Wolff - The Halophylous Vegetation

THE HALOPHILOUS VEGETATION OF THE LAGOONS
OF MESOLONGHI, GREECE
by
W. J. WOLFF
Yerseke
INTRODUCTION
PREVIOUS STUDIES ON MEDITERRANEAN SALT VEGETATIONS
Salt-marshes cover extensive areas of the mediterranean region.

However, compared with the amount of research done on the
atlantic coasts of Europe, the study of the mediterranean salt
vegetations is still in its infancy.
The best known area is undoubtedly the mediterranean coast of
France. Here the investigations of BRAUN-BLANQUEThave been
the basis of all research on mediterranean salt vegetations.
Of the west-mediterranean area a fair amount of local studies
is known, e.g. from the Ebro plains, the Balearics, Algeria, Tuni-
sia, Corsica, Sicily and the Gulf of Venice. Our knowledge of the
east-mediterranean region is much poorer. The only detailed stu-
dies are from the Dalmatian coast of Jugoslavia, northeastern
Greece and Egypt. EIG (I 946) gives only an overall picture of the
Israelian salt vegetations.
In the Balkan Peninsula the halophilous vegetations of the Medi-
terranean connect with those from the inner parts of Asia and
Central-Europe. Therefore we have to consider those vegetations
too. Large inland saltmarshes occur in Austria, Jugoslavia, Hungary
and Rumania. The present state of knowledge was summarized by
WENDELBERGER (I950). These vegetations may be considered as
radiations of the south-russian and central-asiatic saltsteppes. There
are, however, traces of vegetations of the atlantic coasts of Europe
(BEEFTINK I965). These pontic-pannonic vegetations reach the
mediterranean area on the shores of the Black Sea in Rumania.
From here related vegetations radiate toward the Greek coasts,
where OBERDORFER (I952) proposed a S t a t i c i o n o r i e n t a l e as a
transition between the west-mediterranean vegetations and the
asiatic H a l o s t a c h y e t a l i a .
THE PRESENT INVESTIGATION
To obtain a clearer picture of the east-mediterranean halophi-

lous vegetations we chose the western coast of Greece, since the
study of OBERDORFER (1.C.) had already covered the coasts of Thes-
96 W.J. WOLFF
saly, Macedonia and Thrace. The lagoons of Mesolonghi were

chosen because this area will be reclaimed in the near future. We
stayed in the area from April 1963 until July 1963, and in June
i964. In 1964 we also visited the lagoons of Arta on the northern
shore of the Ambracian Gulf and the delta of the river Axios where
OBF.I~I~ORF~I~ made most of his observations (AMos is the Greek
equivalent of the Yugoslav Vardar). Moreover some observations
from other parts of Greece are incorporated in this study.
The aims of this study were:
a qualitative and quantitative description, as complete as possible,
of the halophilous plant-communities of the lagoons of Mesolonghi;
a study of the ecological factors which are responsible for the ob-
served composition and pattern of the vegetation;
a study of the relationship of the observed communities (syntaxa)
with those of other parts of the Mediterranean and
an investigation into the value of the studied area as a nature-re-
serve.
This aspect has been dealt with in a separate paper (WOLFF 1966A,
I966B).
METHODS
The vegetation was studied in accordance with the concepts
and with the methods of the Zfirich-Montpellier school (BRAUN-
BLANQUET I95I , ELLENBERG I956 ). The scale of cover-abundance
used was slightly modified according to S~.GAL & BARK~AN (I960)
(2m = many individuals, but cover less than 5%; 2a = cover
5--12.5 % and 2b = cover 12.5--25 %). As our investigations
have only a preliminary scope the study of the acting ecological
factors has been confined to those recognizable in the field. Only
the salinity of some waters was measured more accurately.
NOMENCLATURE
The nomenclature of the vascular plants follows in general the
flora of HAYEK (I927--I933), except for the following taxa:
Halimione portulacoides (L.) A~LL. Salicorniaradicans S~.
Ranunculus trichophyllus CHALX. Medicago polymorpha L.
Statice limonium L. ssp. serotinum (RcHB.) GAMS.
Anagallis arvensis L. ssp. coerulea (GouAN) VOLLM.
Hedypnois rhagadioloides (L.) WILLD. Juncus arnbiguus Guss.
Scirpus cernuus VAHL. Scirpus litoralis SCHRAD.
Scirpus maritimus L. Cyperus distachyus ALL.
Carex extensa Gooo. var. graeca HAUBKNEGHT.
Elytrigia juncea (L.) N~vsKi ssp. mediterraneumSIMO~ET
HALOPHILOUS VEGETATION OF MESOLONGHI 97
Hordeum marinum HuDs. Puccinelliaf estucif ormis

HOST.
Parapholis incurva (L.) HUBB. Parapholis strigosa (DUM.)
HUBB.
For authorities concerning the taxa mentioned in the text we
refer to the flora and the list mentioned above.
The only observed moss is: TorteUaflaviformis (BRucH.) BROTH.
The only observed lichen is: Xanthoria parietina (L.) TH. FR.
The plant-communities have been named according to the
Braun-Blanquet system. Authority names and year of publication
are mentioned together with the name of the syntaxon.
ACKNOWLEDGEMENT
Our investigations were carried out under the guidance of Dr.
J . J . BARKMAN (Wijster) who was always willing to give advice and
information.
Dr. W. G. BEEFTINK (Yerseke) and Prof. Dr. V. WESTHOFF
(Zeist) gave valuable advice and important criticism.
Mr. D. A. FERNANDES (Grontmij Ltd., De Bilt; this firm re-
claimed a pilot polder in the area) permitted me to read internal
reports on the lagoons of Mesolonghi and moreover send me a copy
of an aerial photograph. Prof. Dr. J. BRAUN-BLAN~UET (Montpel-
lier) made it possible for me to study the many data collected by
him and his collaborators during a stay of a few days in Mont-
pellier. Dr. G. I. LAWRENTIADES(Salonica) very kindly gave us an
introduction into botanical research in Greece. Prof. Dr. H. A.
DIAPOULIS (Athens) and Dr. S.J. VAN OOSTSTROOMand the late Mr.
TH. REICHGELT (Leiden) helped me with the identification of m a n y
plants. Mr. A. T o u w (Leiden) identified our only collected moss
and Mr. E. HENNIPMAN (Leiden) kindly suggested the name of a
lichen. Mr. T. MIEDEMA (Mesolonghi - Vught) presented data on
the hydrology of the lagoons. Prof. Dr. R. Tf3XEN (Stolzenau), Prof.
Dr. V. WESTHOFF (Zeist) and Dr. A. D. HATZIKAKIDES(Athens) sent
me literature concerning vegetation and hydrology of the lagoons.
Dr. C. DEN HARTOG (Leiden) gave information about waterplants
and water-plant communities. The fund "Mr. P. Vollenhoven" of
the State University of Leiden gave financial support. Mr. SHMUEL
BAR-EVEN (Magdn) and Dr. K. F. VAAS (Yerseke) kindly corrected
our English translation.
We would like to thank them all for their invaluable help. Lastly,
we should like to make a special mention of thanks to the inhabi-
tants of the region in which we made our investigations. Without
their hospitality and friendship the perpetual combat with mos-
quitos, dirt and heat would have been difficult to win.
,, I %
+ 7os
L.
.~.
4
,#, PLEURON
;" / - HAG ~.G. "~IA~IA

0
;r,,, ; <;:~>
i ,
7
/
GULF oF PATRAS
Fig. za. The lagoons of Mesolonghi.
H A L O P H I L O U S V E G E T A T I O N OF M E S O L O N G H I 99
Fig. Ib. Greece. The areas where halophilous vegetations were studied, are
shown in black.
ioo W.J. WOLFF
MILIEU
SITUATION AND TOPOGRAPHY OF THE AREA
The lagoons of Mesolonghi are situated in the province of Aito-

lo-Acarnania in south-west Greece around the town of Mesolonghi.
They are connected by a large opening with the Gulf of Patras
which forms part of the Ionian Sea.
The lagoons lie behind a fragmented sandy coastal ridge between
the mouth of the river Evinos and the hill Koutsilaris. Our obser-
vations were made in particular in the vicinity of the village NIa-
goula, but we also visited several localities scattered all over the
lagoons.
The lagoons of Arta lie in western Greece in the province of Arta
(Epirus) on the northern shore of the Ambracian Gulf (Gulf of
Arta), which also is connected with the Ionian Sea. Here too the
lagoons are separated from the sea by a sandy coastal ridge. Our
observations were chiefly made in the vicinity of the former village
Salaora and near Koronisia.
The deltas of the rivers Gallikos, Axios (called Vardar by OBER-
DORFER)and Aliakmon form together a large alluvial plain west
of Salonica (Thessaloniki). Here the landscape near the sea con-
sists of salt-marshes traversed by several old and new riverbeds.
Since the observations of OBERDORFER the area has been greatly
changed by large scale landreclamation. Our observations were made
near Pyrgos between the mouths of the Axios and the Gallikos.
For a survey of the area of Mesolonghi see fig. I a. For the situation
of the other areas see map Ib.
CLIMATE
The climate of the studied areas is typically mediterranean with

hot, dry summers and mild wet winters. There are of course dif-
ferences between the climates of the studies areas, but we do not
have data on them,
HATZlKAKIDES (I952A) gives for Mesolonghi a table with meteo-
rological data obtained from several sources. We give this same
table in a somewhat changed form (table I.). Frost is very rare
around Mesolonghi (pers. comm. Mr. T. MIEDEMA).
GEOLOGY AND GENESIS
The Pindos range and some neighbouring chains form a large

mountain chain in the western half of the Greek continent. To the
south this mountain range is cut off by the Gulf of Corinth, to the
West it slopes down to the Ionian Sea. On several places West and
HALOPHILOUS VEGETATION OF MESOLONGHI IOI
TABLE I
Meteorological data from Mesolonghi, derived from HATZmAKm~S I952a.

Rain observations from the period 1894--1929, temperature observations from
19OO--I 929.
Mean number of days Mean temperature of the

Month with rain air in centigrades
January 12.6 9.9

February 12. I 10.6
March 10.9 19.8
April 9.2 16.2
May 8.0 20.2
June 4.6 24. I
July 1.2 26.9
August 1.9 27- I
September 4.6 24. i
October 9.9 19.5
November 12.2 I 4" 9
December 14.2 I 1.5
South of the Pindos and its parallel chains narrow strips of alluvial
country occur between the mountains and the sea. One of these
strips, along the South coast, is the area of Mesolonghi, which
continues in a northerly direction along the West coast towards
Astakos. In the East of the plain of Mesolonghi we find the delta of
the river Evinos and across the river the mountain Varassova. In
the extreme West of the area the river AcheloSs forms its delta.
Most of the plain is formed by the sediments of these two rivers.
The Evinos has left deposits covering the whole area from Meso-
longhi eastward; the Achelo6s from Mesolonghi to the West. In the
western part of the area the sediments of the Achelo6s surround
several rocks and hills, of which the Koutsi]aris (413 m) is the highest.
All these hills consist of Dachstein- or Pantokratorlime (RENZ i955).
In the western part of the area several former beds of the Ache-
lo6s ma y be distinguished (fig. Ia). From archaeological data it be-
comes clear that the front of the Achelo6s delta reached the hills Of
Trikardos with the city of Oiniadai in the fifth century B. C. In the
Middle Ages the city of Oiniadai was abandoned; apparently be-
cause sediments deposited around the hills prevented vessels from
reaching the city (PowELL I904).
In some places in the area of the lagoons former dune ridges may
be observed: near Aghia Triada, near Aghios Taxyarchis and near
the Megalo Vouno. We assume that these dune ridges each ac-
compagnied an ancient mouth of the Achelo6s.
1o2 W.J. WOLFF
SOIL
Except for some general data only little is known of the soil of the
salt-marshes near Mesolonghi. Most information was obtained
from the reports of the Grontmij Ltd. Nearly the whole area
consists of clay deposited in salt water; only along the riverbeds
do we find clay deposited in fresh water. The saltwater clay has
a blueish colour; freshwater clay is brownish.
It is scarcely necessary to mention that the coastal ridge consists
of sand.
Analysis of clay from the bottom of the lagoons gives:
particles <2/~ 25--45 %, mostly between 30--35 %;
CaCQ lO--3o ~ mostly between 15--25 %
(shells) ;
organic matter 4--8 %;
pH varying between (7.6) 8.o and 8. 7.
Near the coastal ridge the percentage of particles < 2 # and the per-
centage of organic matter become less. The percentages of CaCO3
and organic matter also become small in river deposits.
Nearly all vegetations described in the following sections were
observed growing on the same kind of clay. Important factors gov-
erning the pattern of vegetation seem to be the content of NaC1,
the depth of the reduced layers, the depth of the groundwater table,
the draining of the soil, the amount of influx of fresh water and
very complicated complexes of these factors.
The vegetations on drift material have different substrates: plant-
remains, shells, etcetera.
HYDROGRAPHY
The lagoons of Mesolonghi are connected with the sea by means
of several wide openings in the coastal ridge. Nevertheless some parts
of the lagoons are rather isolated. This results in large fluctuations
in the salinity in the course of the year. In the eastern lagoon the
chlorinity varies according to HATZlKAKIDES (I952A) from 7--
9 ~ Cl' in winter to 25--5 ~ ~ CI' in summer. The salinity in the
shallow part South of Aitolikon and probably near the former mon-
astery Aghios Taxyarchis also fluctuates considerably. It seems
reasonable to assume that in general a large fluctuation in salinity
is connected with a large fluctuation of the temperature of the
water. The mean temperature in winter seems to be about IO~ C;
in summer it may well rise to over 3o~
The lagoons are shallow; the maximum depth is about 2 m, but
large areas are about I m . From the shores to the middle of the la-
goon the bottom very gently slopes down. Only the lake of Alto-
HALOPHILOUS VEGETATION OF MESOLONGHI lO3
likon (North of Aitolikon) differs considerably from the rest (HAT-

