used produces an ordination with ve axes. The distribution of species or releve groups is better revealed on axes 13 than others (Figures 2 and 3). The oristic gradient indicated in Figure 3 can be interpreted ecologically as an overall salinity gradient ranging from low to high saline environments. Also, this gradient related to the structural complexity of the vegetation from pioneer to brushwood communities. The other oristic gradient (axes 12) can be interpreted as related to moisture ranging from hydrophilous halotolerant communities to perennial halophytic communities. Generally, six groups can be recognized on axes 13 (Figures 2 and 3). Similarly, six species and releve clusters have been distinguished in the dendrograms obtained with CAH method (e.g. Figure 4). Partial analysis of groups resulted by AFC method is shown that each group may be divided to some subgroups. Finally, 39 subgroups are distinguished on the basis of partial analysis. The synoptic table (Table 1) is then constructed, using the releve and species clusters obtained by CAH method followed by partial analysis. According to the results of the numerical analysis, all the associations distinguished are to be included in the following classes: Class Halocnemetea strobilacei According to the results of the numerical analysis, most of the communities are characterized by semiwoody shrub and perennial halophytes on muddy and dry salty ats should be included in the class Halocnemetea strobilacei. Our results do not conrm the proposed classication of Zohary (1973). According to Zohary (1973) most of the halophytic communities of Iran should be referred to the class Halocnemetea strobilacei irano-anatolica. It seems, this class should be split into several classes. Also, our syntaxonom-

158

Table 1. Synoptic table of 30 associations and 9 subassociations in the Orumieh lake salt marshes. 1. Salicornietum europaeae, 2. Suaedetum maritimae, 3. Salsoletum sodae, 4. Petrosimonetum brachiatae, 5. Petrosimonietum glaucae, 6. Sclerochloetosum durae, 7. Halocnemetum strobilacei, 8. Climacopteretosum crassae, 9. Phragmitetosum stenophyllae, 10. Halopeplidetosum pygmaeae, 11. Frankenietosum pulverulentae, 12. Halostachyetum caspicae, 13. Kalidietum caspici, 14. Psylliostachyetum leptostachyae, 15. Halimionetum verruciferae, 16. Limonietum carnosi, 17. Limonietum meyeri, 18. Tamaricetum kotschyi, 19. Tamaricetum meyeri, 20. Tamariceto meyeri-octandrae, 21. Tamaricetum octandrae 1 4 3 4 10 3 3 3 6 2 4 2 4 5 21 3 3 3 4 3 4 3 7 3 3 3 4 2 6 9 17 3 10 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 3 4 18 3 4 19 4 11 20 4 12 21 4 7

Association and subassociation no. No. of releves No. of species

100(35) 25(1) 50( 25( 25(1) 33( 100(34) 100(2) 100(2) 100(3) 100(2) 66(12) 100(34) 100(3) 100(2) 20( 33( 100(24) 33( 100(23) 50(

33(

20( 20( 33(

66(

+) +)
50( 33(2)

50(1)

+) 25(+) +) +)
100(34) 20(1) 20(1) 66(1) 50(

+) +) +) +) +) 40(+,1) 40(+) +) +)
50(1)

+) +)

44(12)

66(2)

100(33) 100(3-4) 100(24) 33(1) 44( 33(1) 66(1)

40(1)

100(3) 100(23) 100(34) 100(12) 33(2) 100(34) 100(2) 75(

+,1) 44(+) 33(+) 11(+) 50(+) 55(+)

+)

100(23) 100(34) 50(2) 75( 50(

+,)

Character-taxa of the associations Salicornia europaea Suaeda maritima Salsola soda Petrosimonia brachiata Petrosimonia glauca Halocnemum strobilaceum Suaeda acuminata Eremopyrum triticum Halostachys caspica Kalidium caspicum Halimione verrucifera Scorzonera laciniata Lepidium cartilagineum ssp. pumilum Artemisia fragrans Camphorosma monspeliaca Limonium carnosum Limonium meyeri Tamarix kotschyi Cressa cretica Tamarix tetragyna var. meyeri Spergularia media Tamarix octandra Tamarix ramosissima Cyperus fuscus Crypsis schoenoides Cyperus laevigatus var. distachyos

75(12)

+)

