the halophilous vegetation of the orumieh lake salt marshes
TRANSCRIPT
Plant Ecology 132: 155–170, 1997. 155c 1997 Kluwer Academic Publishers. Printed in Belgium.
The halophilous vegetation of the Orumieh lake salt marshes, NW. Iran
Y. Asri1 & M. Ghorbanli2
1Research Institute of Forests and Rangelands, P.O.Box 13185-116, Tehran-Iran and 2Biology Dept., TarbiatMoalem University, Mofatteh Ave., Tehran-Iran
Received 12 September 1995; accepted in revised form 28 April 1997
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 defined by physiognomic-floristic system.The following six main groups of communitiesare 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) including4 associations, (4) Rush and herbaceous perennial halophytic communities (Class Juncetea maritimi) including 7associations and 1 subassociation, (5) Rush and herbaceous perennial halotolerant communities (Class Agrostieteastoloniferae) including 5 associations, (6) Hydrophilous halotolerant communities (Class Phragmitetea) including 3associations and 2 subassociations. The soil of these communities has been analysed and their habitats are describedand discussed.
Abbreviations: AFC– Analyse Factorielle des Correspondances; CAH– Classification Ascendant Hierarchique;PCA– Principal Components Analysis.
Introduction
The coastal salt marshes comprise areas of land bor-dering the seas and lakes, more or less covered withvegetation and subject to periodic inundation by tide.They have certain qualities, which is related to theproximity to the sea and lake, that distinguishes themfrom inland salt marshes (Chapman 1977). Littoral saltmarshes are essentially fringes of inland deserts, theirlandward boundary being defined by desert conditions.Ecological factors, such as terrain or climate, can beused to delimit the littoral marshes. When there is a nar-row belt along the coast surrounded by a steep barrierof mountains (e.g. part of the studied area), the limitsare clear. But in a broad plain that stretches inland fromthe coast, there may be no distinct physiographic bar-rier. Therefore, other habitat features including veget-ation type have to be used. Vegetation characteristics,related to physiographic attributes reflecting both cli-matic and edaphic factors, provide the best single basis
for delimiting littoral salt marshes. These salt marshesmay be only a narrow belt within the reach of salt spray.They can be a hundred metres wide or they may extendinland for many kilometers (Zahran & Willis 1992).
Iran is the classical country of the great salines andkavirs (Zohary 1973). Halophytic communities of Iranare still among the most poorly known vegetation units.The distribution of halophytic communities has beendepicted cartographically by Mobayen & Tregubov(1970), Mobayen (1976, Kavir-e Lut), Freitag (1977,Turan Biosphere Reserve), Kramer (1984, Gulf Regionnear Bandar-e Rig), Carle & Frey (1977), Frey (1982,Maharlu basin) and Frey et al. (1985, northern peri-pheral region of Dasht-e kavir).
Further physiognomic and ecologic-geographicdata 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-
Gr.: 201002101, PIPS Nr. 139930 BIO2KAP
*139930 veg10516.tex; 28/08/1997; 14:39; v.7; p.1
156
Figure 1. Location map of the study area.
Meyghan) and, Akhani & Ghorbanli (1993). Therehave been relatively few investigations using thephytosociological approach on salt marshes of Iran,including floristic survey of salt desert vegetation byZohary (1963, 1973) and Leonard (1991), littoral saltmarsh vegetation by Asri et al. (1995) and Atri et al.(1995).
The study area
The study area is salt marshes of Orumieh lake betweenwestern and eastern Azarbaijan provinces (Figure 1).The total area is ca. 2000 km2; most of which is locatedin the eastern part of the lake. The average altitude isabout 1284 m. The geological substratum mainly con-sists of Alluvium, Coastal plains and Swamps corres-ponding to the Quaternary and recent times depostis.Other formations are belong to the Oligo-Miocene andCretaceous periods (Iran Oil Co. 1960). A remarkablepart of the soils belongs to Saline soils series. They arechiefly Solonchaks, which fall into the Aridisols cat-egory according to the US comprehensive system ofsoil classification (Dregne 1976). Also, there are Low-Humic Gley and Alluvial soils groups, that belong tothe Inceptisols and Entisols categories, respectively.
The area belongs to the cold-semiarid climatic zoneof Emberger’s system. According to the Gaussan’ssystem, eastern and western parts of the lake belongto hot-arid Mediterranean zone and cold steppe zone,respectively. Means of over 25 years from six meteoro-
logical stations, namely; Miandoab, Maragheh, Tabriz,Sharaf Khaneh, Orumieh and Barandoz-Chay, showthat the total annual precipitation is around 324 mm,with the maximum occuring in the winter and springmonths. Mean maximum and minimum temperaturesare 36.4 �C for July and �13:3 �C for January.
Methods
Sample releves were performed according to theBraun-Blanquet method (Mueller-Dombois & Ellen-berg 1974; Westhoff & Van der Maarel 1978). Thereleve size was determined by establishing a species-area curve in each vegetation type. The releves of herb-aceous perennial and hydrophilous halotolerant com-munities were recorded on an area of 0.24–4 m2, thoseof the halophilous communities on 0.25–16 m2.