ZIKAKIDES I952B ). Its depth is at m a x i m u m 28 m and therefore cir-
cumstances are rather constant in this large body of water. It also
buffers the conditions in the lagoon South of Aitolikon, which is
about 1.5 m deep.
The high salinity in summer caused by evaporation is tempered
to a small degree by fresh water flowing into the lagoons from the
rice-fields and from some small brooks.
I n winter, however, it seems that large amounts of fresh water
are drained into the lagoons. It also seems probable that sometimes
in winter water from the rivers Evinos and Achelo6s flows into the
lagoons.
According to Mr. T. MIED~MA (pers. comm.) the tidal difference
is about 15 cm, but storms m a y cause much greater differences to
about I m.
BIOTIC FACTORS
Influences of plants
The study of the mutual relations between plants is one of the
major tasks of phytocenology. Notwithstanding this only little can
be said on this subject, because a study such as ours is only a star-
tingpoint for research into such problems.
We only want to mention an instance of parasitism. A few times
we encountered in the S a g i n e t a l i a m a r i t i m a e - v e g e t a t i o n s
a serious infection of Cuscuta spec. However, it does not seem that
this infection may change greatly the composition of vegetation.
Influences of animals
We observed only two instances of possible influences of animals.
i. O n several places in the lagoons fairly large colonies of breeding
birds occur (Sterna hirundo, S. albifrons, Gelochdidon nilotica, Glareola
pratincola). These colonies were all found in the communities of
Haloenemum strobilaceum, of Arthrocnemum glaucum and of Matthiola
tricuspidata and Beta maritima. In no instance we could observe any
effect on the pattern or composition of the vegetation.
2. Grazing of sheep and cattle only occurs in some vegetation types,
viz. in those of Cyperus distachyus, the S a g i n e t e a m a r i t i m a e a n d
the J u n c e t e a m a r i t i m i . These vegetations often are strongly in-
fluenced. Vegetation can be very low and only remains undisturbed
under the protection of the prickly Juncus acutus. Probably the com-
position of vegetation may change greatly by grazing but we have
no data on this subject.
Io4 W . J . WOLFF
The vegetations of the L i m o n i e t a l i a , the S a l i c o r n i e t a l i a

f r u t i c o s a e and the C a k i l e t e a m a r i t i m a e are generally left
undisturbed.
Human influences.
The principal occupation of m a n in the lagoons is fishing. To
this purpose elaborate pile-screens are constructed in m a n y creeks
with the stems of the reed Arundo donax. In some places large quanti-
ties of this material are washed ashore and there they can be a sub-
strate for a special type of vegetation ( S a l i c o r n i e t u m f r u t i c o s ae
h a l i m i o n e t o s u m ; page II2). O n some places in the area dams
are made to stimulate accretion of soil. O n their influence we have
no data. In general it can be said that the salt-marshes are not often
visited by man and therefore h u m a n influence is only slight.
Influx of fresh water from the rice-fields is already mentioned
on page IO3.
VEGETATION
DESCRIPTION AND CLASSIFICATIONOF THE VEGETATION
Zosteretea (PIGNATTI I953) DEN HARTIG @g SEGAL I964

Zosteretalia (BI~GUINOT I94I ) DEN HARTOG & SEGAL I964
Posidonion oceanici (BR.-BL. 193 I)
Posidonietum oceanici (BR.-BL.I952 ) em.
From the large amounts of plants of Posidonia oceanicum washed
ashore we assume that the association is common in the Gulf of
Patras. We observed large walls on the beach near Patrai and near
Mesolonghi. Our conception of the P o s i d o n i e t u m o c e a n i c i
is considerably stricter than that of BRAUN-BLANQUET because we
exclude the algae from this association in accordance with DEN
t-IARTOG • SEGAL (I964).
Zosterion (CHRISTL~NSEN I934) DEN HARTOG & SEGAL

I964
Zosteretum marinae (BORGESEN I905) DEN HARTOC & SEGAL I964
Offshore near Nea Kallikrateia (Chalkidike) extensive meadows
of the eel-grass Zostera marina, accompagnied by various algae, may
be observed. The bottom is covered with sand; the depth varies
from o. 5 to 3 m or more. We observed the same association near
Salonica, near Katerini, near Corinth and in the Ambracian Gulf
near Arta (outside the lagoons). Probably this association may be
found everywhere on suitable places along the Greek coasts.
HALOPHILOUS VEGETATION OF MESOLONGHI Io5
Ruppietea (J. T/dXEN I960 ) DEN HARTOG & SEGAL I964

Ruppietalia (J. T/JXEN I960) DEN HARTOG & SEGAL I964
Ruppion maritimae ( B R . ~ B L . I93 I) DEN HARTOG & SEGAL
I964
Ruppietum spiralis IVERSEN 1936

In the lagoons of Arta as well as in the lagoons of Mesolonghi
there occur vast and dense vegetations ofRuppia maritima ssp.
spiralis. Near Mesolonghi we observed once between Ruppia some
plants of Cymodocea nodosa washed ashore, but this species was never
found in the lagoons themselves. Other phanerogams were not
encountered.
The bottom in the R u p p i e t u m s p i r a l i s is m u d d y with m a n y
remains of shells; the depth of the water varies from about 5 cm at
low water until at least I m. Where this vegetation occurs, wave-
action and currents are only weak. The community was also ob-
served in ponds blocked from the lagoon. In one of these ponds
the chlorinitywas 37.5 ~ CI'. In the same kind of ponds without
vegetation, possibly drying out towards the end of the summer, we
measured 57.x and 62.7 ~ CI'. Also in the lagoons themselves the
salinity fluctuates strongly.
According to DEN HARTOG & SEGAL (1964) and SEGAL (I 965) a
large salinity range, combined with weak wave-action and current,
is favourable for the occurrence of the R u p p i e t u m s p i r ali s. The
data from the lagoons of Arta and from Mesolonghi wholly confirm
their hypothesis. We followed DEN HAP.TOG & SEGAL (1.C.) in
keeping separate the R u p p i e t u m from the algal communities.
The R u p p i e t u m s p i r a l i s is known from large parts of Europe.
Potametea Tx. & PRSO. 1942

Parvopotametalia DEN HARTOG & SEGAL 1964
Callitricho-Batrachion DEN HARTOG & SEGAL I964
Ranunculeturn baudotii BI~.-BL. 1952 Table 2 : 1 - 1 3 .
In ditches in the vicinity of the village Magoula near NIesolonghi
we observed in the cultivated area with fresh or oligohaline water an
aquatic vegetation of rather poor floristic composition, in which
Zannichellia palustris ssp. pedicdlata dominated. Also a poor-flowering
form of Ranunculus trichophy[lus (R. paucistamineus TscH.) regularly
occurred. We assume that this vegetation will prove to be the
same as the R a n u n c u l e t u m b a u d o t i i described from southern
France, although this association was based on only one vegetation
record. SEGAL (1965) includes the R a n u n c u l e t u m baudotii
BR.-BL. 1952 in the C a l l i t r i c h o - R a n u n c u l e t u m baudotii
DEN HARTOG I963.
io6 W . J . WOLFF
TABLE 2.
Nr. I u 3 4 5 6 7 8 9
Date 19-IV I9-IV 92-IV 94-IV 22-IV 92-IV 24-IV ~9-IV ~2-I ~
'63 '63 '63 '63 '63 '63 '63 '63 '63
Area M M 1V~ M 1V[ M M M M
Surface (m s) 4 4 9 19 9 4 7 400 io
Cover of plants emerging above the water (%) -- -- 5% 50% Io% .~% 1o% 9o
H e i g h t of plants emerging above the w a t e r (era) - - - - 1oo ? ? ? I00 70
Characteristic species combination

Scirpetam maritimi et litoralis:
Seirpus marltimus ~ -- -- -- () -- 1.9 2m.5 z.9
Sclrpus lltoralis -- -- -- +.9 -- -- I.~ 9a.3
T y p h a cf. angustata -- -- -- I'~ 3.3 1.5 +.I +.2 +.I
Characleriatic species Phragmitetalia and Phragmitetea:
Phragmites communis -- -- . . . . . . r
Allsma cA'.plantago-aquatica -- --
Companion:
S a l i c o m i a frutieosa -- --
Cover of submerged plants ( % ) : 5o % xoo % so% ioo% ~o% xo% 2o% 5o% 4oC
Depth o f the water (cm) 5 o 0 5 IO ? 0----I5 5--I0 IO
Characteristic species combination Ranunadetum baudotii:

Callitrlche t r u n e a t a
Oallitriehe spcc.
Zannichellia palustris ssp. pcdicellata 3.4 5.5 3.3 5.5 2a.3 L9 2a. 3 3.5 3.5
Ranunculus trichophyllus . . . . . . . . 9a.3
Characteristic species of the Parvopotamion:

Potamogcton crlspus . . . . . . . .
Potamogeton el, pusillus . . . . . . . .
Potamogeton natans . . . . . . . .
Companion:
C h a r a spee. -- . . . . . ~a.3 x.9
O c c u r r i n g in record 16: Chlorophyceae + ; 2 i : Juncus maritiraus + . 9 , Aeluropus litoralis + . 2 , Salleornia herbacea s.1. + , t ; 99:Rupl0
sp. t . 9 ; 93: Juncus acutus x.2.
M ~ lagoons of Mesolonghi.
Well developed vegetations occur in stagnant or slowly running

water, io to 50 cm deep on a clay or mudbottom. In very shallow
ditches which desiccate in summer, the association is reduced to a
facies of Zannichellia palustris ssp. pedicellata. In a single case the
chlorinity was measured: 1. 3 ~ o CI'. The R a n u n c u l e t u m b a u -
d o t i i is only mentioned before from France. The C a l l i t r i c h o -
R a n u n c u l e t u m b a u d o t i i DEN HARTOG 1963 was described from
the Netherlands.
Although not belonging to the halophilous communities, we
would like to mention here two types of aquatic vegetation from
Mesolonghi:
a. vegetation ofPotamogeton crispus and P. cf. pusillus (table 2: 14-19) ;
b. vegetation of Potamogeton natans (table 2: 2o).
The former community occurs, generally in low densities, in shallow
HALOPHILOUS VEGETATION OF MESOLONGHI lO7
IO II 12 13 I4 t5 I6 I7 18 I9 20 21 22 23 24
23-IV ~3-IV 23-IV a3-IV 24-IV ~9-IV ~4-IV 2I-IV 24-IV ~4-IV 28-IV I3-VI 29-IV x-V I-V
'63 '63 '63 '63 '63 '63 '63 '63 '63 '63 '63 '63 '63 '63 '63
M M M M M M M M M M M M M M M
8 2 5 4 20 9 6 15 IO IO 150 IO0 IO0 IO0 IO0
I0% I0% 20% 5--I0% I0% 20% ? 50% 5% I00% I00%
I50 IO0 I00 50 ~ 200 ~ I00 30 IO0 IO0
-- -- 2m.3--4 2a,2 +.t 1.2 • -- +.~ I°-2 ~m.5 5"5 5-5

H
-- 1,2 2 m . 3 - - 4 1.2 1.2 -- +.2 +.3 3.5 +.2
-- 2a.3 1.2 2a.2 2a.2 1.2 1.2 1.2
1.3 . . . . 2a.3 --
+.2 -- -- +.2 -- -- --
I°.5 I~.5
xoo% 95% xoo% ~oo% 5o% 3o% 5% 45% 75% 2o% ? . . . .

5° 30 3° 4° 5--15 I0--20 I0--20 I0 I0---20 20--30 30 O IO--I 5 0 0
+.2 - - 2m. 5 -- --
- - 3 . 4 - - + . 2 - -
5.5 1.2 1.5 -- +.2

- - 4-4 5-5 1.2 - -
3.5 -- -- 2b.3 4.5 2a'3 ] . . . . .

d __ - -
+.z 3.3 2a.2 3.3 --
1,2 +.o -- 5,5 I ~-.I
ditches (lO--3o cm) with in summer slowly running fresh water

originating from the rice-fields. It is possible that in spring and
a u t u m n a weak influence of NaG1 may be felt. The latter vegetation
was observed in a ditch filled with fresh water and about I m deep.
Any influence of sodium chloride must be excluded here. The syn-
taxonomic position of these vegetations remains doubtful.
Thero-Salicornietea (Tx. 1954) Tx. & OBERD. 1958