Table 1. (continued) 1 4 4 3 10 3 3 3 6 2 4 2 4 5 21 3 3 3 4 3 4 3 7 3 3 3 4 2 6 9 17 3 10 3 4 3 4 4 11 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 4 12 21 4 7

Association and subassociation no. No. of releves No. of species

100(3) 20( 100(23) 100(2) 100(45) 33(1) 100(3) 11( 33( 11(1) 100(23) 33(

20(

33(

+) +)
50(1)

25(2)

66(

+,1) +)
50( 20( 66( 20(1) 20( 66( 33( 20(1)

+) +) +) +) +)
33(1) 11( 33(

66(1)

50(1)

Differential-taxa of the subassociations Sclerochloa dura Climacoptera crassa Phragmites australis var. stenophylla Halopeplis pygmaea Frankenia pulverulenta Psylliostachys leptostachya
33(

+)

+) +)
33(1) 100(23) 75(2) 25(1) 25(2) 22(1) 33(23) 33(1) 25(1) 25(1) 25(1) 50(1) 11(1) 33(1) 33( 25(1) 50(1) 100( 75(23) 75(23) 50(

25(1) 25( 25(1) 33(

+)

+) 11(+)

25( 25(

+) 25(+) +)
100( 20(1) 20( 33(

+) +)
33(

+) 20(+) 20(+) +) +) 33(+)

+) +) +,1) +)

+)
33(

25(1)

+)
50( 20(1) 20(

+) +)

25(

+)

+)
11( 11(

75(

+) +) +)
25( (33)(

+) +)
25(

Companions Aeluropus littoralis Hordeum geniculatum Frankenia hirsuta Crypsis aculeata Halanthium rariorum Puccinellia bulbosa ssp. bulbosa Alhagi maurorum Atriplex tatarica Puccinellia distans Psylliostachys spicata Suaeda altisissima Parapholis incurva Spergularia marina Suaeda microphylla Limonium gmelini Heliotropium samoliorum Lepidium aucheri Erysimum sisymbrioides Plantago maritima ssp. salsa Lycium ruthenicum Koelpinia linearis

+)

159

160

Table 1. (continued) 22. Aeluropodetum littoralis, 23. Puccinellio distantis- Aeluropodetum littoralis, 24. Hordeetosum geniculati, 25. Puccinellio bulbosae- Aeluropodetum littoralis, 26. Juncetum orientalis, 27. Juncetum inexi, 28. Juncetum maritimi, 29. Juncetum libanotici, 30. Trifolio- Cynodontetum, 31. Iridetum musulmanicae, 32. CaricoJuncetum acuti, 33. Carico-Juncetum orientalis, 34. Carico-Juncetum inexi, 35. Bolboschoenetum maritimi, 36. Crypsidetosum aculeatae, 37. Alopecuretosum arundinacei, 38. Eleocharetum palustri, 39. Alismatetum plantaginis- quaticae 22 2 1 2 5 2 3 7 10 3 8 3 5 3 6 3 4 6 10 3 14 4 15 4 10 2 11 4 9 23 24 25 26 27 28 29 30 31 32 33 34 35 36 2 8 37 3 12 38 3 7 39 2 3

Association and subassociation no. No. of releves No. of species

100(5) 100(4) 100(3) 100(25) 100(3) 100(2) 100(1) 100(13) 100(45) 66( 33(1) 100(34) 100(45) 50(1) 28(1) 100(3) 66(1) 100(23) 66(12) 100(45) 100(34) 33(1) 25(1) 100(35) 100(23) 100(13) 100( 66(1) 66(1) 66(1)

66(2)

100(12) 100(23)

50(12)

33(2)

+,1)

33(2)

100(34)

100(4) 25(

+)
100(2)

50(

+)
100(23)

+,1)

75(12)

100(1) 25(1) 100(35) 100(4) 100(1) 50(1) 100(34) 100(45) 100(1) 100(5)

Character-taxa of the associations Aeluropus littoralis Puccinellia distans Limonium bellidifolium Puccinellia bulbosa ssp. bulbosa Juncus heldreichianus ssp. orientalis Inula aucheriana Saussurea salsa Juncus inexus Juncus maritimus Juncus gerardii ssp. libanoticus Cynodon dactylon Trifolium fragiferum var. pulchellum Iris spuria ssp. musulmanica Juncus acutus Bolboschoenus maritmus Eleocharis palustris Alisma plantago-aquatica
50(1) 28(