The releve and species groups were defined bythe AFC method, using the computer program ofAnaphyto (Briane 1991). The most popular ordina-tion technique has been PCA, although the strategy ofAFC has become an effective tool for phytosociolo-gical clustering and table sorting. The AFC methoddeveloped by Benzecri (1969), allows all the points(species and releves) to be represented on the same dia-gram. The species with the most similar patterns formgroups, which are placed in, or near, groups of releveswith similar species composition. The ordination is afloristic one, and the interpretation of a floristic axis interms of environmental factors can only be tentative.Important environmental factors may be reflected ontwo or more floristic axes.
The releve and species clusters have been definedby numerical analysis of AFC data, using a CAH tech-nique included in the program. CAH is essentially aclustering procedure based on releve similarity, com-bined with a procedure for obtaining a diagonal struc-ture of clusters in the table. In the phytosociologicaltable, species and releves are ordered in such a waythat the species with a more or less similar distribu-tion pattern over the releves are grouped together and,similarly, releves with a more or less similar speciescontent are placed nex to one another. The names ofsyntaxa correspond with the codes of phytosociologic-al nomenclature (Barkman et al. 1976).
The nex step comprises the replacement of eachassociation data by a column in which for each parti-cipating species the presence degree is indicated. Sucha table is called synoptic table. After comparison of thesynoptic table with those from other types of vegeta-
veg10516.tex; 28/08/1997; 14:39; v.7; p.2
157
tion from the same region an idea can be formed aboutthe local diagnostic species groups in the table understudy. The syntaxonomical research step starts when avegetation type is to be fitted into the hierarchic syn-taxon tables.
A few samples of the upper layer of the soil weretaken from each community. Analysis of soil sampleshas been carried out following Richards (1954) andJackson (1960).
Results
The program used produces an ordination with fiveaxes. The distribution of species or releve groups isbetter revealed on axes 1–3 than others (Figures 2 and3). The floristic gradient indicated in Figure 3 can beinterpreted ecologically as an overall salinity gradientranging from low to high saline environments. Also,this gradient related to the structural complexity of thevegetation from pioneer to brushwood communities.The other floristic gradient (axes 1–2) can be inter-preted as related to moisture ranging from hydrophil-ous halotolerant communities to perennial halophyticcommunities.
Generally, six groups can be recognized on axes1–3 (Figures 2 and 3). Similarly, six species and releveclusters have been distinguished in the dendrogramsobtained with CAH method (e.g. Figure 4). Partial ana-lysis of groups resulted by AFC method is shown thateach group may be divided to some subgroups. Finally,39 subgroups are distinguished on the basis of partialanalysis. The synoptic table (Table 1) is then construc-ted, using the releve and species clusters obtained byCAH method followed by partial analysis. Accordingto the results of the numerical analysis, all the associ-ations distinguished are to be included in the followingclasses:
Class Halocnemetea strobilacei
According to the results of the numerical analysis,most of the communities are characterized by semi-woody shrub and perennial halophytes on muddy anddry salty flats should be included in the class Halocne-metea strobilacei. Our results do not confirm the pro-posed classification of Zohary (1973). According toZohary (1973) most of the halophytic communitiesof Iran should be referred to the class Halocnemeteastrobilacei irano-anatolica. It seems, this class shouldbe split into several classes. Also, our syntaxonom-
ical scheme (Table 2) is not in accordance with thescheme of European salt marsh vegetation, proposedby Chapman (1974). According to Chapman (1974)this vegetation type should be assigned in the classHalostachyetea.
The synoptic table (Table 1) clearly shows thatthe Halocnemetum strobilacei association includesthe following four subassociations of perennial andannual halophytic plants: Halopeplidetosum pygmae-ae, Phragmitetosum stenophyllae, Climacopteretosumcrassae and Frankenietosum pulverulentae. These sub-associations are distributed along a gradient of decreas-ing salt in the soil. Also, the synoptic table (Table 1)shows that Kalidietum caspici includes the Psylliosta-chyetosum leptostachyae subassociation. The syntaxo-nomical scheme of this class is shown in Table 2.
Class Thero-Salicornietea
The communities characterized by annual halophytessettled on soils subject to natural or artificial dis-turbances have been assigned to the class Thero-Salicornietea. This is in accordance with the classific-ation of European salt marsh vegetation, proposed byChapman (1974), Ellenberg (1986) and Biondi (1989).The associations belonging to this class are the follow-ing (Table 1): Salicornietum europaeae, Suaedetummaritimae, Salsoletum sodae, Petrosimonietum bra-chiatae and Petrosimonietum glaucae. Our results sug-gest the syntaxonomical scheme for this class (Table 2).
Class Tamaricetea
Zohary (1973) described the class Tamaricetea salinafor salt marsh brushwood communities of Iran. Buthe did not give a hierarchical classification for it. Theassociations belonging to the class Tamaricetea arethe following (Table 1): Tamaricetum meyeri, Tam-aricetum octandrae, Tamaricetum kotschyi and Tam-ariceto meyeri-octandrae. In all the associations asubstratum of Aeluropus littoralis occurs (Table 1).The syntaxonomical scheme of this class is shown inTable 2.