Thero-Salicornietalia (Tx. 1954) Tx. & OBERD. 1958
Thero-Salicornion (BR.-BL. 1931 ) Tx. 195o
Salicornietum europaeae (WARMING19o6 ) BEEFTINK 1962. TABLE 3"
In places where fresh water from the rice-fields flows into the la-
goons and in some places in the bare zone between the Halocnemura
~o8 W.J. WOLFF
and the Ruppia-vegetations a vegetation occurs consisting nearly ex-

clusively of Salicornia herbacea s.1. Since this taxon could not be in-
dentified more exactly, it seems best to include these vegetations
into the association group of the S a l i c o r n i e t u m europaeae
sensu BI~EFTINK 1962. Certainly this association can be defined in a
better way because the Salicornias clearly belong to one or perhaps
two types. The S a l i c o r n i e t u m h e r b a c e a e VAI~ LANGENDONCK
1933 as distinguished by PIGNATTI (1953) may provisionally be
included in this association group.
TABLE 3"
Hr. 25 26 ~7 28 29 30 3~ 32 33 34 35 36 37
Date zo--Vl 2--VI 2--VI zz--V zz--V ~-VI 2--VI il~V zl--V 2o--IV z3--VI 24--1V24--1V
'63 '64 '64 '63 '63 '64 '64 '63 '63 '63 '63 '63 '63
Area M A A M M M A M M M M M M
S u r f a c e ( m s) ZOO ZOO ZOO 1OO ZOO ZOO ZOO ZOO ZOO 400 100 400 4OO
Cover (%) 0. 5 Io 5~ 20 20 3o 5~ 20 2o 5 z5 2 5
H e i g h t (era) zo 9. ? 3~ 3~ ? ? 25 25 20 20 I0---15 I0-'-20
Characteristic species
Salieornietum europaeae:
S a l i c o m i a h e r b a c e a ssp. i,z 2m, z 3--4,5 em.2 2m.5 3.5 5--4.5 2m.~ 2m.2 2m.5 2m.5 2m.5 2m.5
Characteristic spedes
Salicornion fruticosae:
Salleornia fruticosa . . . . . 1.2 +.2 - - - - - - z.2 - - +.t
Salieornia radicans . . . . . . +.2 +.z +.z +.z +.2 +.z +.z
Puccinellia festuciformis -- -- -- z. z +, I - - -- 4.2 . . . . .
Aeluropus litoralis . . . . z.3 + .2 - - +.I Z.I +.Z - - - - - -
Companions:
Haloenemum strobilaceum . . . . . .
. . . . +.i +.z +.z
Scirpus marltimus -- -- -- t~ z~ -- -- i~ z~ z,i - - - -
Scirpus litoralis -- -- - - I~ z~ 4~ - - z O . z Z~ -- _ _ - - +~
Juncus acutus . . . . . . - - - - +.z(?) +.2 - - - - - -
Cyanophgceae 5.5 . . . . 5.5 . . . . 5.5 -- --
Occurrin3 in record 3I: Halimione portulacoides 4.2, S t a t i c e l i m o n i u m ssp. s e r o t i n u m 4 . 2 ; 32: J u n c u s m a r i t i m u s 4.2; 35: Cy!oerus
d i s t a c h y u s (?) 4 . 2 .
M = l a g o o n s o f M e s o l o n g h i ; A = l a g o o n s of A r t a .
The soil consisted in all cases of clay and was always very wet.
Often it was covered with a thin layer of water; it also smelled of
H2S.
The association occurs in those areas which are subjected to
changes in two sets of factors: waterlevel and salinity. The impor-
tance of these factors lies not in any specific magnitude but in
the instability of this magnitude.
The association and close related ones in the Mediterranean are
also known from Venice (PIGNATTII953) , Sicily (FREI 1937) and
Macedonia (OBF,RDORFER 1952). The relations with the Salicornia-
communities of north-west Europe and central Europe are not clear.
HALOPHILOUS VEGETATION OF MESOLONGHI Io9
Salicornietea fruticosae (BR.-BL. & Tx. 1943) Tx. & OBERD. I958
Salicornietalia fruticosae (BR.-BL. 1931) Tx. & OBERD. I958
Salicornion fruficosae BR.-BL. 193 I
Syn.: Halo-Puccinellion PIGNATTI 1953 p.p. (suballiances Boreo-
Puccinellion PIGNATTI 1953 and Puccinellion mediterraneum
PIGNATTI 1953 p.p.)
Salicornietum radicantis BIn-BE. 193 I. TABLE 4.
The S a l i c o r n i e t n m r a d i c a n t i s occurs near Mesolonghi on
well-aerated clay and sandy clay soils. The soil is reduced at a depth
of lO--2o cm and always somewhat wet. The toplayer of the soil
may be desiccated in some instances. The description of the milieu
by BEEFTINK (1965) from Brittany and Normandy is in accordance
with our data from Greece: a rather high salinity, well-aerated and
fairly wet. The data of BRAUN-BLANQUETC.S. (1952) therefore are
difficult to explain. In southern France the association should be
situated on the lowest places a]ong the shores of the lagoons @tangs).
Perhaps it may be explained by assuming that the situation of this
association on the edge of the water is responsible for a good drain-
age which induces aeration of the soil. On the other hand the
proximity of the water will cause these places to be wet and saline.
Near Mesolonghi two variants may be distinguished: one poor
in species and another rich. The poor variant is characterized by
the occurrence of Salicorniafruticosa; the rich one by a large number
of predominantly annual L i m o n i e t a l i a and J u n c e t a l i a m a r i -
t i m i species. We suppose that the difference between these two
variants may be found in the level of the soil water, but we do not
have enough data on it. The rich form is always encountered in
well-drained places, e.g. the steep shores of little ponds. This drier
and richer variant forms a transition between the L i m o n i e t a l i a ,
t h e J u n c e t a l i a m a r i t i m i and the S a l i c o r n i e t a l i a f r u t i c o s a e .
OBERDORFER (1952) mentions this association (two records: one
of each variant) from the Axios-delta near Salonica. We ourselves
met with it near Arta and near Mesolonghi. According to BEEFTINK
(1.C.) the association is known from Wales and southern England,
the atlantic coast of France, Spain and Portugal, and in the Medi-
terranean from France, Sicily, Venice and Macedonia. In all these
instances only the poor variant is recorded; the rich one is only
known from Greece, but will without doubt be found in other med-
iterranean countries.
Salicornietumfruticosae BR.-BL. 1928. TABLE 5"
Around the lagoons of Mesolonghi extensive vegetations of Sa-
licornia fruticosa occur. Often a variant very poor in species is met
with; only Salicorniafruticosa forms a fairly high and very dense ve-
1 io W.J. WOLFF
TABLE 4"
Nr. 38 • 39 40 4x 4~ 43 44 45 46 47
Date 3-VI 28-V 3o-V 5-VI 28-V 28-V 28-V 27-V 27-V 3o-V
'63 '63 '64 '64 '63 '63 '63 '63 '63 '64
Area M M A M M M M M M A
S u r f a c e ( m 2) 5 49 5 50 5o 49 Ioo 5o 5o 3o
Gover (%) x5 50 75 50 3o 3° 50 4° 40 90
H e i g h t (era) 2o Io ? ? 2o 2o IO 30 40 ?
Charaeterlsti¢ species Salieornietum radicantis:

S a l i c o r n i a radicmas 2b,3 3.5 4.5 3.5 2b.2 3.5 3.5 2b.3 ~b.3 4.5
Differential species of the wet variant:
Salicornia fruticosa . . . . x.2 +.2 i.~ 2b.3 2b.3 +oj
Differential species of the dry variant:

Plantago eoronopus
Spergularia salina
Juncus subulatus m
Hordeum marlnum m
Juneus ambiguus
Centaurium spicatum m u
Characteristic species of the Limonietalia:

Arthrocnemum glaucum m + . I I.I - -
Haloenemum strobilaceum
Parapholis strigosa ~-.I -- -- -- 2b.3
Parapholis incurva
Characteristic species of the Salieomietalia frutieosae:
Puceinellla coavoluta +.2 I. 2 -- -~-.o _ _ _ _ i°.2 - - +.I +.o
Statice limonium ssp. serotinum -- -- +°.x +,2 -- -- 2a.2
Halimioae portulacoides - - - - - - + . !
I°.2 +°.2 --
Aeluropus litoralis - - - - + . 3 1.21UX -- -- 2m.5
Companions
S a l i c o r n i a h e r b a e e a s.1. + , I I.I I.I - - 2b.2 I.I I.! I.l I.I +.I
Juncus acutus __ 1 --
Phragmites cornmunis +°.t +°.I +.I
O c c u r r i n g in r e c o r d 4 o : S u a e d a m a r i t i m a + ° d ; 4 7 : J u n c u s m a r i t l m u s + . 2 ; 4 9 : T r i g l o c h i n b u l b o s u m i o i ; 5 0 : P u c c i n e l l i a f e s t u c i f o r m i s
1.2; 5 I : P o I y p o g o n m a r l t i m u s + . 2 ; 52: A t r i p l e x h a s t a t a r . x , S u a e d a m a r i t i m a + ° . x ; S o n c h u s sp. r; 5 3 : T r i g l o c h i n b u l b o s u m + . a , I n u l a
c r i t h m o i d e s + . 2 ; 5 7 : S c i r p u s e e r n u u s + . x ; 6 5 : P o l y p o g o u m a r i t i m u s 1.2.
i = l a g o o n s of M e s o l o n g h i ; A = l a g o o n s of A r e a .
TABLE 5"
Nr. 68 69 7° 71 72 73 74 75 76 77 78 79
Date xz-IV iz-IV ~7-V 27-V Is-VII 6-VI 7-V 6-VI x5-VI x-V 6-VI 6-VI
'63 '63 '63 '63 '64 '64 '63 '64 '64 '63 '64 '64
Area M M M M Th M M M Th M M M
S u r f a c e ( m 2) too I2 50 I0o too Io0 Ioo 50 Ioo 400 Ioo Ioo
Cover (%) 95 ioo 60 70 50 IOO IOO 90 30 IOO 75 7°
Height 4o 3° 30 3° ? ? 4° ? ? 40--5 ° ~. ?
Characteristic species combination

Salieornietum fruticosae:
Puceinellia fesmciformis -- +.I -- -- -- r.~ +.2 . 1.2 -- 1.2 -- +.o
Salicoruia fruticosa 5-5 5.5 4.5 4.5 3--4°.5 5.5 5.5 5.5 3-5 5-5 4-5 4,5
Differential species of the subass.
halimionetosum :
Salicornia radicans . . . . r° . . . . . . .
Suaeda maritima . . . . . . . . +. I -- -- --
Preferential species of the subass.
halimionetosum :
Halimione por tulacoides +.I -- -- -- 1.2 -- t.2 +.~ +.x +.2 +.2 +.2
Diff. species of the subass.
halimionetosum in Macedonia:
L i m o n i u m cf. g m e l i n i . . . . . . . . . . . .
Species of the Salicornletalia
frutieosae:
Juncus subulatus . . . . . . . . . +.3 --
S t a t i c e l i m o n i u m ssp. s e r o t i n u m : . . . . +.2 -- -- -- 1.2 -- --
Companions:
Phragmites communis -- -- t°.2 x°.~ . . . . . . . .
S a l i c o r n i ~ h e r b a c e a s.l. -- +°.x +°.l . . . . l.t L3 2re.x--2 L2
O c c u r r i n g i n r e c o r d 78: P u c c i n e I l i a c o n v o l u t a t . ~ ; 79: t M c c i n e l l i a c o n v o l u t a + . 2 , A r t h r o c n e m u m g l a u c u m + . 2 ; 82: S c i r p u s m a r l t i m u s

-¢-.I, J u n c u s a c u t u s 2 m . 5 ; 8 7 : A r t h r o c u c m t t m g l a u c u m + . 2 ; 88: A t r i p l e x h a s t a t a L 2 ; 9o: T r i g l o c h i n b u l b o s u m 1.2, I n u l a c r l t h m o i d e s
+ . 2 , T a m a r i x sp. 1.2; 93: A s t e r t r i p o l l u m 1.2; 9 4 : A s t e r t r l p o l i n m + , z.
M = l a g o o n s o f M e s o l o n g h l ; T h = d e l t a of I~. A x i o s n e a r S a l o n i c a .
HALOPHILOUS VEGETATION OF MESOLONGHI Il I
48 49 50 51 52 53 54 55 56 57 58 59 60 61 62
3-VI 7-VI 7-VI 7-VI 3o-V 13-VI 27-IV 27-IV 3-VI 27-IV 27-IV 29-IV 29~V 3-VI 3-VI
'63 '64 '64 '63 '64 '63 '63 '63 '63 '63 '63 '63 '63 '63 '63
M M M M A M M M M M M IV[ M M N£
7 1o 4° too 5 4 2 2 IO 4 IO 8 8 zo 20
I5 75--80 1oo 40 go 90 50 50 20 40 50 60 45 60 70
25 ? ? 1o(30) ? I0 I0----20 ? I5 to'--20 25 I5 I5 X5 20
2b.3 4.5 3.4 3.3 22.3 3.5 2b.3 2b.3 2b.3 2b.3 2b.2 2b.3 3.3 3.3 4.5
+.2 1.2 3"3 1.2 +o.i +.2 . . . . . . . . .
-- -- x.t -- +.x x.t +.1 2a,2 1.I +.I +.~,

-- -- I.I +.I I.l I,i t,I -- +.°I +,2 +°o
+°.2 i,zt -- -- I,I -- 1.2 ':'m.5 -- 2m. 4 2m. 5 1.2 1.2 1,3 1.2
1,2 ~m,~ +,I -- +,I -- -- 1,2 I,~ I,I l,I
- - I,2 1.2 - - - - I , I - - I,I I,I - - - -
- - - - - - r -- +.i 1.2 . . . . . . +.i --
+.2 -- +.2 +.1 ~ -- +.t +.t +.x +,2 +.i -- -- +.1 --

- - - - - - I.I . . . . r -- + . I . . . .
-- +.2 1.2 +.2 2a.3 2m.5 +.I -- +.I I.I +.2 1.2 1.2 +,I +.2
-- -- +.2 . . . . . . . . . . .
+.2 22.2 22.2 1.2 -- 1.2 2b.3 2b.3 1.2 2b°.2 +°.2 2b.3 22.2 2b. 3 22.3
-- +.2 1.2 +.2 2b.3 2b.3 +.I -- +.2 -- -- 1.2 -- +.2 +.2
+Q.2 1,2 3.3 +,2 +,2 +,2 +,2 -- -- +.2 -- 22,2 1.2 i°.3 l°.2
1,2 +.2 +.2 1.2 -- +,2 +,2 -- I,I -- -- 22.2 +.2 +.2 1.2
+,i -- -- i,i +.I -- -- -- +.x . . . . +.x --
-- -- -- +.2 2a.3 +.2 --

-- r -- +.2 -- --
80 81 8° 83 84 85 86 87 88 89 90 91 92 93 94 95
t-V 2-V x-V 2-V x6-IV x-V x-V 6-VI x-V x-V ~,-V 2-V ~,-V x5-VI I5-VI t5-VI
'63 '63 '63 '63 '63 '63 '63 '64 '63 '63 '63 '63 '63 '64 '64 '64
M M M M M M M M M 1V[ M M M Th Th Th
400 30 IO0 5° 30 400 I2 IO0 40 20 3° 30 3° 20 3° I0
Ioo 80 IOO 90 95 xoo xoo xoo 70 Ioo Ioo IOO 95 too xoo 95
50 4° 5° 50 50 5° 50 - 30 4° 23 (4 o) 25 ~5 (4 o) ? ? -
+.2 2a,2 -- +.i 1.2 1.2 -- 22.2 +.2 2a.3 4.4 2a.3 22.3 -- -- --
5.5 5.5 3-5 5.5 4,5 2b.3 3-4 1,2 2b.3 2b.3 22.2 r.2 t.2 3--4.5 3.3 4.5
-- +.~ -- +.x 1.2 2a.3 3.4 -- x.~ --

-- -- 2b.2 +,2 -- -- -- 2~ °-2 I°,2--3 I ~.2
i.~ t.a 3.5 1.2 ~b.3 +.~ 4.5 5-5 3.3 3-3 3.3 4.5 4.4 3.3 3.3 ~a.2
. . . . . . . . . . . . . 2a.2 2b.3 i2b.3
-- -- +.2 -- -- +.2 . . . . . . . . . .
+.2 +.2 +.~ +.2 -- +.x -- -- +.~ i.o :t.2 z,~ 1.2 z.2 +.2 x.2
-- -- +.r . . . . . . r.~ -- +.2 . . . .