+,1)

100(4) 50(1)

50(

+)
100(34) 100(23) 66(

Differential-taxa of the subassociations Hordeum geniculatum Crypsis aculeata Alopecurus arundinaceus Beckmania eruciformis ssp. eruciformis

+)

Table 1. (continued) 22 23 24 25 2 1 2 5 2 7 3 10 3 8 3 5 3 6 3 6 4 10 3 14 4 15 4 10 2 11 4 9 2 8 3 12 3 7 2 3 26 27 28 29 30 31 32 33 34 35 36 37 38 39

Association and subassociation no. No. of releves No. of species

75(12) 50(1) 33(1) 66(1) 66(12) 66(12) 75(12) 17(1) 33( ) 50(1) 33( 33( 66(1) 50(1) 25(1) 50( 50(1) 33(1) 25(1) 50(1) 50(1)

33(1)

33(1)

50(1)

+,2) +) 25(+)
75( 25(

+)
50(

+ 33(+) 66(+) 17 (+) 33(+) +,)

50( 25(

+) 50(+) 50(+) +)
50( 25(

33( 66( 33(

50(

+) +) +) +)
33( 33( 25( 33(

14( ) 14(1) 33( 66( 33(1) 33(1) 33(1) 25(1) 25(

+ + +)
66( 33(1) ) 66(12)

+) 33(+) 33(+) +)

+) +)

+) +)

+) +) +)
100( 25(

+) +)

17(

+)

28( 28(3)

+)

+)
25(

+)
50(1) 33( 50( 25( 75(1) 75(12) 25(

+) +) +) +)
33(1) 33(

+)

14(

+)
33( 28(

+) +)
33(1)

Companions Phragmites australis var. australis Agrostis stolonifera Agropyrum elongatum Carex divisa var. ammophila Lotus tenuis Plantago major ssp. major Plantago maritima ssp. salsa Butomus umbellatus Veronica anagalis- aquatica Rumex conglomeratus Rumex crispus Phragmites australis var. stenophylla Limonium meyeri Taraxacum sp. Catabrosa aquatica Zingeria trichopoda Aster tripolium Festuca arundinacea Polypogon semiverticillata Lepidium cartilagineum ssp. pumilum Dactylorhiza umbrosa var. longibracteata Alhagi maurorum Potentilla recta Polypogon monspeliensis Batrachium trichophyllus Poa trivialis Atriplex hastata Mentha longifolia Epilobium angustifolium Tragopogon graminifolius Spergularia marina Carex distans Cirsium alatum Frankenia hirsuta Halanthium rariorum Halimione verrucifera Veronica becca-bunga
66(

+)

161

162

Figure 2. Releves ordination based on AFC (axes 13).

belong to the class Juncetea martimi. Our results support the syntaxonomical schemes proposed by Corbetta et al. (1989), Gehu et al. (1989a) and Gehu et al. (1989b), in which the perennial halo-

philous communities are referred to the class Juncetea maritimi (Table 2). The associations belonging to this class are the following (Table 1): Juncetum maritimi, Juncetum orientalis, Juncetum liban-

163
Table 2. Syntaxonomical scheme of vegetation Halocnemetea strobilacei Halocnemetalia strobilacei Halocnemion strobilacei Halocnemetum strobilacei Halostachyetum caspicae Kalidietum caspici Halopeplidetosum pygmaeae Phragmitetosum stenophyllae Frankenietosum pulverulentae Climacopteretosum crassae Psylliostachyetosum leptostachyae Halimionion Halimionetum verruciferae Limonietum meyeri Limonietum carnosi

Juncetea maritimi Juncetalia maritimi Juncion maritimi Juncetum maritimi Juncetum inexi Juncetum orientalis Puccinellietalia distantis Puccinellion bulbosae Puccinellio bulbosae-Aeluropodetum littoralis Aeluropodetum littoralis Juncetum libanotici Puccinellion distantis Puccinellio distantis-Aeluropodetum littoralis Hordeetosum geniculati

Thero-Salicornietea Thero-Salicornietalia Thero-Salicornion Salicornietum europaeae Thero-Suaedion Suaedetum maritimae Salsoletum sodae Petrosimonion Petrosimonietum brachiatae Petrosimonietum glaucae Sclerochloetosum durae Tamaricetea Tamaricetalia Tamaricion tetragynae Tamaricetum meyeri Tamaricetum octandrae Tamaricetum kotschyi Tamariceto meyeri-octandrae