Class Juncetea maritimi
Similar the others, e.g. There are vast stands of herb-aceous perennial halophytic communities in most partsof Orumieh lake salt marshes. They are accompan-ied by mosaics or rush plants. This vegetation typeaccording to the syntaxonomical scheme (Table 2)
veg10516.tex; 28/08/1997; 14:39; v.7; p.3
158
Tabl
e1.
Syno
ptic
tabl
eof
30as
soci
atio
nsan
d9
suba
ssoc
iatio
nsin
the
Oru
mie
hla
kesa
ltm
arsh
es.
1.Sa
licor
niet
umeu
ropa
eae,
2.Su
aede
tum
mar
itim
ae,
3.Sa
lsol
etum
soda
e,4.
Petr
osim
onet
umbr
achi
atae
,5.
Petr
osim
onie
tum
glau
cae,
6.Sc
lero
chlo
etos
umdu
rae,
7.H
aloc
ne-
met
umst
robi
lace
i,8.
Clim
acop
tere
tosu
mcr
assa
e,9.
Phr
agm
iteto
sum
sten
ophy
llae,
10.
Hal
opep
lidet
osum
pygm
aeae
,11
.F
rank
enie
tosu
mpu
lver
ulen
tae,
12.
Hal
osta
chye
tum
casp
icae
,13.
Kal
idie
tum
casp
ici,
14.P
sylli
osta
chye
tum
lept
osta
chya
e,15
.Hal
imio
netu
mve
rruc
ifera
e,16
.Lim
onie
tum
carn
osi,
17.L
imon
ietu
mm
eyer
i,18
.Tam
aric
etum
kots
chyi
,19
.Tam
aric
etum
mey
eri,
20.T
amar
icet
om
eyer
i-oc
tand
rae,
21.T
amar
icet
umoc
tand
rae
Ass
ocia
tion
and
12
34
56
78
910
1112
1314
1516
1718
1920
21su
bass
ocia
tion
no.
No.
ofre
leve
s4
43
32
25
33
33
33
29
33
34
44
No.
ofsp
ecie
s3
103
64
421
34
47
34
617
104
411
127
Cha
ract
er-t
axa
ofth
eas
soci
atio
nsSa
licor
nia
euro
paea
100(
3–5)
25(1
)33
(+
)20
(+
)66
(+
)50
(1)
Suae
dam
ariti
ma
50(+
)10
0(2–
4)33
(+
)50
(+
)50
(+
)20
(+
)
Sals
ola
soda
25(+
)25
(+
)10
0(2–
3)33
(+
)
Petr
osim
onia
brac
hiat
a25
(1)
100(
3–4)
33(+
)
Petr
osim
onia
glau
ca33
(+
)10
0(3)
100(
2)20
(+
)50
(+
)
Hal
ocne
mum
stro
bila
ceum
100(
3–4)
100(
2)10
0(2)
100(
3)10
0(2)
66(1
–2)
33(2
)50
(1)
44(1
–2)
66(2
)
Suae
daac
umin
ata
40(+
�
1)
Ere
mop
yrum
triti
cum
40(+
)
Hal
osta
chys
casp
ica
100(
3–4)
Kal
idiu
mca
spic
um20
(1)
100(
3–3)
100(
3-4)
Hal
imio
neve
rruc
ifera
20(1
)66
(1)
100(
2–4)
33(1
)33
(1)
Scor
zone
rala
cini
ata
44(+
�
1)
Lep
idiu
mca
rtila
gine
umss
p.pu
milu
m44
(+
)66
(1)
Art
emis
iafr
agra
ns33
(+
)
Cam
phor
osm
am
onsp
elia
ca11
(+
)
Lim
oniu
mca
rnos
um40
(1)
100(
3)
Lim
oniu
mm
eyer
i50
(+
)55
(+
)10
0(2–
3)
Tam
arix
kots
chyi
100(
3–4)
Cre
ssa
cret
ica
100(
1–2)
Tam
arix
tetr
agyn
ava
r.m
eyer
i33
(2)
100(
3–4)
100(
2)
Sper
gula
ria
med
ia75
(+
)
Tam
arix
octa
ndra
100(
2–3)
100(
3–4)
Tam
arix
ram
osis
sim
a50
(2)
Cyp
erus
fusc
us75
(+�
)
Cry
psis
scho
enoi
des
75(1
–2)
Cyp
erus
laev
igat
usva
r.di
stac
hyos
50(+
)
veg10516.tex; 28/08/1997; 14:39; v.7; p.4
159
Tabl
e1.
(con
tinue
d)
Ass
ocia
tion
and
12
34
56
78
910
1112
1314
1516
1718
1920
21su
bass
ocia
tion
no.