. . . . +.x -- -- __ +.~ . . . . . +.x --
II2 W.J. WOLFF
TABLE 4 (Cont.)
Nr~ 63 64 65 66 67
Date 3-VI $-VI ~-VI ~-VI 28-V
'63 '63 '64 '64 '63
Area M M A A M
Surface (m2) 2o 6 20 IO0 49
Cover (%) 20 5~ 50 4~ 30
Height (era) I5 20 ? ? 15
Characteristic species Sallcornietum radicantls:

Salieornia radieans 2b.3 3.5 3.3 3.3 3.5
Differential species of the wet variant:
Salicornia fruticosa
Differential spedes of the dry variant:
Plantago eoronopus -- I,I I,I I.I --
Spergularia salina 1.2 -- I,I I.I I,I
Juneus subulatus I.I 1.2 -- -- --
Hordeum marinum -- I.I 1.2 1,2 --
Juneus ambiguus
Centaurium spieatura
Characteristic spedes of the Limonietalia:
Arthroenemum glaucum @.I . . . .
ttaloeaernum strobilaeeum
Parapholls strlgosa
Parapholis ineurva -- -- I.I ~-.I --
Characteristic species of the Salicornietalia fruticosae:

Pueelnellia eonvoluta 1.2 1.2 -- -- 1.2
Statice limonium ssp. serotinum +.2 +.2 1.2 +.2 +~
Hallmione portulacoides ~~ I~ . . . .
Aeluropus litoralis 1,2 I,I 1,2 ~*I --
Companions:
Salieornia herbacea s.I. +,I JF.I l,I -- I.I
Juncus aeutus
Phragmites eommunis
getation in which only a few individuals of Puccinelliafestuciformis,

Limonium vulgare ssp. serotinum and Halimione portulacoides occur. The
soil is completely reduced and smells of H2S; the toplayer is often
covered with Cyanophyceae. Moreover the soil is covered with
a thin layer of water, probably all the year round. This type of
vegetation we want to distinghuish as the subassociation t y p i c u m
subass, nov. Compared with the next type we may regard this sub-
association as a back-swamp vegetation. Where the S a li c o r ni e-
turn f r u t i c o s a e is bounded by the open lagoon, sometimes a
natural levee is formed, especially by remains of the reed Arundo
donax (see page IO4). These levees are characterized by the strong
dominance of Halimioneportulacoides and the occurrence of Salicornia
radicans and Suaeda maritima. The physiognomy of this vegetation
reminds one of the atlantic H a l i m i o n e t u m p o r t u l a c o i d i s , but
its floristic composition is quite different. We want to distinguish
these vegetations as S a l i c o r n i e t u m f r u t i c o s a e h a l i m i o n e t o -
s u m subass, nov. The difference with the milieu of the S a l i c o r -
n i e t u m r a d i c a n t i s is not always quite clear; perhaps the soil is
better aerated in the subassociation described here. The descrip-
tion of the milieu in the mediterranean area by BEEFTINK (1.C.)
does not seem applicable. We may characterize it by a very good
aeration, a fairly high salinity and a moist substrate. Remarkable
HALOPHILOUS VEGETATION OF MESOLONGHI 1i 3
is the fact that Puccinelliafestuciformis is optimally developed on and

especially just behind the levees with Halimioneportulacoides.
In the delta of the river Axios the levees of very small creeks are
characterized by the occurrence of Limonium cf. gmelini.
The S a l i c o r n i e t u m f r u t i c o s a e is known from the Iberian
peninsula and mediterranean France (BRAUN-~BLANQUETC.S. 1952),
from Sicily (FI~.I I937) , from Venice (PIGNATTI1953) and from
Dalmatia (HoRvATIC 1934). OB~.RDOI~I~ER (1.C.) did not mention
the association from northeastern Greece, but we noticed it near
Salonica and near Porto Lago (Thrace).
Some measurements of the chlorinity of the water-layer just above
the soil gave the values: 17.6 , 16.8, 23.6, 36.4 and 38.1 ~ CI'. In
all these cases these values were about the same or slightly higher
than the value for the water of the neighbouring lagoon. The sam-
ples were taken between 3 and 6 J u n e 1964 .
Limonietalia BR-BL. & DE BOLOS 1957

Syn.: Suball. Limonion ferulacei PIGNATTI 1953 and Puccinellion
mediterraneurn PIGNATTI I953 p.p.
Staticion galloprovincialis BR.-BL. 1931
Salicornio-Halocnemetum strobilacei ass. nov. TABLE 6.
Syn.: Arthrocnemum glaucum-Halocnemum strobilaceum-Ass. OBER-
DORFER I952 p.p.
Large areas of the dry parts of the salt-marshes near Arta, Meso-
longhi and Satonica are "covered" by a monotonous vegetation
of Halocnemum strobilaceum and a very distinct form of Salicornia
herbacea.
Except for these two dominant species only a few species pointing
to a relation with the L i m o n i e t a l i a and probably the S t a t i -
c i o n g a l l o p r o v i n c i a l i s , are seen to occur sparingly in this associ-
ation. Another relation may also be defended, because it is possible
to regard this association as the westernmost radiation of the east-
ern H a l o s t a c h y e t a l i a into the northern mediterranean area.
But as long as this order is so imperfectly known, it seems better to
place this association into the L i m o n i e t a l i a . Moreover several
species occurring in the association also occur in the other communi-
ties of the L i m o n i e t a l i a .
In the environments of Mesolonghi the boundary between the
S a l i c o r n i o - H a l o c n e m e t u m s t r o b i l a c e i , as we propose to call
this association, and the A r t h r o c n e m e t u m Bm-BL. 1928 is very
sharp. The dominating species Halocnemum strobilaceum and Arthroc-
nemum glaucum are mutually exclusive. The A r t h r o c n e m e t u m is
situated on higher levels than the S a l i c o r n i o - H a l o c n e m e t u m
s t r o b i l a c e i (see fig. 2). Several species occurring in the A r t h r o c -
114 W . J . WOLFF
TABLE 6.
Nr. 96 97 98 99 ioo IOi IO2 IO3 Io4 zo5 xo6 lO7

Date 2-VI II-IV ix-IV 2-VI ~-VI 2-Vt ~-VI I5-VI I5-VI IS-VI ts-VI I5-VI
'64 163 '63 '64 '64 '64 '64 '64 '64 '64 '64 '64
Area A M M A A A A Th Tit Th Th Th
Surface (1212) ioo ioo I00 IOO I00 I0o IO0 35 IO0 IOO IO0 IO0
Cover ( % ) xo 5 5--Io io 3~ 3~ ~,~ 50 50
H e i g h t (era) ? 15 I5 39. . ~ . 5 . io . ~. .~ . 9 . ~
Characteristic species Salicornio-

Halocnemetum strobilacei:
I-/aloeneraum strobiIaceum 2a.2 i. i 9a.2 1.2 2a.2 1.2 2am ~b.3 2b.5 2b.5 3.4--5 3-5
Salicorniaherbaceassp. -- - - - - I .I 2m, I - - 2 2ra. I +.I 2ra.! 2m, 5 i~ I.I +.I
Characteristic species of the
Limonietalia :
Sphenopus divaricatus . . . . . . . . . . . .
Parapholis incurva . . . . . . . . . . . .
Plantago coronopus . . . . . . . . . . . .
Arthrocneraum glaucum . . . . . . . . . . . .
F r a n k e n i a pulverulenta . . . . . . . . . . . .
Spergularia salina . . . . . . . . . . . .
Occurring in record I oo: Salicornia radicans + . 2 ; 103: Suaeda raaritima + . I, Salicornia fmticosa + . i ; 1o4: Salicornla fruticosa + . I
io6: Suaeda raaritiraa r, Hallrainne portulacoides r; tx8: H o r d e a r a raarintma x.2; I2o: Salicornia radicans + . I , Aeluropus lltoralls v
x~6: Parapholis strigosa + . x , H o r d e u m raarinum 1.2; 128: Polypogon raaritiraum L I ; z29: Parapholis strigosa [ . I ; I32: Xanthor~
parietlna on sterns of Arthrocneraura glaucum.
A = lagoons of Arta; M = lagoons of Mesolonghl; T h = delta of R. AMos near Salonlca.
C ARTHROCNENE
TUN
LAGOON' BARE ISALICORNIO- ,IJ SALICORNIE-{'JUNCUS PHRAG

CLAY\I-IALOCNENEj
0,N)TOMFRUTI-;ACUTUS
IHITES
[ETLMCOSAE (+J.MARI-ICOMMU
! ,,:> C';,:J
i/i~' ,'i"i~, :
Fig. 2. A pattern of zonation observed between a limestone hill and the lagoon.
Mesolonghi, May 1963. Scale I : 5oo.
nemetum are lacking in the S a l i c o r n i o - H a l o c n e m e t u m

s t r o b i l a c e i : e.g. Triglochin bulbosum, Parapholis strigosa, Salicornia
radicans a n d Statice limonium ssp. serotinum.
I n the salt-marshes n e a r A r t a a n d n e a r Salonica we o b s e r v e d the
s a m e vegetations. I t m u s t be r e g r e t t e d t h a t we only were able to
HALOPHILOUS VEGETATION OF MESOLONGHI xI 5
I08 I09 IIO III I12 II3 II 4 II 5 II6 II 7 IIS II 9 120 I2I 122 I23
l-IV 19-IV 2 9 - I V 22-IV 2-VI I9-IV 22-IV a9-IV II-IV II-IV 29-IV 29-IV 24-IV 2o-IV 2 r-V 21 - V
'63 '63 '63 '63 '64 '63 '63 '63 '63 '63 '63 '63 '63 '63 '63 '63
lVi M M M A M M M M M M 1'¢I M M M IV[
400 400 IOO 400 Ioo 400 400 IOO too 6o loo zoo 400 ioo ioo ioo
io io 15 Io IO 20 IO xo 40 5o 15 15 5 20 3 IO
Io I5--2o 15 15 ~• 15 IO 15 25 20--25 15 I5 IO 20 tO 20
oa°,2 oa.2 2b.3 2a.3 2a.2 2b.2 2a.~ ~a°.2 3.3 3,3 ~a°.2 ~b°.2 1.2 2b.3 1,2 2a.~
+.¢' 2m, I +.2 1.2 +.I 2rn.I 2m. I--2 2m. 5 2 rrl, I 2rn, i 1,2 2m.5 I.I 2111,1 21n,2 +,I
+.x -- +.~ +.2 1.2 -- -- 2a.~ 1.2 +.I 1.2 I.I l.I
-- +.I +.2 2a.2 1.2 I.I 1.2 --
. . . . . . . . +.2
. . . . . . . l.I ~m. 3
- - +.I I.I 1,2 ~.o +.r +.z -- +.~ J:.i +.:r +.t +.x
discover one instance of an A r t h r o c n e m e t u m near Arta and none

near Salonica. But rearranging table 6 of OBERDORFER (1,C.) we
see a distinction between the A r t h r o c n e m e t u m and the S a l i c o r -
nio-Halocnemetum s t r o b i l a c e i , also in northern Greece.
However the constant occurrence of Salicorniafruticosa in the records
of OBERDORFER proves that there must be slight differences between
the association near Mesolonghi and near Salonica. We ourselves
observed in the latter region vegetations with Salicornia fruticosa
near the shore and vegetations without this species more inland.
The other differences between the vegetation records of OBERDOR-
I~ER and those of ourselves seem to be unimportant. Consequently
we split the association described by OBERDORFER in the A r t h r o c -
n e m e t u m BR.-BL. I928 and the association described here. PIG-
N A T T I (1953) also saw the heterogeneity of OBERDORFER'Sassociation,
but he brought the records discussed here to the H a l o c n e m e t u m
s t r o b i l a c e i (KELLER 1923) T e A 1939. The A r t h r o c n e m e t u m
resulting from this splitting is hardly different from the association
from southern France. Therefore we can only place it in the Sta-
t i c i o n g a l l o p r o v i n c i a l i s . For its accompanying species we
ought to place also the S a l i c o r n i o - H a l o c n e m e t u m s t r o b i l a -
cci into the S t a t i c i o n g a l l o p r o v i n c i a l i s , if we regard this as-
sociation as belonging to the L i m o n i e t a l i a . The Petrosimonia
crassifolia-Bupleurum gracile-Ass. OBERDORFER 1952-) has some species
not occurring in the communities of the S t a t i c i o n g a l l o p r o v i n -
c i a l i s , but these species can be regarded as characteristic for this
association. It has no species in common with other communities
I 16 W, J. WOLFF
TABLE 6. (Cont.)
Nr. I~4 I25 i26 I27 I28 x29 I3 ~ I3I I32
Date ~9-IV xI-IV 29-IV I9-IV ~o-IV o7-IV 2-V I6-IV x6-IV
'63 ~63 '63 '63 '63 '63 '63 '63 '63
Area M M M M M M M M M
Surface (m ~) IOO 400 I oo 400 l oo 400 4oo 4o0 ~25
Cover (%) I5 x5 I0 3~ 20 20 25 5~ 4~
H e i g t h (era) I5 2o I5 20(40) 5(I5) 20 IO 20---30 20
Characteristic species Salieorn:eto-