Agrostietea stoloniferae Agrostietalia stoloniferae Trifolio-Cynodontion Trifolio-Cynodontetum Iridetum musulmanicae Carico-Juncion Carico-Juncetum inexi Carico-Juncetum orientalis Carico-Juncetum acuti Phragmitetea Phragmitetalia Bolboschoenion maritimi Bolboschoenetum maritimi Eleocharetum palustri Alismatetum plantaginis-aquaticae Crypsidetosum aculeatae Alopecuretosum arundinacei

otici, Puccinellio distantis-Aeluropodetum littoralis, Puccinellio bulbosae-Aeluropodetum littoralis and Aeluropodetum littoralis. The synoptic table (Table 1) shows that Puccinellio distantis-Aeluropodetum littoralis includes the Hordeetosum geniculati subassociation. Class Agrostietea stoloniferae There is a type of herbaceous perennial halotolerant vegetation accompanied with mosaics of rush plants in part of the littoral salt marsh. According to our results these communities are to be included in the class

Agrostietea stoloniferae. This is in accordance with the classication of European salt marsh vegetation (Ellenberg 1986; Canullo et al. 1988; De Foucault et al. 1992). The associations belonging to this class are the following (Table 1): Trifolio-Cynodontetum, Iridetum musulmanicae, Carico-Juncetum orientalis, Carico-Juncetum inexi and Carico-Juncetum acuti. In our opinion the syntaxonomical scheme (Table 2) needs further analysis.