No.
ofre
leve
s4
43
32
25
33
33
33
29
33
34
44
No.
ofsp
ecie
s3
103
64
421
34
47
34
617
104
411
127
Dif
fere
ntia
l-ta
xaof
the
suba
ssoc
iati
ons
Scle
roch
loa
dura
100(
3)20
(+
)33
(+
)
Clim
acop
tera
cras
sa25
(2)
66(+
�
1)20
(+
)10
0(2–
3)33
(+
)50
(1)
Phr
agm
ites
aust
ralis
var.
sten
ophy
lla10
0(2–
3)11
(1)
66(1
)50
(1)
Hal
opep
lispy
gmae
a10
0(2)
Fra
nken
iapu
lver
ulen
ta10
0(4–
5)33
(+
)
Psy
llios
tach
ysle
ptos
tach
ya33
(+
)50
(+
)20
(+
)66
(+
)33
(1)
100(
3)11
(+
)33
(+
)
Com
pani
ons
Ael
urop
uslit
tora
lis20
(1)
33(1
)33
(1)
100(
2–3)
75(2
)75
(2–3
)75
(2–3
)
Hor
deum
geni
cula
tum
25(1
)20
(+
)11
(+
)25
(1)
50(+
)
Fra
nken
iahi
rsut
a25
(+
)20
(+
)66
(+
)11
(+
)25
(+
)
Cry
psis
acul
eata
25(+
)20
(+
)33
(+)
25(2
)25
(+
)
Hal
anth
ium
rari
floru
m25
(1)
33(+
)10
0(
+
)33
(+
)
Puc
cine
llia
bulb
osa
ssp.
bulb
osa
20(1
)22
(1)
25(1
)25
(1)
Alh
agim
auro
rum
33(2
–3)
33(1
)25
(1)
50(1
)
Atr
iple
xta
tari
ca25
(1)
50(1
)10
0(
+�
1)
Puc
cine
llia
dist
ans
20(1
)11
(1)
33(1
)
Psy
llios
tach
yssp
icat
a20
(+
)33
(+
)
Suae
daal
tisis
sim
a33
(+
)25
(1)
Para
phol
isin
curv
a33
(+
)50
(+
)
Sper
gula
ria
mar
ina
25(+
)75
(+
)
Suae
dam
icro
phyl
la33
(+
)
Lim
oniu
mgm
elin
i20
(1)
Hel
iotr
opiu
msa
mol
iflor
um20
(+
)
Lepi
dium
auch
eri
11(+
)
Ery
sim
umsi
sym
brio
ides
11(+
)
Pla
ntag
om
ariti
ma
ssp.
sals
a25
(+
)
Lyci
umru
then
icum
25(+
)
Koe
lpin
ialin
eari
s(3
3)(+
)
veg10516.tex; 28/08/1997; 14:39; v.7; p.5
160
Tabl
e1.
(con
tinue
d)22
.A
elur
opod
etum
litto
ralis
,23
.P
ucci
nelli
odi
stan
tis-
Ael
urop
odet
umlit
tora
lis,
24.
Hor
deet
osum
geni
cula
ti,25
.Puc
cine
llio
bulb
osae
-A
elur
opod
etum
litto
ralis
,26
.Jun
cetu
mor
ient
alis
,27.
Junc
etum
infle
xi,2
8.Ju
ncet
umm
ariti
mi,
29.J
unce
tum
liban
otic
i,30
.Tri
folio
-C
ynod
onte
tum
,31.
Irid
etum
mus
ulm
anic
ae,3
2.C
aric
o-Ju
ncet
umac
uti,
33.
Car
ico-
Junc
etum
orie
ntal
is,
34.
Car
ico-
Junc
etum
infle
xi,
35.
Bol
bosc
hoen
etum
mar
itim
i,36
.C
ryps
idet
osum
acul
eata
e,37
.A
lope
cure
tosu
mar
undi
nace
i,38
.Ele
ocha
retu
mpa
lust
ri,3
9.A
lism
atet
umpl
anta
gini
s-qu
atic
ae
Ass
ocia
tion
and
2223
2425
2627
2829
3031
3233
3435
3637
3839
suba
ssoc
iatio
nno
.N
o.of
rele
ves
22
27
33
33
63
44
24
23
32
No.
ofsp
ecie
s1
53
108
56
410
1415
1011
98
127
3
Cha
ract
er-t
axa
ofth
eas
soci
atio
nsA
elur
opus
litto
ralis
100(
5)10
0(4)
100(
3)10
0(2–
5)66
(2)
100(
1–2)
100(
2–3)
50(1
–2)
33(2
)
Puc
cine
llia
dist
ans
100(
3)10
0(2)
66(1
)66
(1)
Lim
oniu
mbe
llidi
foliu
m10
0(1)
Puc
cine
llia
bulb
osa
ssp.
bulb
osa
100(
1–3)
66(1
)
Junc
ushe
ldre
ichi
anus
ssp.
orie
ntal
is10
0(4–
5)33
(2)
100(
3–4)
Inul
aau
cher
iana
66(+
�
1)
Saus
sure
asa
lsa
33(1
)
Junc
usin
flexu
s10
0(3–
4)10
0(4)
Junc
usm
ariti
mus
100(
4–5)
Junc
usge
rard
iiss
p.lib
anot
icus
50(1
)28
(1)
100(
3)66
(1)
25(1
)25
(+
)50
(+
)
Cyn
odon
dact
ylon
100(
3–5)
100(
2–3)
100(
1–3)
100(
2)10
0(2–
3)
Trifo
lium
frag
iferu
mva
r.pu
lche
llum
100(
2–3)
66(1
–2)
100(
+�
1)75
(1–2
)10
0(1)
Iris
spur
iass
p.m
usul
man
ica
100(
4–5)
Junc
usac
utus
100(
3–4)
25(1
)
Bol
bosc
hoen
usm
aritm
us50
(1)
28(+
�
1)10
0(3–
5)10
0(4)
100(
3–4)
Ele
ocha
ris
palu
stri
s33
(1)
100(
1)10
0(4–
5)
Alis
ma
plan
tago
-aqu
atic
a50
(1)
100(
1)10
0(5)
Dif
fere
ntia
l-ta
xaof
the
suba
ssoc
iati
ons
Hor
deum
geni
cula
tum
100(
4)50
(+
)
Cry
psis
acul
eata
50(1
)10
0(3–
4)
Alo
pecu
rus
arun
dina
ceus
100(
2–3)
Bec
kman
iaer
ucifo
rmis
ssp.