Halocnemetum strobilacei :
Halocnemum strobilaceum 2b.2 2b.I 2a.2 2b.2 2a.2 2b.5 2b.5 3.3 2b.2
S a l i e o r n i a h e r b a c e a ssp. +. i I. I r 2m.I -- 2m. 5 -]-.2 -- --
Charaeteriaic species of the
Limonietalia:
Sphcnopus divaricatus -- +. I -- ~-.I I.I r -- 1 1
Parapholis incurva x.2 +.2 +.2 +.i ~.x +.I -- -- --

P l a n t a g o coronopus +. I r -~oI -- 1.3 -~.o -- -- --
Arthrocnemum glaueurn -- -- -- 2a.I q-.x 1.2 +.2 I.~ 2b.~
Frankenia pulverulenta -- --
Spergularia salina +.t 2m.i "q-.I r am.I I.~ -- -- --
except those of the S t a t i c i o n g a l l o p r o v i n c i a l i s . We therefore

do not maintain the S t a t i c i o n o r i e n t a l e OBERDORFER I952 but
regard the communities of this alliance as belonging in the S t a t i -
c i o n g a l l o p r o v i n c i a l i s . Moreover the position of the Petrosi-
monia-Bupleurum-association to the S t a t i c i o n g a l l o p r o v i n c i a l i s
is the same as that of the S a l i c o r n i o - H a l o c n e m e t u m stro-
b i l a c ei. Their characteristic species, considered by OBERDORFERas
characteristic for the S t a t i c i o n o r i e n t a l e , exclude one another
as becomes clear from the tables of OBERDORFER.
So the Petrosimonia-Bupleurum-associationhas to be placed in the
S t a t i c i o n g a l l o p r o v i n c i a l i s if one wishes to place the S a l i -
cornio-Halocnemetum s t r o b i l a c e i in the L i m o n i e t a l i a .
However, if one wishes to classify the S a l i c o r n i o - H a l o c n e m e -
t u r n s t r o b i l a c e i in the H a l o s t a c h y e t a l i a , it seems possible
to do the same with the Petrosimonia-BupIeurum-association, but it is
perhaps better to maintain in that case a separate alliance S t a t i -
c i o n o r i e n t a l e belonging to the L i m o n i e t a l i a for the latter as-
sociation. The Cambhorosma ruthenica-community described by
OB~RDORSER (1.e.) is not well enough known to be placed defini-
tively.
In the area of Mesolonghi the S a l i c o r n i o - I - t a l o c n e m e t u m
s t r o b i l a c e i grows on clay soils, desiccated and covered with salt-
crusts in summer. The upper 1--2 cm of the soil can often be re-
moved, because they consist of loose shales. The ground water table
lies at a depth of 15--2o cm; the soil is well-aerated to a fairly large
depth. The conditions of the milieu on the growing place of the S ali-
cornio-Halocnemetum s t r o b i l a c e i are very extreme. The sa-
linity in the toplayer of the soil may increase until saltcrusts appear
upon the soil. Temperatures may raise to extreme great heights
HALOPHILOUS VEGETATION OF MESOLONGHI 1I 7
especially when the soil has recently been wet and the soil may be-
come desiccated until the toplayers are as firm as stone.
In the S a l i c o r n i o - H a l o c n e m e t u m s t r o b i l a c e i all occur-
ring species, with the exception of the Salicornia, have a distinct pref-
erence for the cover of the low shrubs of Halocnemum. Only Salicor-
nia grows independently also on the bare spots between the Haloc-
nemum-plants. One time a distribution as figured in fig. 3 was ob-
served.
In April and M a y the aspect of the vegetation is given by Haloc-
nemum together with a number of annual L i m o n i e t a l i a - s p e c i e s ;
in J u n e and J u l y the latter have disappeared and are all replaced by
the above mentioned form of Salicornia herbacea s.1.
Fig. 3. Distribution of different age-classes ofHalocnemum strobiIaceum. Mesolonghi,

April 1963 .
The H a l o c n e t u m s t r o b i l a c e i TOpA 1939 described from

Rumania has a quite different floristic composition and should be
placed in some still undescribed alliance of the H a 1o s t a c h y e t a li a
(ToPA 1939). Also the H a l o c n e m e t u m s t r o b i l a c e i TADROS
1953 is different from our association; its syntaxonomic position is
doubtful.
Probably the S a l i c o r n i o - H a l o c n e m e t u m s t r o b i l a c e i is
restricted to the Balkan Peninsula, possibly it may be found in
Asia Minor.
Arthrocnemetum BR.-BL. 1928. TABLE 7"
Syn.: Arthrocnemetum glaucum-Halocnemum strobilaceum-Ass. OBER-
DORFER I952 p.p.
i8 W . J . WOLFF
TABLE 7
Nr. 133 134 135 136 I37 x38 I39 14 ~ t4I 142 t43 t44
Date 16-IV I6-IV 19-IV 22-IV ~2-IV 22-IV 27-IV 29-IV 29-IV 2-V 2-V 7-V
'63 ~63 '63 '63 '63 '63 '63 '63 '63 '63 '63 ' 6 3-
Area M M M M M M M
M M M M M
S u r f a c e ( m 2) I2 75 I00 15 14 I00 I00 30 Ioo 4~ 20 70
Cover (%) 60 80--00 ~o 50 45 25 4~ 50 t5 80 45 30
Height (era) 3o 50 35 50 40 4o 70 40 I5 60 40 5o
Characteristic speciesof the

Arthrocnemetum:
Ar throenemum glaueum 3-3 5.5 2b.2 2b.~ 2b.2 2b.2 3.2 3.2 2b.2 4.5 3.3 2b.2
Frankenia pulverulerata . . . . . . . . -- -- -- +.1
Local characteristicspecies:
Triglochin bulbosum . . . . . . I. x -- -- -- I.~ -- +,!
Characteristicspeciesof the Staticion

gallobrovincialisand of the
Limonietalia:
Parapholis ineurva -- -- r -- -- +. I -- +.
Sphenopus divarieatus -- -- 1,2
Parapholis strigosa -- -- -- 2a~ x.2 1.2 +. I I. 3
Halocnemum strobilaceum -- -- 1.2 -- +,o -- -~-,I --

Plantago eoronopus -- -- +.I I,I i,I I . IIDX. +,I I,I I,l -- l ~ ,I
Spergularia salina -- -- -- i. i i. i i. i I. i 1.2 I,I "~-,I -- I.I
Characteristicspeciesof the
Sali~oenieteafruticosae:
Salieornia frutieosa -- -- -- +.2 1.2 -- +.2 --
Salieornia radieans -- -- 1.2 1.2 1.2 +. I 1.2 x.2 -- t.2 +.2
Puceinellia convoluta +,I -- -- -- +.2 2t-,2 1.2 -- -- 2a.2 ~a,2
Statlee ]imonium ssp. serotinum +.2 . . . . . +.I - -
Halimione portulacoides 2b.2 1.2 2a.2 3.3 3-3 ~a.2 2a,~ 2b.3 -- i .2 1.2 +.]
Companions:
Salieornia herbacea s.l. -- -- -- +. I +. I +. I x. I -- I.I -- I.l I,]
Hordeum marlnum -- - - - - 2b. 3 +.I +,2 -- 1.2
Differential species of the variant of

natural levees:
Salsola soda . . . . . . . .
Atriplex hastata -- -- -- r . . . . -- -- +.I --
Matthiola tricuspidata . . . . . . . .
Matricaria chamomina . . . . . . . .
Suaeda maritima . . . . . . . .
Species of the Saginlon

mediterraneum:
Polypogon maritimum . . . . . . . .
Crepis bulbosus . . . . . . . .
Centaurium tenuiflorum . . . . . . . .
Centaurium spicamm . . . . . . . .
Romulea sp. -- . . . . . . .
The A r t h r o c n e m e t u m Bm-BL. 1928 occurs in the area of

Mesolonghi on places situated higher than the S ali c o r n i o-H a 1o c-
n e m e t u m s t r o b i l a c e i . From table 7 it becomes clear that the
Greek vegetations bear close relationship to the A r t h r o c n e m e -
t u r n from southern France. Also the milieu is about the same: very
dry and salt clay which submerges only during high floods. Often
there is a distinct difference in height between the A r t h r o c n e m e -
t u r n and the neighbouring S a l i c o r n i o - H a l o c n e m e t u m s t r o -
b i l a c e i . In comparatively high places in the lagoons the associa-
tion becomes more reminiscent of the alliance S a g i n i o n m e d i t e r-
r a n e u m (page 121).
H A L O P H I L O U S V E G E T A T I O N OF M E S O L O N G H I I I9
I45 146 147 I48 149 15o I51 152 153 154 155 x56 157 158 159 16o
7-V 7og 2I-V I-VI 2-VI 22-IV 1-Vl 6-VI I-VI ~9-1V 2-V 3o-V x-VI x-VI ~-VI 22-IV
'63 "63 '63 '63 '64 "63 "63 '63 '63 '63 '63 '63 '63 '63 '63 '63
M M M M A M M M M IV£ M M N[ M 3¢[ NI
400 400 ZOO ZOO 25 75 zoo 50 6o Io I0 1o0 40 20 zoo ?
30 3o 30 30 90 4o 25 60 4° 60 50 3o 90 60 70 ?
50 40 40 50 ? 60 50 40 40 30 50 3° 5o 5° 50 ?
3.2 2b.2 2b.2 2b.2 5.5 2a.2 2b.2 3.3 2b.2 2b,2 3.2 2b.2 3.2 3.5 2a.2 2b.2
+.2
-- 2a.2 +.2 -- 1.2 -- -- +.i -- -- 1, 3 . . . . +.2

+.t +.~ 1.2 1.2 -- 0 +.a 1.2 - - +.2 -- +.i 1.2 -- x.2 2a.3
1,2 -- 1.2 2a.i 1.2 1.2 1.2 2a. 3 +.2 2a.3 1.3 1.2 1.2 I. 3 3"4 2a.3
. . . . +,2 -- -- -- +.2 1.2 -- 1.2 -- -- -- +oI
I*I 1.I I°l I.I -- I.I I.I +.2 +.I -- +.I 1.2 1,2 +.I ~a,I l.II=.
1.I 1.2 I,I I,I I,I I.I 2111.1 I°I "~-,I 1,2 1.2 I.I 1.2 1.I I,I 1.I
-- -- -- 1.2 -- 1,2 -- -- 1,2 -- -- -- 1.2 -- -- --

-- -- I,~ ia.2 1,2 1.2 -- +.2 1,1 1.2 . . . . +.2 --
. . . . . 1ol -- -- 1. ° -- "['.st -- -- +.2 -- +.I
-- -- - - +.2 -- +.1 -- +°2 I, 0 . . . . . . .
+.t -- +.2 2b.2 -- ~b.3 -- 3.3 J:.2 3.3 ~b.3 +.i 3.2 2b.3 1.2 +.x
I.I 1.1 +,I 1.I -- +.I I,l i.i 1.i -- 1.1 I.I 1,2 -- +.I I.!
-- 1,2 1,2 . . . . . . . l.~ -- -- -- 1.2 2b.3
J -- +.I +.I +.x -- -- +.I