164
Legend to Figure 3 . 0165 Catabrosa aquatica (L.) Beauv. 0390 Juncus inexus L. 0509 Phragmites australis (Cav.) Trin et Steud. var. stenophylla (Boiss.) Bor 0538 Potentilla recta L. 0908 Poa trivialis L. 1117 Aeluropus littoralis (Gouan) Parl. 1118 Psylliostachys spicata (Willd.) Nevski 1119 Psylliostachys leptostachya (Boiss.) Roshk. 1132 Limonium carnosum (Boiss.) O. Kuntze 1133 Halocnemum strobilaceum (Pall.) Bieb. 1141 Halanthium rariorum C. Koch. 1142 Salicornia europaea L. 1303 Suaeda maritima (L.) Dumort 1304 Petrosimonia brachiata (Pall.) Bge. 1305 Halimione verrucifera (Bieb.) Aellen 1306 Limonium meyeri (Boiss.) O. Kuntze 1307 Lepidium cartilagineum (J. Meyer) Thell. ssp. pumilum 1308 Puccinellia bulbosa (Grossh) Grossh ssp. bulbosa 1309 Juncus gerardii Loisel. ssp. libanoticus (Thieb.) Snog. 1310 Bolboschoenus maritimus (L.) Pall. 1312 Alhagi maurorum Medikus 1313 Climacoptera crassa (Bieb.) Botsch. 1314 Camphorosma monspeliaca L. 1316 Hordeum geniculatum All. 1317 Spergularia marina (L.) Griseb. 1318 Frankenia hirsuta L. 1319 Frankenia pulverulenta L. 1320 Alopecurus arundinaceus Poir. var. arundinaceus 1323 Veronica becca-bunga L. 1324 Beckmania eruciformis (L.) Host. ssp. eruciformis 1327 Zingeria trichopoda (Boiss.) P. Smirn 1328 Eleocharis palustris (L.) Roem & Schult. 1330 Batrachium tricophyllus (Chaix) Bossche 1331 Rumex crispus L. 1332 Veronica anagalis-aquatica L. 1333 Phragmites australis (Cav.) Trin ex Steud. var. australis 1335 Kalidium caspicum (L.) Ung.-Sternb. 1336 Salsola soda L. 1337 Puccinellia distans (Jacq.) Parl. 1338 Limonium bellidifolium (Goun) Dumort 1339 Lycium ruthenicum Murray 1340 Juncus heldreichianus marsson ex Parl. ssp. orientalis Snog. 1341 Inula aucheriana DC. 1343 Tamarix octandra (Bieb.) Bge. 1344 Petrosimonia glauca (Pall.) Bge 1345 Cressa cretica L. 1346 Crypsis aculeata (L.) Ait. 1348 Plantago maritima L. ssp. salsa (Pall.) Rech.f. 1349 1350 1351 1356 1358 1360 1361 1368 1369 1371 1373 1374 1375 1378 1383 1385 1386 1387 1390 1391 1392 1393 1398 1399 1400 1401 1403 1404 1405 1406 1407 1408 1411 1413 1415 1417 1418 1423 1424 1426 1432 1436 1437 1438 1445 1447 1448 1449 1451 Eremopyrum triticum (Gaerth.) Nevski Halopeplis pygmaea (Pall.) Bge. Sclerochloa dura (L.) Beauv. Tamarix kotschyi Bge. Polypogon monspeliensis (L.) Desf. Halostachys caspica (Bieb.) C.A. Mey. Koelpinia linearis Pallas Parapholis incurva (L.) C.E. Hubbard Cynodon dactylon (L.) Pers. Suaeda altissima (L.) Pall. Plantago major L. ssp. major Agrostis stolonifera L. Rumex conglomeratus Murr. Spergularia media (L.) presl. Lotus tenuis Waldst. et. Kit. Alisma plantago-aquatica L. Butomus umbellatus L. Crypsis schoenoides (L.) Lam. Cyperus fuscus L. Cyperus laevigatus L. var. distachyos (All.) Maire & Weiller Trifolium fragiferum L. var. pulchellum Lange Mentha longifolia (L.) Hudson. Carex divisa Huds. var. ammophila (Willd.) K uk Taraxacum sp. Iris spuria L. ssp. musulmanica (Fomin) Takht. Agropyrum elongatum (Host) P. Beauv. Cirsium alatum (S.G. Gmelin) Bobrov Tragopogon graminifolius DC. Festuca arundinacea Schreb. Artemisia fragrans Willd. Limonium gmelini (Willd.) O. Kuntze Juncus maritimus Lam. Suaeda acuminata (C.A. Mey) Moq. Heliotropium samoliorum Bge. Atriplex tatarica L. Epilobium angustifolium L. Tamarix tetragyna Ehrenb. var. meyeri (Boiss.) Boiss. Atriplex hastata L. Polypogon semiverticillata (Forssk.) Hyl. Suaeda microphylla (L.) Dumort Saussurea salsa (Pall.) Spreng. Juncus acutus L. Carex distans L. Tamarix ramosissima Ledeb. Scorzonera laciniata L. Erysimum sisymbrifolium C.A. Mey. Lepidium aucheri Boiss. Aster tripolium L. Dactylorhiza umbrosa (Kar. & Kir.) Nevski var. longibracteata Renz.

165

Figure 3. Species ordination based on AFC (axes 13).

Class Phragmitetea The communities characterized by hydrophilous plants on margins of salty and brackish swamps, streams, areas with high ground-water and localities where fresh water ows down into the salt marsh are to be included

in the class Phragmitetea. This class was suggested by Ellenberg (1986) and Best (1988) for the vegetation type. The associations belonging to this class are the following (Table 1): Bolboschoenetum maritimi, Alismatetum plantaginis-aquaticae and Eleocharetum palustri. The synoptic table (Table 1) shows that

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Figure 4. Dendrogram produced from CAH clustring.