eruc
iform
is66
(+
)
veg10516.tex; 28/08/1997; 14:39; v.7; p.6
161
Tabl
e1.
(con
tinue
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veg10516.tex; 28/08/1997; 14:39; v.7; p.7
162
Figure 2. Releves ordination based on AFC (axes 1–3).
belong to the class Juncetea martimi. Our resultssupport the syntaxonomical schemes proposed byCorbetta et al. (1989), Gehu et al. (1989a) andGehu et al. (1989b), in which the perennial halo-
philous communities are referred to the class Jun-cetea maritimi (Table 2). The associations belong-ing to this class are the following (Table 1): Jun-cetum maritimi, Juncetum orientalis, Juncetum liban-
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163
Table 2. Syntaxonomical scheme of vegetation
Halocnemetea strobilacei
Halocnemetalia strobilacei Juncetea maritimi
Halocnemion strobilacei Juncetalia maritimi
Halocnemetum strobilacei Juncion maritimi
Halostachyetum caspicae Juncetum maritimi
Kalidietum caspici Juncetum inflexi
Halopeplidetosum pygmaeae Juncetum orientalis
Phragmitetosum stenophyllae Puccinellietalia distantis
Frankenietosum pulverulentae Puccinellion bulbosae
Climacopteretosum crassae Puccinellio bulbosae-Aeluropodetum littoralis
Psylliostachyetosum leptostachyae Aeluropodetum littoralis
Halimionion Juncetum libanotici
Halimionetum verruciferae Puccinellion distantis
Limonietum meyeri Puccinellio distantis-Aeluropodetum littoralis
Limonietum carnosi Hordeetosum geniculati
Thero-Salicornietea Agrostietea stoloniferae
Thero-Salicornietalia Agrostietalia stoloniferae
Thero-Salicornion Trifolio-Cynodontion
Salicornietum europaeae Trifolio-Cynodontetum
Thero-Suaedion Iridetum musulmanicae
Suaedetum maritimae Carico-Juncion
Salsoletum sodae Carico-Juncetum inflexi
Petrosimonion Carico-Juncetum orientalis
Petrosimonietum brachiatae Carico-Juncetum acuti
Petrosimonietum glaucae
Sclerochloetosum durae Phragmitetea
Phragmitetalia
Tamaricetea Bolboschoenion maritimi
Tamaricetalia Bolboschoenetum maritimi
Tamaricion tetragynae Eleocharetum palustri
Tamaricetum meyeri Alismatetum plantaginis-aquaticae
Tamaricetum octandrae Crypsidetosum aculeatae
Tamaricetum kotschyi Alopecuretosum arundinacei
Tamariceto meyeri-octandrae
otici, Puccinellio distantis-Aeluropodetum littoral-is, Puccinellio bulbosae-Aeluropodetum littoralis andAeluropodetum littoralis. The synoptic table (Table 1)shows that Puccinellio distantis-Aeluropodetum littor-alis includes the Hordeetosum geniculati subassoci-ation.
Class Agrostietea stoloniferae
There is a type of herbaceous perennial halotolerantvegetation accompanied with mosaics of rush plantsin part of the littoral salt marsh. According to ourresults these communities are to be included in the class
Agrostietea stoloniferae. This is in accordance withthe classification of European salt marsh vegetation(Ellenberg 1986; Canullo et al. 1988; De Foucaultet al. 1992). The associations belonging to this classare the following (Table 1): Trifolio-Cynodontetum,Iridetum musulmanicae, Carico-Juncetum orientalis,Carico-Juncetum inflexi and Carico-Juncetum acuti.In our opinion the syntaxonomical scheme (Table 2)needs further analysis.
veg10516.tex; 28/08/1997; 14:39; v.7; p.9
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Legend to Figure 3 .
0165 Catabrosa aquatica (L.) Beauv. 1349 Eremopyrum triticum (Gaerth.) Nevski
0390 Juncus inflexus L. 1350 Halopeplis pygmaea (Pall.) Bge.
0509 Phragmites australis (Cav.) Trin et Steud. 1351 Sclerochloa dura (L.) Beauv.
var. stenophylla (Boiss.) Bor 1356 Tamarix kotschyi Bge.