-- @.I r +°I +.2 +,2 -- -- r
. . . . . . r m m -- -- r ° +.x +.2
+.2 +o.I ro ro
+°I . . . . . . +.1 +.x -- +.2 -- x.1 +.~ -- +.2
-- -- 1.2 . . . . . m -- +.2
. . . . . . . . +.2 -- +.2
m -- +,I +.2
m -- +.I
-- +.2
Along the shores of the lagoons there is a natural levee consisting

of shells deposited upon a clay soil. O n the sides of these levees an
Arthrocnemetum also occurs, often intermingled with species
belonging to the C a k i l e t e a m a r i t i m a e (fig. 4).
The soil normally consists of clay or sandy clay and is very dry
in summer. The soil is well-aerated but salinity does not rise as high
as in the soil of the S a l i c o r n i o - H a l o c n e m e t u m strobilacei.
The association is common near Mesolonghi. Near Arta we ob-
served only one place with Arthrocnemum-vegetafionsand near Salo-
nica none at all. But in our opinion record 443 of OSERDORFER
(1.C.) belongs to the A r t h r o c n e m e t u m . Also PIGNATTI (1953)
I~ W.J. WOLFF
TABLE 7 (Cont.)
Occurring in record:
Nr. I6r x62 x63 164 144: H y p o c h a e r i s a e t h n e n s i s +.x
Date 4-V 21-V a-u xo-u I53: Juncus subulatus L3
'63 '63 '63 '63 Melilotus indica r
Area M M M M I5o: Hordeum murinum +.t
S u r f a c e ( m e) 60 IOO zoo 40 A l l i u m sp. +. I
Cover (%) 50 3o 50 40 I54: S a g i n a m a r i t i m a L2
Height (cm) 20 50 70 25 Melitolus sicula +. x
I55: Melilotus sicula r
z58: C a k i l e m a r i f i m a +.x
Characteristic species of the Juncus subulatus +. I
Arthrocnemetum : x6o: B e t a v u l g a r i s ssp. m a r i t l m a +,I
Ar throcnemurn glauetxm 2a.2 3.3 3.2 2b.2 Salsola kali r
Frankenia pulverulenta -- +. I +. I +. I Rumex conglomeratus +.2
i61 : Me]ilotus indica +.2
Local characteristic species: I62: Sagina maritima +.2
Trigloeh~ bulbosum . . . . x63: C a k i l e m a r l t i m a r
I64: Aeluropus litoralis +.2
Characteristic species of the Statidon Evax pygmaea +. I
galloprovindalis and of the Tortella flavovirens +.2
Limonictalia:
Parapholis incurva 2b.3 -- -- +.z
Sphenopus divarieatus 1.2 1.1 +.2 1,2 A = lagoons of Arta
Parapholis strigosa -- 1.2 2b.3 1.2
M = lagoons of Mesolonghi
Halocnemum strobilaceum . . . .
Plantago coronopus I. 1 I, I i. 1 I, I
Spergularia salina I.I 1.2 1.I 1.I
Salicornletea fruticosae:
Salieornia fruticosa . . . .
Salicornia radicans 2a.2 --- -- 1.2
Puccinellia convoluta . . . .
S t a t i c e l i m o n i u m ssp. s e r o t h a u m 1.2 -- -- --
Halimione portulacoides 2b.3 +.2 2b.3 +,2
Companions
S a l i c o r n i a h e r b a c e a s.]. -- +,I I,I - -
Hordeum marinum +.x 1.2 +.2 1.2

Differential species of the variant of
natural levees:
Salsola soda . . . .
Atriplex hastata . . . .
Matthiola tricuspldata . . . .
Matricaria chamomilla . . . .
Suaeda maritima __ __ ,3(_ I --
Species of the Saglnlon
mediterraneum:
Polypogon marifimum +. I 1.2 -- +.2
Crepis bulbosus +.2 -- -- --
Centaurium tenuiflorum +. 1 -- +, I +. i
Centaurium spicatum -- -- +,z I.I
Romulea sp. I.I +.2 -- +.I
considers part of the records of OBERDORFER as belonging to the

Arthrocnemetum.
The A r t h r o c n e m e t u m BR.-BL. 1928 was already known from
Portugal, Spain, mediterranean France and Sicily (BRAuN-BEAN-
QUET C.S. 1952 ) . The A r t h r o c n e m e t u m g l a u c i described by
EIG (1946) from Israel differs considerably from our association.
Saginetea maritimi WESTHOFF,VAN LEEUWEN & ADRIANI

1961
Saginetalia maritimi WESTHOI~I%VAN LEEUWEN &
ADRIANI 1961
HALOPHILOUS VEGETATION OF MESOLONGHI 12I
Saginion mediterraneum all. nov.

ROMULEO-SAGINETUMMARITIMAEass. nov. TABLE 8.
At a level between that of the A r t h r o c n e m e t u m and that of
the inland meadows of the T h e r o - B r a c h y p o d i e t a l i a we find
an association belonging to the recently created class of the S a g i n e-
t e a m a r i t i m a e (TOx~.N & WESTHOFF, 1963). This class comprises
the vegetations of the dry bordermilieus between the saltmarshes
and the fresh inland vegetations, e.g. the T h e r o - B r a c h y p o -
d i e t a l i a . For this reason we find in the records a number of typi-
cal L i m o n i e t a l i a - s p e c i e s , but also several species belonging to
the T h e r o - B r a c h y p o d i e t a l i a . For the description of the milieu
we refer to TUXEN ~2 WESTHOFF (1.c.).
For the related vegetations of the atlantic coast of Europe the
alliance S a g i ni o n m a r i t i m a e WESTHOFF,VAN LEEUWEN & ADRI-
ANI 1961 was created, but in our opinion the mediterranean vege-
tations of this kind should be placed together in a new alliance,
which we propose to call S a g i n i o n m e d i t e r r a n e u m all. nov.
The characteristic species of the alliance are probably: Bellis annua
and Polypogon maritimum. Moreover the alliance possesses a number
of differential species compared with the S a gi ni o n m a r i t i m a e,
e.g. the species occurring also in the L i m o n i e t a l i a : Sphenopus
divaricatus and Parapholis incurva.
In this alliance S a g i n i o n m e d i t e r r a n e u m we want to in-
clude our new association the R o m u l e o - S a g i n e t u m m a r i t i m a e ,
but also the S a g i n o - m a r i t i m a e - T o r t e l l e t u m flavovirentis
BEEFTINK, Tx. & WESTHOFF 1963 described from the Camargue
(TOxEN & WESTHOFF, 1.C.). The relation with the S t a t i c e t u m
e c h i o d e s (TALLON) BR.-BL. 1931 should also be studied.
Ecologically the S a g i n i o n m e d i t e r r a n e u m and the associa-
tions in it may be sketched as the dry borderzone between the S ali-
c o r n i e t e a f r u t i c o s a e and some fresh inland vegetations, mostly
belonging to the T h e r o - B r a c h y p o d i e t e a . The surface-layers of
the bottom of these associations may be temporarily rich in NaC1
(see also T/dXEN & WESTHOFF, 1.c.) and for this reason there is a
close relationship between the S a g i n e t a l i a m a r i t i m a e and the
L i m o n i e t a l i a . In the latter the toplayers are also enriched in
salts, because in these vegetations we find in summer an upward
waterstream caused by evaporation. In this way the occurrence of
several species in both the L i m o n i e t a l i a and the S a g i n e t a l i a
m a r i t i m a e may be explained: e.g. Plantago coronopus, Parapholis
strigosa, P. incurva, Sphenopus divaricatus and Hordeum marinum. There-
fore it is difficult to separate the L i m o n i e t a l i a from the S a g i -
n e t a l i a m a r i t i m a e and perhaps we ought to consider the S a g i -
n i o n m a r i t i m a e as an atlantic radiation of the L i m o n i e t a l i a .
i22 W.J. WOLFF
TABLE 8.
iNr. 165 I66 I67 168 169 I7O I7 I 172 173 I74 I75
Date 2I-V 22-IV 29-IV I9-IV 27-IV 6-VI 29-IV 29-IV 2o-IV 2o-IV o7-IV
'63 '63 '63 '63 '63 '63 '63 '63 '63 '63 '63
Area M M M M M M M M M IV[ M
S u r f a c e ( m 2) 25 75 IOO 15o 75 IO0 100 Ioo 35 25 8
Cover (%) 90 75 9o 5o 7o 70 50 4~ 7~ 50 3~
Height (era) ? 5o 5(2o) 25 I5(4o) oo 5(oo) 15 3~ I5 5
Characteristic speciescombinationof the

Romuleo-Saglnetum maritimae:
Romulea bulboeodium - +-~ - }L,
Romu]ea roliei ) 1.1 )I.I )1,i 27:: )I.I )2m, 3 } +,o
Anagallis arvensis ssp. eoerulea -- 1.2 I.I -- +.2 -- +.I - - - - 1.2 - -
Sagina maritima -- I.O -- I.X 1.2 -- -- - - 3.4 1.1 4 . I
Characterlstle speciesof the Sagirdon

mediterraneum:
Bellis annua t.2 2a.3 1.2 1,2 4.o +.I x.2 1.2 1 . 2 - - ~ 1.2 - -
Polypogon maritimum 2a.3 I*2 ob.o--3 -- oa.3 -- 1.2 2a.o -- -- --
Speczes of the ffuncetalia mantzmz:

Juneus aeutus 4.2 -- +.I . . . .
Juncus ambiguus I.o -- 1.2 4.I -- -- ore.5 IoI - - ~ - -
C e g l t a u r i u m t e n u i ~ t o r uYlx 1,1 ) +,, ) I.I } 4.1 )4.2 I.I I,I } I.I } I.I } I.I -
- -
Centaurium spieatum -- +. i --
Trifolium resurpinatum +.2 -- 1.2 -- -- -- x.o - - - - 4 . 2 - -
Trifolium tomentosum 1.2 -- 1.2 -- -- -- 4.0

Ranunculus sardous -- -- +.2 -- r -- +.o
Species of the Limonletalia:

Plantago coronopus 3.3 3-5 2a.2 0m.I I.I 2m.3 2m.5 2a.2 3.4 2a.5 ob.5
Parapholis strigosa -- -- I.O -- 1.2 4.2 1.2 1.2 -- 2a,3 1.0
Parapholis incurva 1.2 +.o -- +. I 4,2 -- -- -- 2a.o -- I~
Sphenopus divarieatus +.2 l.O -- 4.I I.I 4.I I.O -- I.I oa.5 --
Hordeura marinum 2a.o ~ -- 4.I -- I.o -- 1.2 -- -- --
Spergularla salina 1.2 -- -- i.i 4.1 om.I -- +.2 +.x -- 1.2
Triglochin bulbosum +.2 z.2 . . . . +.I - - - - 1,2 - -
Arthrocnemum glaucum -- oa.o -- 4.2 4.2 -- 2a.2 2a.2 -- I.O +.I

I-Ialoenemum strobilaceum . . . . . . 1.0 -- oa.O -- --
Salicornia radicans oa.o -- -- 1.2 -- 4.2 -- 1.2 - - - - - -
Halimione portulaeoides -- 1.o -- +.o 3.3 -- -- 4.2 -- --

Puccinellia convoluta -- 1.2 -- -- -- +. I -- 4.2 -- -- --
Statice limonium ssp. serotinum +.1 2a.o +.x I.O l.o 2a.o --
Specie, of the Thero-Brachypodietalia:

Allium sphaerocephalum -- 1.2 -- +.o +.2 1.2 --
Hedypnois rhagadioloides +.o -- I .O -- -- -- 4.2
- - - - + . O - -
Medicago polymorpha 1.2 -- +.I . . . .
Crepis bulbosa -- I.O -- -- 1.2 1.2 -- 1.2 -- -- --
Evax pygmaea -- +.2 1.3 . . . .
Poa jubata -- -- 2a,2 . . . . +.o +.i -- --
Eryngium eL creticum +. I 4. I . . . . .
Iris sisyrhynchium x. I +.2 +. I 4. I -- -- --
Koeleria hispida -- -- I.O -- -- +. I --
Companions:
Beta vulgaris ssp. maritima 4.1 52 -- 4,I 4.0 4.0 --
Asphodelus microcarpus 4. I I. i 4. I +.2 -- -- --
Lolium loliaceum -- -- -- -- +.2 +.o -- 4.2 -- -- --
Melilotus indicus +.o x .', 1.2 -- +. i -- 4.2 4.0 -- -- --
Hypoehaeris aethnensis -- -- -- 4.2 -- -- -- - - 4 . 2 - - - -
- - - - - - + . I
Matricaria chamomilla 4.1 -- +~ -- +.2 -- +.o
Lolium perenne I.O . . . . +.O --
Indet. Compositae -- 1.2 -- +.2 -- 4.2 --
O c c u r r i n g i n r e c o r d 165: P l a n t a g o l a g o p u s + . 2 , L y t h r u m s p . ~ . 2 , H y p o c h o e r i s c r e t e n s i s 4 . I , A t r i p l e x h a s t a t a + . I , L o l i u m s t r i c t u m I .
C i c h o r i u m s p . 4 . 0 ; 166: P l a n t , a g o l a n c e o l a t a 2 a . 3 , A e l u r o p u s l i t o r a l i s + . x , E u p h o r b i a e x i g u a 4 . 1 , V a i l l a n f i a h i s p i d a + . I , H o r d e u
m u r i n u m + ~ 2 , c f . G a u d i n i a f r a g i l i s 4 . 2 , l ~ u p h o r b i a h e l i o s c o p a 4 . 2 , M a l v a s p . + . x ; 1 6 7 : P l a n t a g o l a n c e o l a t a I . o , S c l r p u s c e r n u u s x.
C a r l i n a s p . ('~) + . x , P a r e n t u c e t l i a l a t i f o l i a + . I , Cuscuta sp. r.2, Trifolium campestre 4.2; 168: P o a a n n u a r , C a r e x d i v i s a 4 . 2 ; 16
M a t t h i o l a t r i c u s p i d a t a 4 . I , S o n c h u s s p . r ( j u v ) , C u s c u t a s p . 4 . 2 , T o r i l i s n o d o s a 4 . I ; 17o: C h e n o p o d i u m a l b u m r , S a l i e o r n i a h e r b a e ,
s . l . 4 . 1 ; I 7 3 : E r a n k e n i a p u l v e r u l e n t a 4 . o ; 174: S c i r p u s m a r i t i m u s 1 . x , L y t h r u m c f . t r i b r a c t e a t u m r , V a i l l a n t i a h i s p l d a + . x .
1Vi = l a g o o n s o f M e s o ] o n g h l .
r
.&_
9
EBETO-
MATTHIOLETUH
TRICUSPIDATAE
[ HELL- REMAINS
f SALICORNIO-
HA LOCNEHE-fUH
STROIglLAC EI
t
ARTHROCNENUM
Fig. 4. Vertical section through a stand of the Beto-Matthioletum tricus-
pidatae. Mesolonghi,June 1963.
To justify this view one should have much more data, so we only
mention the possibility.
The Greek association of Romulea and Sagina grows on well-aer-
ated clay and sandy clay soils on a level above that of the A r t h r o c -
n e m c t u m . The differential species combination of the R o m u 1e o-
S a g i n e t u m m a r i t i m a e is Anagallis arvensis ssp. coerulea, Romulea
bulbocodium, Romulea rolld and Sagina maritima. The vegetation is
well-developed in April and May; later on the mostly annual spe-
cies desiccate. The aspect of the association is a dry, short vege-
tation with here and there an high plant. The vegetations are
grazed, particularly by sheep.
Juncetea maritimi BR.-BL. 193 I
Juncctalia maritimi (BR.-BL. 1931 ) era.
Syn.: Caricetalia fuscae PmNATTI 1953
Alliance?
Juncetum maritimi (I~i]IBEL 193o ) PIGNATTI 1953. TABLE 9"
Around all hills and rocks near Mesolonghi surrounded by salt-
marshes there is a belt of vegetations dominated by two species of
Juncus: j . acutus and J. maritimus. Our only record of these vegeta-
tions represents their composition fairly well. They much ressemble
PIGNATTI'S J u n c e t u m m a r i t i m i , which, however, certainly
should not be placed in the J u n c i o n m a r i t i m i PIGNATTI 1953.
This alliance is a very heterogenous assemblage of associations
which in our opinion belong to three different classes. The J u n c e -
t u r n m a r i t i m i may be placed in the J u n c e t a l i a m a r i t i m i and
perhaps in a new alliance.
124 W.J. WOLFF
TABLE 9-
Hr. 176 I77 I78 x79