167 Bolboschoenetum maritimi including two subassociations: Alopecuretosum arundinacei and Crypsidetosum aculeatae. The soil features are apparently one of the main factors inuencing the plant growth of the littoral salt marshes, the plant cover, distribution and also the zonal pattern of the vegetation types (Zahran 1977). Table 3 shows the mean values for soil characteristics in stands of different communities. The pH values show that these soils are saline and alkaline, with little differences between associations and subassociations. The high rate of evaporation results in the accumulation of salts on the surface of the soil, especially where the water table is high. The salts are mainly chlorides and sulphates. Data of the mechanical analysis reveal that most of the soils have medium and heavy texture with considerable differences between them. When data for total water-soluble salts are considered, it can be seen that the soils of the rush and herbaceous perennial halotolerant communities, and hydrophilous halotolerant communities have the lowest salt concentrations. The associations belonging to the vegetation types are the following: Trifolio-Cynodontetum, Iridetum musulmanicae, Carico-Juncetum acuti, Carico-Juncetum inexi, Carico-Juncetum orientalis, Alismatetum plantaginis-aquaticae, Bolboschoenetum maritimi, Eleocharetum palustri; while those of the semi-woody shrub and perennial halophytic communities, and annual halophytic communities such as Halocnemetum strobilacei, Limonietum meyeri, Limonietum carnosi, Halopeplidetosum pygmaeae, Phragmitetosum stenophyllae, Salicornietum europaeae and Suaedetum maritimae have the highest salt concentrations. association (Halimionetum verruciferae.) The vegetation types belong to six classes; that with the exception of classes Halocnemetea strobilacei, Phragmitetea and Tamaricetea, the others i.e. Thero-Salicornietea, Juncetea maritimi and Agrostietea stoloniferae are recorded for the rst time from Iran. The vegetation of the coastal salt marshes is generally characterized by simplicity of the structure and uniformity of the species composition like other salt marsh vegetation. Each association has one sometimes two dominants with or without associated species. They vary greatly in structure area, some are only about 4 square meters and the others are a few square kilometers. The following associations play an important role in this region: Halocnemetum strobilacei, Alhagietum maurori, Salicornietum europaeae, Tamaricetum meyeri, Tamariceto meyerioctandrae, Puccinellio distantis-Aeluropodetum littoralis, Puccinellio bulbosae-Aeluropodetum littoralis. In general, Halocnemetum strobilacei is typical of vast areas of littoral marshes with high salinity and high ground-water level. This association is very poor in species, and it is often monodominant. The associated species occur in the margins or in the transition zones of the neighbouring associations of the saltland vegetation. The growth of Halocnemum strobilaceum occurs in two forms: (1) circular patches on at tidal muddy ground, or (2) sheets of irregular shaped patches far away from shoreline. In addition to the Halocnemetum strobilacei, the stands of brushwood associations particularely, Tamaricetum meyeri and Tamariceto meyeri-octandrae are the most important vegetation units of the Orumieh lake salt marshes. They form thickets in saline habitats, saline river beds, and areas with a relatively high water-table. Alhagietum maurori is an alien type to salt habitat (Kassas & Zahran 1967). This association occupies the stands inhabited by Halimionetum verruciferae, Puccinellio distantis-Aeluropodetum littoralis and Puccinellio bulbosae-Aeluropodetum littoralis. Alhagi mourorum has a long root system that may extend several meters in depth, reaching to the permanent wet soil layers with less salinity. In the inland and littoral zones it is an abundant species dominating a characteristic salt marsh community. Therefore, it was considered by Zahran & Willis (1992) as a cumulative halophyte. Salicornietum europaeae forms a pure association on high salty and wet soils at low, frequently inundated mudbanks. It seems that a protected and more or less water-saturated muddy substratum is an essential factor

Discussion The Orumieh lake salt marshes show a highly varied vegetation pattern including a number of halotolerant or halophytic associations and subassociations that are clearly characterized from the oristic viewpoint and well dened in their ecology. Inference about the ecology of the studied vegetation have been drawn from AFC. The ordination of vegetation types along axes 13 (Figure 3) corresponds to a gradient of salinity decreasing from annual halophytic association (Salicornietum europaeae) to salt marsh brushwood association (Tamaricetum meyeri.) The moisture gradient appears with different ordination along axes 12 from hydrophilous halotolerant association (Alismatetum plantaginis-aquaticae) to perennial halophytic