0538 Potentilla recta L. 1358 Polypogon monspeliensis (L.) Desf.
0908 Poa trivialis L. 1360 Halostachys caspica (Bieb.) C.A. Mey.
1117 Aeluropus littoralis (Gouan) Parl. 1361 Koelpinia linearis Pallas
1118 Psylliostachys spicata (Willd.) Nevski 1368 Parapholis incurva (L.) C.E. Hubbard
1119 Psylliostachys leptostachya (Boiss.) Roshk. 1369 Cynodon dactylon (L.) Pers.
1132 Limonium carnosum (Boiss.) O. Kuntze 1371 Suaeda altissima (L.) Pall.
1133 Halocnemum strobilaceum (Pall.) Bieb. 1373 Plantago major L. ssp. major
1141 Halanthium rariflorum C. Koch. 1374 Agrostis stolonifera L.
1142 Salicornia europaea L. 1375 Rumex conglomeratus Murr.
1303 Suaeda maritima (L.) Dumort 1378 Spergularia media (L.) presl.
1304 Petrosimonia brachiata (Pall.) Bge. 1383 Lotus tenuis Waldst. et. Kit.
1305 Halimione verrucifera (Bieb.) Aellen 1385 Alisma plantago-aquatica L.
1306 Limonium meyeri (Boiss.) O. Kuntze 1386 Butomus umbellatus L.
1307 Lepidium cartilagineum (J. Meyer) Thell. 1387 Crypsis schoenoides (L.) Lam.
ssp. pumilum 1390 Cyperus fuscus L.
1308 Puccinellia bulbosa (Grossh) Grossh ssp. bulbosa 1391 Cyperus laevigatus L. var. distachyos (All.) Maire
1309 Juncus gerardii Loisel. ssp. libanoticus (Thieb.) & Weiller
Snog. 1392 Trifolium fragiferum L. var. pulchellum Lange
1310 Bolboschoenus maritimus (L.) Pall. 1393 Mentha longifolia (L.) Hudson.
1312 Alhagi maurorum Medikus 1398 Carex divisa Huds. var. ammophila (Willd.) Kuk
1313 Climacoptera crassa (Bieb.) Botsch. 1399 Taraxacum sp.
1314 Camphorosma monspeliaca L. 1400 Iris spuria L. ssp. musulmanica (Fomin) Takht.
1316 Hordeum geniculatum All. 1401 Agropyrum elongatum (Host) P. Beauv.
1317 Spergularia marina (L.) Griseb. 1403 Cirsium alatum (S.G. Gmelin) Bobrov
1318 Frankenia hirsuta L. 1404 Tragopogon graminifolius DC.
1319 Frankenia pulverulenta L. 1405 Festuca arundinacea Schreb.
1320 Alopecurus arundinaceus Poir. var. arundinaceus 1406 Artemisia fragrans Willd.
1323 Veronica becca-bunga L. 1407 Limonium gmelini (Willd.) O. Kuntze
1324 Beckmania eruciformis (L.) Host. ssp. eruciformis 1408 Juncus maritimus Lam.
1327 Zingeria trichopoda (Boiss.) P. Smirn 1411 Suaeda acuminata (C.A. Mey) Moq.
1328 Eleocharis palustris (L.) Roem & Schult. 1413 Heliotropium samoliflorum Bge.
1330 Batrachium tricophyllus (Chaix) Bossche 1415 Atriplex tatarica L.
1331 Rumex crispus L. 1417 Epilobium angustifolium L.
1332 Veronica anagalis-aquatica L. 1418 Tamarix tetragyna Ehrenb. var. meyeri (Boiss.)
1333 Phragmites australis (Cav.) Trin ex Steud. Boiss.
var. australis 1423 Atriplex hastata L.
1335 Kalidium caspicum (L.) Ung.-Sternb. 1424 Polypogon semiverticillata (Forssk.) Hyl.
1336 Salsola soda L. 1426 Suaeda microphylla (L.) Dumort
1337 Puccinellia distans (Jacq.) Parl. 1432 Saussurea salsa (Pall.) Spreng.
1338 Limonium bellidifolium (Goun) Dumort 1436 Juncus acutus L.
1339 Lycium ruthenicum Murray 1437 Carex distans L.
1340 Juncus heldreichianus marsson ex Parl. 1438 Tamarix ramosissima Ledeb.
ssp. orientalis Snog. 1445 Scorzonera laciniata L.
1341 Inula aucheriana DC. 1447 Erysimum sisymbrifolium C.A. Mey.
1343 Tamarix octandra (Bieb.) Bge. 1448 Lepidium aucheri Boiss.
1344 Petrosimonia glauca (Pall.) Bge 1449 Aster tripolium L.
1345 Cressa cretica L. 1451 Dactylorhiza umbrosa (Kar. & Kir.) Nevski
1346 Crypsis aculeata (L.) Ait. var. longibracteata Renz.
1348 Plantago maritima L. ssp. salsa (Pall.) Rech.f.
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Figure 3. Species ordination based on AFC (axes 1–3).
Class Phragmitetea
The communities characterized by hydrophilous plantson margins of salty and brackish swamps, streams,areas with high ground-water and localities where freshwater flows down into the salt marsh are to be included
in the class Phragmitetea. This class was suggestedby Ellenberg (1986) and Best (1988) for the vegeta-tion type. The associations belonging to this class arethe following (Table 1): Bolboschoenetum maritimi,Alismatetum plantaginis-aquaticae and Eleocharetumpalustri. The synoptic table (Table 1) shows that
veg10516.tex; 28/08/1997; 14:39; v.7; p.11
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Figure 4. Dendrogram produced from CAH clustring.
veg10516.tex; 28/08/1997; 14:39; v.7; p.12
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Bolboschoenetum maritimi including two subassoci-ations: Alopecuretosum arundinacei and Crypside-tosum aculeatae.
The soil features are apparently one of the mainfactors influencing the plant growth of the littoral saltmarshes, the plant cover, distribution and also the zonalpattern of the vegetation types (Zahran 1977). Table 3shows the mean values for soil characteristics in standsof different communities. The pH values show thatthese soils are saline and alkaline, with little differ-ences between associations and subassociations. Thehigh rate of evaporation results in the accumulation ofsalts on the surface of the soil, especially where thewater table is high. The salts are mainly chlorides andsulphates. Data of the mechanical analysis reveal thatmost of the soils have medium and heavy texture withconsiderable differences between them. When data fortotal water-soluble salts are considered, it can be seenthat the soils of the rush and herbaceous perennial halo-tolerant communities, and hydrophilous halotolerantcommunities have the lowest salt concentrations. Theassociations belonging to the vegetation types are thefollowing: Trifolio-Cynodontetum, Iridetum musul-manicae, Carico-Juncetum acuti, Carico-Juncetuminflexi, Carico-Juncetum orientalis, Alismatetumplantaginis-aquaticae, Bolboschoenetum maritimi,Eleocharetum palustri; while those of the semi-woodyshrub and perennial halophytic communities, andannual halophytic communities such as Halocnemetumstrobilacei, Limonietum meyeri, Limonietum carnosi,Halopeplidetosum pygmaeae, Phragmitetosum sten-ophyllae, Salicornietum europaeae and Suaedetummaritimae have the highest salt concentrations.
Discussion
The Orumieh lake salt marshes show a highly variedvegetation pattern including a number of halotolerantor halophytic associations and subassociations that areclearly characterized from the floristic viewpoint andwell defined in their ecology. Inference about the eco-logy of the studied vegetation have been drawn fromAFC. The ordination of vegetation types along axes1–3 (Figure 3) corresponds to a gradient of salinitydecreasing from annual halophytic association (Sali-cornietum europaeae) to salt marsh brushwood asso-ciation (Tamaricetum meyeri.) The moisture gradi-ent appears with different ordination along axes 1–2from hydrophilous halotolerant association (Alismat-etum plantaginis-aquaticae) to perennial halophytic
association (Halimionetum verruciferae.) The vegeta-tion types belong to six classes; that with the exceptionof classes Halocnemetea strobilacei, Phragmitetea andTamaricetea, the others i.e. Thero-Salicornietea, Jun-cetea maritimi and Agrostietea stoloniferae are recor-ded for the first time from Iran.
The vegetation of the coastal salt marshes is gen-erally characterized by simplicity of the structureand uniformity of the species composition like oth-er salt marsh vegetation. Each association has onesometimes two dominants with or without associ-ated species. They vary greatly in structure area,some are only about 4 square meters and the othersare a few square kilometers. The following associ-ations play an important role in this region: Halocne-metum strobilacei, Alhagietum maurori, Salicornietumeuropaeae, Tamaricetum meyeri, Tamariceto meyeri-octandrae, Puccinellio distantis-Aeluropodetum littor-alis, Puccinellio bulbosae-Aeluropodetum littoralis.
In general, Halocnemetum strobilacei is typical ofvast areas of littoral marshes with high salinity andhigh ground-water level. This association is very poorin species, and it is often monodominant. The associ-ated species occur in the margins or in the transitionzones of the neighbouring associations of the saltlandvegetation. The growth of Halocnemum strobilaceumoccurs in two forms: (1) circular patches on flat tid-al muddy ground, or (2) sheets of irregular shapedpatches far away from shoreline.
In addition to the Halocnemetum strobilacei, thestands of brushwood associations particularely, Tam-aricetum meyeri and Tamariceto meyeri-octandrae arethe most important vegetation units of the Orumiehlake salt marshes. They form thickets in saline habit-ats, saline river beds, and areas with a relatively highwater-table.
Alhagietum maurori is an alien type to salt hab-itat (Kassas & Zahran 1967). This association occu-pies the stands inhabited by Halimionetum verrucifer-ae, Puccinellio distantis-Aeluropodetum littoralis andPuccinellio bulbosae-Aeluropodetum littoralis. Alhagimourorum has a long root system that may extend sev-eral meters in depth, reaching to the permanent wet soillayers with less salinity. In the inland and littoral zonesit is an abundant species dominating a characteristicsalt marsh community. Therefore, it was consideredby Zahran & Willis (1992) as a cumulative halophyte.