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Table 3. The mean values for soil characteristics of different vegetational types Associations and subassociations Halocnemetum strobilacei Halostachyetum caspicae Kalidietum caspici Halimionetum verruciferae Limonietum meyeri Limonietum carnosi Alhagietum maurori Halopeplidetosum pygmaeae Frankenietosum pulverulentae Climacopteretosum crassae Psylliostachyetosum leptostachyae Phragmitetosum stenophyllae Salicornietum europaeae Suaedetum maritimae Salsoletum sodae Petrosimonietum brachiatae Petrosimonietum glaucae Sclerochloetosum durae Tamaricetum kotschyi Tamaricetum octandrae Tamaricetum meyeri Tamariceto meyeri-octandrae Puccinellio distantis-Aeluropodetum Puccinellio bulbosae-Aeluropodetum Aeluropodetum littoralis Juncetum orientalis Juncetum inexi Juncetum maritimi Hordeetosum geniculati Trifolio-Cynodontetum Iridetum musulmanicae Carico-Juncetum acuti Bolboschoenetum maritimi Eleocharetum palustri Alismatetum plantaginis-aquaticae Dept (cm) 050 050 050 050 030 030 050 030 030 030 030 050 030 030 030 030 030 030 050 050 050 050 030 030 030 050 050 050 030 030 050 050 030 030 030 pH Ec mmhos cm, 1 147 60.7 51.3 34.9 86.8 84.7 13.6 110.2 55.9 87.4 32.6 103.3 182 167.6 55 70.5 42.8 6 46.1 36.5 9.5 12.9 36.9 54.4 59.6 19 29 30.5 7.2 6.8 5.7 4.1 4.4 4.1 4 Textural Class SiC SiC SiL SCL LS SiL C SiCL CL CL SiL SiCL SCL SiC SiC SiC C SiC SiC-C SiL SiL SiC C SiCL SiCL L-SCL SiC-C SiL SiC SiL SiCL SiL LS LS LS Na+ Mg2+

+ Ca2+

Cl, (meq L,1 ) 1388 604 504 366 756 732 102 1192 318 772 282 1026 1750 1968 506 682 414 17.6 460 374 73 92 332 484 438 172 252 292 49 34 31 15 6 11 9.5

, SO2 4
135.7 40.7 80 45 135 177.5 25 82 240 207.2 82 129.3 204 34 62.7 51.7 62.7 53.2 35 12 32.1 40.5 79.9 95 160 36 51.8 25 23.8 41 30 25.2 35 30 32

HCO, 3 1.6 1.6 5.6 1.2 3.6 2 8.1 2.2 2.6 1.6 4.8 1.6 2 3.2 1.6 1.2 2.4 2 4 3.2 2.4 2.4 2.4 3.2 2.8 4.4 3.6 4.4 2.4 6.4 4.2 7.6 2 2.8 2.4

7.5 7.7 7.8 8.1 8.5 8 9.4 7.9 7.9 7.2 7.8 8 7.6 7.4 8.4 7.6 7.8 7.8 7.4 7.5 8 7.9 7.5 7.6 9.2 8.5 7.9 7.6 7.7 8.4 7.8 8.5 7.7 8 8.5

1240 440 385 350 860 750 135 1050 430 780 290 960 1680 1440 440 540 340 35 225 175 78 107.5 300 470 551 185 250 265 50 38 25.3 20.3 9 11.3 10

278 208.2 209 60 42 166 8 234 136 156 84 184 310 564 128 198 131 36.8 272 216 28 25 108 116 60 24 60 60 24 50 38 31 36 37 38

for the growth of this species (Halwagy & Halwagy 1977). The herbaceous perennial halophytic communities dominated by Aeluropus littoralis occupy one of the littoral and inland zones of the Orumieh lake salt marshes. Also A. littoralis forms dense patches or mats. Sometimes, it has been covered by spray-like crusts of salt, indicating that the plants may have been temporarily covered by saline water. This clearly visible in the northwest of the lake, namely around of Zanbil mount.

Edaphic factors play a paramount role in the distribution of plant associations and subassociations in the regions. Among the soil variables analysed in the present study, texture and the relative concentrations , , of Na+ , Ca2+ , Mg2+ , SO2 4 and Cl are probably the most important factors in controlling the vegetational pattern in the study area. The role of these factors in delimiting plant associations has been stressed by many authors (e.g. Abdel-Razik et al. 1984; Ayyad & El-Ghareeb 1982). Apparently, saline water table is another factor determining for establishment of plant

169 communities. Halocnemetum strobilacei, Suaedetum maritimae and Salicornietum europaeae dominate the lower marshes which are subject to periodic inundation for varying periods. The constant occurrence of these associations in this habitat may suggest that salt-water inundation plays the main role in plant distribution. Inundation seems to act mainly through increasing soil moisture and affecting soluble salts content to levels suitable for inhabitation of the plants.
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Acknowledgements We would like to express our gratitude to the Research Institute of Forests and Rangelands for providing various facilities which made this work possible. We are also grateful to Dr M Assadi and Mr V Mozaffarian for identication of some species and Mr B Hamzehee for his joint in the eld and identication of a few species.

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