Salicornietum europaeae forms a pure associationon high salty and wet soils at low, frequently inundatedmudbanks. It seems that a protected and more or lesswater-saturated muddy substratum is an essential factor
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Table 3. The mean values for soil characteristics of different vegetational types
Associations Dept pH Ec Textural Na+ Mg2++ Ca2+ Cl� SO2�
4 HCO�3and subassociations (cm) mmhos cm�1 Class (meq L�1)
Halocnemetum strobilacei 0–50 7.5 147 SiC 1240 278 1388 135.7 1.6
Halostachyetum caspicae 0–50 7.7 60.7 SiC 440 208.2 604 40.7 1.6
Kalidietum caspici 0–50 7.8 51.3 SiL 385 209 504 80 5.6
Halimionetum verruciferae 0–50 8.1 34.9 SCL 350 60 366 45 1.2
Limonietum meyeri 0–30 8.5 86.8 LS 860 42 756 135 3.6
Limonietum carnosi 0–30 8 84.7 SiL 750 166 732 177.5 2
Alhagietum maurori 0–50 9.4 13.6 C 135 8 102 25 8.1
Halopeplidetosum pygmaeae 0–30 7.9 110.2 SiCL 1050 234 1192 82 2.2
Frankenietosum pulverulentae 0–30 7.9 55.9 CL 430 136 318 240 2.6
Climacopteretosum crassae 0–30 7.2 87.4 CL 780 156 772 207.2 1.6
Psylliostachyetosum leptostachyae 0–30 7.8 32.6 SiL 290 84 282 82 4.8
Phragmitetosum stenophyllae 0–50 8 103.3 SiCL 960 184 1026 129.3 1.6
Salicornietum europaeae 0–30 7.6 182 SCL 1680 310 1750 204 2
Suaedetum maritimae 0–30 7.4 167.6 SiC 1440 564 1968 34 3.2
Salsoletum sodae 0–30 8.4 55 SiC 440 128 506 62.7 1.6
Petrosimonietum brachiatae 0–30 7.6 70.5 SiC 540 198 682 51.7 1.2
Petrosimonietum glaucae 0–30 7.8 42.8 C 340 131 414 62.7 2.4
Sclerochloetosum durae 0–30 7.8 6 SiC 35 36.8 17.6 53.2 2
Tamaricetum kotschyi 0–50 7.4 46.1 SiC-C 225 272 460 35 4
Tamaricetum octandrae 0–50 7.5 36.5 SiL 175 216 374 12 3.2
Tamaricetum meyeri 0–50 8 9.5 SiL 78 28 73 32.1 2.4
Tamariceto meyeri-octandrae 0–50 7.9 12.9 SiC 107.5 25 92 40.5 2.4
Puccinellio distantis-Aeluropodetum 0–30 7.5 36.9 C 300 108 332 79.9 2.4
Puccinellio bulbosae-Aeluropodetum 0–30 7.6 54.4 SiCL 470 116 484 95 3.2
Aeluropodetum littoralis 0–30 9.2 59.6 SiCL 551 60 438 160 2.8
Juncetum orientalis 0–50 8.5 19 L-SCL 185 24 172 36 4.4
Juncetum inflexi 0–50 7.9 29 SiC-C 250 60 252 51.8 3.6
Juncetum maritimi 0–50 7.6 30.5 SiL 265 60 292 25 4.4
Hordeetosum geniculati 0–30 7.7 7.2 SiC 50 24 49 23.8 2.4
Trifolio-Cynodontetum 0–30 8.4 6.8 SiL 38 50 34 41 6.4
Iridetum musulmanicae 0–50 7.8 5.7 SiCL 25.3 38 31 30 4.2
Carico-Juncetum acuti 0–50 8.5 4.1 SiL 20.3 31 15 25.2 7.6
Bolboschoenetum maritimi 0–30 7.7 4.4 LS 9 36 6 35 2
Eleocharetum palustri 0–30 8 4.1 LS 11.3 37 11 30 2.8
Alismatetum plantaginis-aquaticae 0–30 8.5 4 LS 10 38 9.5 32 2.4
for the growth of this species (Halwagy & Halwagy1977).
The herbaceous perennial halophytic communitiesdominated by Aeluropus littoralis occupy one of the lit-toral and inland zones of the Orumieh lake salt marshes.Also A. littoralis forms dense patches or mats. Some-times, it has been covered by spray-like crusts of salt,indicating that the plants may have been temporarilycovered by saline water. This clearly visible in thenorthwest of the lake, namely around of Zanbil mount.
Edaphic factors play a paramount role in the dis-tribution of plant associations and subassociations inthe regions. Among the soil variables analysed in thepresent study, texture and the relative concentrationsof Na+, Ca2+, Mg2+, SO2�
4 and Cl� are probably themost important factors in controlling the vegetation-al pattern in the study area. The role of these factorsin delimiting plant associations has been stressed bymany authors (e.g. Abdel-Razik et al. 1984; Ayyad& El-Ghareeb 1982). Apparently, saline water table isanother factor determining for establishment of plant
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communities. Halocnemetum strobilacei, Suaedetummaritimae and Salicornietum europaeae dominate thelower marshes which are subject to periodic inundationfor varying periods. The constant occurrence of theseassociations in this habitat may suggest that salt-waterinundation plays the main role in plant distribution.Inundation seems to act mainly through increasing soilmoisture and affecting soluble salts content to levelssuitable for inhabitation of the plants.
Acknowledgements
We would like to express our gratitude to the ResearchInstitute of Forests and Rangelands for providing vari-ous facilities which made this work possible. We arealso grateful to Dr M Assadi and Mr V Mozaffarian foridentification of some species and Mr B Hamzehee forhis joint in the field and identification of a few species.
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