Date 5-VI x-V r-V x-V
'64 '63 '63 '63
Area M M M M
Surface (m s) Io0 IO0 25 9
Cover (%) go ioo ioo ioo
Height (cm) ? 50 50 50
Characteristic species combination of the

ff'uncetum rnaritimi:
Juneus maritimus +.2
Carex extensa var. graeca
I n u l a crithmoides
Characteristic species combination of the

ffuncus acuhts - Salicornia frutieosa -
consociation:
Juncu~ aeutus 1.2 ~m.5 3.5 4-5
Salieornia fruticosa +. x 3.5 3.5 3.5
Companions:
Sallcornia radleans r.~
Salicornia herbacea s.1. +. I
Halimione portulacoides 3.5 x.2 - -
Statice limoniurn ssp. serotinum + . I - - - -
Phragmites communis +. I
Scirpus maritimus +.I 1.3 1,2
M = lagoons of Mesolonghi
The soil is mostly reduced and very wet. We obtained the im-
pression that the mostly stagnant water is rather brackish. The
chlorinity of the water of a creek which drained these vegetations
was 6. 7 ~ CI' (fig. 2).
The ecological position of these vegetations seems to be the same
as that of the atlantic suballiance L o t o - T r i f o l i o n as provision-
ally described by WESTHOFF et al. (1961). They have about the
same position as this suballiance in the zonation of vegetation be-
tween salt and fresh wet milieus. Also the role ofJuncus maritimus
does not seem to differ very much. In our opinion the J u n c e t e a
m a r i t i m i ought to be seen as the mediterranean equivalent of the
suballiance L o t o - T r i f o l i o n of the A g r o p y r o - R u m i c i o n cris-
p i. Their ecological position may be described as the transition
zone between fresh and salt milieus in a wet or at least temporarily
wet milieu. We want to include in this order the alliances J u n ci o n
m a r i t i m i BR.-BL. 1931 , P l a n t a g i n i o n c r a s s i f o l i a e BR.-BL.
1931 , T r i f o l i o n m a r i t i m i BR.-BL. I93i and T e t r a g o n o l o b i o n
s i l i q u o s i PIGNATTI 1953. Vegetations related to the J u n c e t u m
m a r i t i m i are known from southern France (BRAuN-BLANQUET
et al. i952: J u n c e t a l i a m a r i t i m i ) , Venice (PIGNATTI I953:
Caricetalia extensae, Juncetum m a r i t i m i ) , Macedonia
(OBERDORrER I952: Wiesen von Juncus acutus), Palestine (ZOHARY
I947: J u n c o - P h r a g m i t i o n ) and Egypt (TADROSI953: association
o f t h e J u n c i o n m a r i t i m i ; TADROS & ATTA I958: P l a n t a g i n i o n

crassifoliae).
Plantaginetea maioris Tx. & PRSO. I950

Plantaginetalia maioris Tx. 195o
Polygonion avicularis BR.-BL. 1931
Community of Hordeum marinum and Cynodon dactylon TABLE I O.
In the area of the lagoons of Arta a dry and elevated clay soil
was covered with a very short and dense vegetation with Hordeum
marinum and Cynodon dactylon as dominant species. This meadow was
intensively grazed by sheep. This community seems to be closely
related to the Cynodon dactylon-Plantago coronopus-Ass. (HoRvATIC
I934) Tx. I95O from Dalmatia and to the subassociation t i p i c a
PIGNATTI I953 from Venice, both belonging to the alliance Poly-
g o n i o n a v i c u l a r i s BR.-BL. I93I. According to TtiXEN (I950)
this alliance has to be placed in the P l a n t a g i n e t e a m a i o r i s
Tx & PRSG. I950. The association mentioned by TOXEN is only
known from the Adriatic coasts.
Cakiletea maritimae Tx. & PRSG. 1950

Thero-Suaedetalia (BR.-BL. & DE BOLOS 1957) BEEFTINK
I962
Thero-Suaedion (BIn-BE. i93 I) Tx. I95O
Suaedo-Kochietum hirsutae BR.-BL. 1928 TABLE I I.
Near Arta along the shores of the lagoon north-east of Salaora
we observed a vegetation of Bassia hirsuta ( = Kochia hirsuta) and
some other nitrophilous species on banks of plant-remains (Ruppia)
washed ashore in winter and deposited on wet clay. Without doubt
these vegetations must be placed in the S u a e d o - K o c h i e t u m
h i r s u t a e described by BRAuN-BLAN•UET from southern France.
OBERDORFER (1.C.) mentions the same vegetation from the AMos-
delta near Salonica, but in our impression only his record 454,
which clearly differs from the other ones, has to be incorporated
into this association. His other records probably belong to the
Thero-Salicornion.
Near Arta on the banks mentioned above but older than one
year, Rumex pulcher dominated. We distinguish the latter vegeta-
tions as S u a e d o - K o e h i e t u m r u m e c e t o s u m subass, nov.
According to BEEFTINK (I965) this association has a discontin-
uous areal:
1. the coasts of Denmark, Germany and the Netherlands;
2. the coast of the Mediterranean from France to Venice and
3. the shores of the Black Sea.
126 W . J . WOLFF
TABLE I O.
Nr. i8o 181

Date 2-VI 2-VI
'64 %4
Area A A
S u r f a c e ( m 2) 25 25
Cover (%) 7o 8o
H e i g h t (era) 5--1o 5--m
Dominant species:
Hordeum marinum 4.5 2m.5
Cynodon daetylon 1.2 3-5
Species of the ffuneetalia maritimi:
Centaurium tenuiflorum +. I I. z
Juncus ambigaus I. : 2m. I
T r l f o l i u l n cf. f i ' a g i f e r u m -- rJ',
Species of the Saginetalia maritimae
and the Limonietalia:
Plantago eoronopus 2m.5 2m.I
Parapholis strigosa -- +. I
Sphenopus divaricatus -- I. I
Spergularia salina 1.~ - -
S t a t i c c l i m o n i u m ssp. s e r o t l n u m +.2 --
Companions:
cretfca
C r e s s a x. I --
Matricaria chamomilla -- +.
Cirsium arvcnse r
Koeleria hispida -- +.2
Lolium perenne -- +.~
A = lagoons of Arta.
TABLE I I.
Nr. i82 i83 i84 i85 Ob, 186 I87 188

Date 2-VI 2-Vl 2-Vl 2-Vl 3-1X 2-VI 2-VI 2-Vl
'G4 '64 '64 '64 '4r '64 '64 '64
Area A A A A Th A A A
S u r f a c e ( m ~) 20 2o :o io ? 2o 20 40
Cover (%) 80 50 50 30 60 70 40 30
CharaeterlsKe species of the Suaedo-Kochletum

hirsutae:
Bassia hirsuta 4-5 3.3 3-3 2a,3 +.2 4.5 2a.2 1.2
Characteristic species of the subassodation

rumeeetosum:
Rumex pulcher . . . . .
Characteristic species of the Thero-Suaedlon:

Suaeda maritima +.2 +.2 i.o 1,2 1.2
Salsola soda +.r +.I +.I i.o +.2 -- 1.2 --
Characteristic Cedes of the Cakiletea

nuaritlmae:
Cakile maritlma r +. I - - - - - - 1.2 1.2 2a.2
Atriplex hastata -- +. I +.2 -- -- -- +.: --
Companions:
S a l i c o r n i a h e r b a c e a s.1. 1.2 1.2 -- 1.2 4.4 1.2 I.I I.I
Halimione portulacoides +.2 -- -- - - +.2 1.3
Seirpus maritimus +. I . . . . +.~ +.: --
( 3 h c n o p o d k l r n spec, - - + . :~ - - - - - - -- +.: L~
O c c u r r i n g i n r e c o r d 182: P h r a g m i t e s c o m m m a l s + . i ; : 8 3 : P u c c l n e l l i a festuciform~s + . 2 ; i 8 4 : A s t e r t r i p o K u m + . I ; O b . : A g r o s t l s a l b a
fo. ( + ) , S u a e d a s p l e n d e n s ; i 8 7 : S a l i c o r n i a r a d i c a n s 2a. 2, S t a t i c e l i m o n i u m ssp, s e r o t i n u m + . 2 , S a l i c o r n i a f r u t i c o s a 1.2.
Ob. = record 454 of OBERDORFER (I952). A = lagoons of Arta, Th = delta of R. Axios near Salonica.
H A L O P H I L O U S V E G E T A T I O N OF M E S O L O N G H I 127
The above-mentioned Greek records more or less fill the gap be-
tween the west-mediterranean and the pontic areal. FREI (1937)
mentions the association from Sicily.
Euphorbietalia peplis Tx. 195o

Syn.: Cakiletalia maritimae PIGNATTI 1953
Euphorbion peplis Tx. 195o
Syn.: Salsolo-Polygonion maritimi PIGNATTI I953
Cakilion litoralis PIGNATTI 1953
Association of Xanthium strumarium and Salsola kali OBERD. ~2 Tx.
195o. TABLE I2.
On the beach of the outer sandy coastal ridge near Mesolonghi
we discovered degenerated examples of the association of Xanthium
strumarium and Salsola kali. Some intermingling with dune-plants
was observed. Also near Nea Kallikrateia (Chalkidike) this associa-
tion was met with, but in a much richer form. As we had here no
opportunity for making surveys, we noted only its occurrence.
OBERDORFER (I.c.) gives related records from Macedonia and
Thrace, which were used by T/dXEN (I950) for the creation of these
association mentioned above. The Ass. a Cakile maritima e Xanthium
(BEG. I94I ) PmNATTI I953 seems very closely related.
The drift consists of plant-remains and shells deposited on a sandy
beach. The association is only known from Greece, but probably
has a wider distribution.
Community of Cakile maritima and Chenopodium spec. TABLE 12.
O n sandy beaches along the shores of the Gulf of Arta near Ko-
ronisia we observed a vegetation of Cakile maritima and Chenopodium
spec. on drift material. We were not able to identify this species of
Chenopodium because ripe fruits were lacking. However we record
this vegetation as we suppose that it represents an unknown asso-
ciation.
Atriplicetalia littoralis SISSINGH I946

Alliance?
Beto-Matthioletum tricuspidatae ass. n o v . TABLE ! 3-
Along the shores of the lagoons of Mesolonghi big banks of shells
washed ashore m a y be observed. On these banks a luxuriant vege-
tation of Matthiola tricuspidata, Matricaria chamomilla, Beta vulgaris
ssp. maritima and other nitrophilous species occurs. O n the slopes
of these walls we observed an A r t h r o c n e m e t u m which more or
less intermingles with the previous vegetation. Therefore we have
to describe the previous vegetation as a kind of "Teppich-Gesell-
I28 W.J. WOLFF
II I II III 4~
~+~
f~ 4
~÷ 4 11 I
I f
l E
l 1 II I
~ : ~ ~°o ee~
ee~
J~
+_~.~
a ~ ~.
÷
~ ~ .
+ °
~,o ~ :~'~ i~ ~ ~o
~Q
H A L O P H I L O U S V E G E T A T I O N OF MESOLONGHI I29
TABLE 13.
Mr, 206 207 208 209 210 2II 212 213 2I~ 2t5 216 217
Date 2-V 22- I V 27- I V 27- I V 27-IV 2-V 3o-V I-VI 1-VI I-VI 30"V $o'V
'63 '63 '63 '63 '63 '63 '63 '63 '63 '63 '63 '63
Area M M M M M M M M M M Yl lVi
Surface ( i # ) IO 20 50 20 20 15 60 40 3o 20 20 4~
Cover (%) 2o x5 50 4o 20 70 4o 50 3~ 40 60 4o
H e i g h t (cm) 3~ 20 ? ? 3~ 5~ 3o 4~ 4~ 50 40 50
Characteristic species combination of

the Beto-Matthioletum tricuspidatae:
Matthiola tricnspidata 1.2 1.2 1.2 2b.3 +.2 +.2 +.2 2b.3 1.2 1.2 +.2 --
Beta vulgaris ssp. m a r i t l m a 2a.2 I~ I,2 I.I I~ +.2 +.I 1.2 ~-~ +,2 ~ +,2
Atriplicetalia litoralis:
Atriplex hastata +.i . . . . . . +.2 +.i +.2 +.2
Senecio vulgaris - - +.x +.i 1.2 +.1 -- - - +.2 - - - - ~ ~ I
M a t r i c a r i a ehamomilla 1.2 I,2 2b.3 +~ 2a.2 +.2 +.2 2b.3 1.2 2a.3 +.~ 1.2
Cakiletea maritimae:
Cakile m a r i t i m a . . . . . . . - - - - + . 2 - - - -
Salsola kall . . . . . . . +.t +.i - - - - - -
Salsola soda . . . . . . . +,I - - +.I -- - -
Suaeda m a r i t i m a - - +.i . . . . . . i.i +.x 1.2 +.2
Nitrophitous spades:
Atriplex cf. rosea . . . . . . . +.i +.t +.x -- - -
Hordeum murinum I. 5 - - r 1,2 - - - - - - +.2 . . . .
Species of the LimonietaIia:

Arthrocnemum glaucum +.2 +.o -- 2b.3 +.~ 2a.2 3.3 2b.2 2a.2 2b.2 t.o ob.3
Parapholis incurva -- +.I 1,2 -- 1.2 . . . . . . .
Sphenopns divaricatus -- --~o 1.2 -- 1.2 "~-*I -- "~.I I~ ~ +~ -+~ "~-.X
Plantago coronopus -- +.I 2b,2 +.I x.2 x.3 +.x +.t +.I +.x ~ +.t
H a l o c n e m u m strobilaceum - - 1 . 2 . . . . . . I 9 2 1 . 2 - - - -
Sagina m a ritilna 1
Parapholis strigosa - - x~ 2b.2 2a.3 2b.3 4.5 3.3 2a.3 2a.3 1.2 4,5 ob.3
Spergularia salina l + " I l,I l I.I 1.3 1.2 -- I.I I.I 1.2 1.2
Polypogon i n a r i t i m u m - - - - 2b. 3 - - - - - - +.2 +-3 . . . .
Hordeum marinum - - +.i +a - - +.~ +.x - - - - +.2 - - +.~ ~.~

Species of the Salicornietalia fmticosae:
Salicornia fruticosa - - - - - - 1.2 - - - - x,2 -- -- - - +.~ +.2
Salieornia radicans . . . . +.2 . . . . 2a.~
Puecinellia convoluta -- +.2 -- -- -- +.i -- -- i.~ +.i +.2 +.x
Halimione portulacoides 1.2 2a.2 +,x x.2 -- +.2 i.o -- -- +.2 2b.3 x.2
Companions:
Salieornia herbaeea s.1. -- -- +.I -- -- -- I.I -- I.I I.I ~ I,l
Melilotus sicula +.i x.i +.i -- +.x -- -- i.l -- -- +.e --
Melilotus indica -- -- -~.i -- -- dl-.i -- I~ I*I "~,I "+.l ~-.I
Sonchus asper . . . -- . -- +3 +.2 +.X -- -- +.X
Crepis bulbosa -- -- +.2 -- 0 1,2 -- +.2 +.2 -- ~.2 --
Aliinm sphaerocephainm - - - - X.2 - - 1.2 +.2 - - I.I -- - - +.2 --
R o m u l e a sp. - - i.z i.t - - - - +.2 - - +.~ . . . .
Lolium loliaceum -- -- +.2 +3 +.i . . . . . . .
O c c u r r i n g in record 207: Amaranthus albus r; 208: Juncus ambiguus + . 2 , X a n t h i u m spinosum r, Bellis a n n u a + . 2 , Chenopodlum sp. r;
21 o: Allium sp. t.2; 21 x: Medicago polyinorpha + . x, Vaillantia hlspida + . x; o 15: Chenopodium r u b r u m + . I, R a n u n c u h s sp. r, Sclrpus
maritimus r ~
M = lagoons of Mesolonghi.
schaft" (Tf3xEN 1950 ) in the vegetation. Fig. 4 p r o b a b l y clears u p

this situation.
T h e relationship between these two associations can be f o u n d
in the substrate. T h e fairly high elevation accounts for the occur-
rence o f the A r t h r o c n e m e t u m, whereas the influx of fresh organic
i3o w . J . WOLFF
matter accounts for the occurrence of the nitrophilous community.

When the influx of organic matter becomes less only the A r t h r o c-
n e m e t u m remains, as we observed in many instances. The Arthroc-
nemum-vegetation does not function as an agent catching drift ma-
terial, at least not to a large extent, because the banks of shells
also occur without a vegetation of Arthrocnemum.
In relation to the high elevation of the association the occurrence
of some elements of the S a g i n i o n m e d i t e r r a n e u m is possible.
In I964 we only observed some depauperated fragments of the as-
sociation on places where it occurred in a luxuriant state in I963.
The syntaxonomic position of this association is doubtful. It
bears a ressemblance to the atlantic A t r i p ! i c e t a l i a l i t t o r a l i s
SISSING~ I946. But according to T/3XEN (i95o) this order (called
C a k i l e t a l i a m a r i t i m a e ) is replaced in the mediterranean area
by the E u p h o r b i e t a l i a p e p l i s . However, also PIGNATTI (1.C.)
records vegetations ressembling the A t r i p l i c e t a li a from Venice.
Therefore it will be useful to re-examine the relationship between
the A t r i p l i c e t a l i a and the E u p h o r b i e t a l i a . We think it pos-
sible that the E u p h o r b i o n p e p l i s Tx. I95o has to be included in
the order of the A t r i p l i c e t a l i a l i t t o r a l i s .
Consociation belonging to the order Juncetalia maritimi

Consociation of Juncus acutus. TABT.~ 9"
Near the town of Aitolik0n in the province of Mesolonghi we
observed an extensive area with an aberrant vegetation dominated
by Juncus acutus in a vegetation of Salicornia fruticosa. Also Scirpus
maritimus and Phragmites communis seemed to be characteristic. In
the lower layer of the vegetation we only observed Salicorniafruticosa.
This type of vegetation was observed in b o t h subassociations of the
S a l i c o r n i e t u m f r u t i c o s a e as described on page II2.
The conditions are in general the same as in these subassociations
but the difference seems to be in a fairly large influx of fresh water
into the Juncus acutus-consociation.
The classification of this consociation is difficult and we would
prefer not to choose a definite standpoint.
Not classified
Sociation of Phragmites communis
In the zonation of vegetation-units around hills and rocks in the
saltmarshes the sociation of Phragmites communis forms the innermost
belt. It is found between the feet of the hills and the Juncus-vege-
tations (page II4, fig. 2) on a very weak and wholly reduced
TABLE 14 .
Nr~ 218 219 22o 22x 2oo 223 224 2o5 226 227 228 229 230
Date 24-IV 24-IV 2o-IV 24-IV 2-V 2-V lI-V 24-V 24-V o4-V 24-IV o4-IV 24-IV
'63 '63 '63 '63 '63 '63 '63 '63 '63 '53 '63 '63 '63
Area M M M M M M M IV[ M M M M M
Surface ( m s) 400 4oo lOO 400 200 200 50 605 625 625 400 400 400
Cover (%) IO 20 IO 25 60 5~ 40 3~ X5 30 20 I5 5
Height (cm) I S ( I 0 0 ) IS(IOO ) 30 X5(IO0 ) 2O 20 70 oo oo 2o(Ioo) I5(IOO) I5 I5
Charaaeristic species?:
Cyperus distaehyus oa.3 oa.2--3 2a.2 L3 4.5 4.5 ob.3 3.5 2b,5 2b.5 1.3 1.3 +.2
Polypogon monspeliensls -- +.0 -- r +.I 1.I 1.2 x,O +.2 1.2 2a.o -- --
Agrostis verficillata - - +.2 -- -- -- 1.I +.I - - 1.0 - - - - - -
Species of the Phragmltion:

Scirpus maritimus 0m~ 2m,2--3 I.I -- +.x +.I I.I +o.! +o.i +~ I~ I~ I.O
Seirpus litoralis x.0 1.0 -- +.2 +.2 +.0 0a.3 -- -- -- +.i +.x +.t
T y p h a of. a n g u s t a t a +.2 - - +.i . . . . . r~ - - +%i +~
Spedes of the Sallcornion

fruticosae:
S a l i e o r n l a frufieosa +.2 -- +.2 -- -- +.i -- -- -- +.2 +.0 --
Salicornla radieans +.2 +.2 -- 1,2 -- +.2 2a..2 -- -- -- 1,2 I.O 1.2
Puecinellla fcstuciformis -- - - +.I +.2 +.O - - I.I -- -- -- 1,2 -- - -
Aeluropus litoralis 1.2 1.2 +.2 2b.2 +.2 +.2 1.2 -- I.o +.I I.I 2&.2 t. o
Companions:
S a l i e o m i a h e r b a c e a s.L -- -- +.x 1.2 I.I I.Z t.2 +.I +.I +.I I.I 2m. 5 om. 5
Juncus acutus 1.2 1.2 +.I oa.2 -- -- 1.2 +~ -- 1.0 2a.2 -- --
I-Ialotmemum
strobilaccum - - +.z . . . . . . . . . +.z
Atriplex hastata r . . . . . r +.I r -- -- --
C h l o r o p h y c e a e +.2 +o i o . . . . . . . + __ __
O c c u r r i n g i n r e c o r d 218: Z a n n i c h e l l i a p a l u s t r i s s s p . p e d i c e n a t a + . 2 ; 2 2 z : Juncus m a r i t i m u s t . 2 ; 222: J u n c u s s u b u l a t u s + . 2 , T a m a r i x sp.

+ . x ; 2 2 4 : S t a t i c e l i m o n i u m ssp. s e r o t i n u m + . t ; 2 2 8 : P a r a p h o l i s i n c u r v a x.2, S p h e n o p u s d i v a r i c a t u s + . 2 , G y a n o p h y c e a e 5.5.
soil. The reed m a y become fairly high (I.5--~ m), but remains
sterile. Sometimes the undergrowth of the vegetation is formed by
Salicornia fruticosa.
We do not want to classify this sociation.
Not classified
Community of C.yperusdistachyus and Polypogon monspeliensis
TABLE 14 .
On places where before a S a l i c o r n i o - H a l o c n e m e t u m stro-

b i l a c e i occurred and which are flooded in summer by fresh water
from the rice-fields a remarkable vegetation is found. The physio-
gnomy of this vegetation is determined by Cyperus distachyus.
The combination of species and the milieu of the vegetation res-
semble the order P a s p a l o - H e l e o c h l o e t a l i a BI~.-BL. i95~ de-
scribed from southern France. Also the community ofCr_ypsis aculea-
ta and Heleochloa schoenoides and that of Fimbristylis dichotoma and
C.yperusfuscus described by OBERI)ORFER (1.C.) seem to be related.
The community described here, however, includes a fairly large
number of species of the S a l i c o r n i o n f r u t i c o s a e and the
P h r a g m i t i o n . The species belonging to the latter are badly
developed. For these reasons the syntaxonomic position of these
I32 W.J. WOLFF
communities becomes very difficult to determine. Indeed the milieu

is nearly unique. Just before our second visit these vegetations had
been flooded and afterwards burnt by heavy sunshine, so we could
not make further observations in 1964 .
Phragmitetea Tx. 1942
Phragmitetalia (KocI-I 1926 ) Tx. 1942
Phragmition BR.-BL. 193 I
Scirpetum maritimi et littoralis BR.-BL. 1931 TABLE 2: 1-22.
Syn.: Ass. a Scirpus littoralis PmNATTI I953
In places with more or less brackish water, e.g. in ditches in the
vicinity of the lagoons and along creeks with fresh water running
into the lagoons, a vegetation of Scirpus maritimus, S. litoralis and
Typha cf. angustata occurs. Although the observed vegetation is
rather poor in species, it belongs undoubtedly to the S c i r p e t u m
m a r i t i m i et l i t t o r a l i s . This association is known from southern
France, Italy, Dalmatia (BRAUN-BLANQUETC.S. 1952 ) and Egypt
(TADROS • ATTA 1958).
Consociation belonging to the alliance Phragmition.
Consociation of Scirpus maritimus TABLE 2: 23-24.
In one place where an influx of fresh water occurred in a S a l i -
c o r n i e t u m f r u t i c o s a e a vegetation of Scirpus maritimus with only
Salicornia fruticosa as a companion was observed. The individuals
of Scirpus were well developed, those of Salicornia more or less
poorly so.
This vegetation was situated at a greater distance from the lagoon
than the sociation of Juncus acutus mentioned on page 13o , so we
suppose that the salinity and the fluctuations in the salinity will be
less in the vegetation described here.
Owing to the poor development of the Salicornias we regard this
vegetation as a consociation of the P h r a g m i t i o n .
SuccESSION
When considering succession it is very useful to bear in mind the
difference between allogenic and autogenic succession as distin-
guished by TANSLEY (1928). As we do not have data on succession
we have to derive our conclusions from observations on the pattern
of vegetation. However, when doing this, one has to be very careful
in distinguishing between zonation and succession.
From our observations we have indications of only one instance
of autogenic succession anywhere in the saltmarshes studied. O n
page 125 we already mentioned this case of autogenic succession
from the S u a e d o - K o c h i e t u m h i r s u t a e to the subassociation
Suaedo-Kochietum hirsutae rumecetosum.

This does not mean that this phenomenon does not occur more
often, but it is at least fairly rare.
Allogenic succession is more common but one should be careful
when judging it. It seems probable that in many instances a pat-
tern of vegetation which seems to be the result of allogenic succes-
sion, is in fact the result of a contemporary formation of the soil-
profile. Therefore we think that the differentiating into e.g. S ali-
c o r n i e t u m r a d i c a n t i s and S a l i c o r n i e t u m f r u t i c o s a e is not
a result of allogenic succession but of a contemporary sedimenta-
tion of different soil-profiles. The same seems true for the differ-
ences between S a l i c o r n i o - H a l o c n e m e t u m s t r o b i l a c e i and
Arthrocnemetum and for the chain: sociation of Phragmites
communis-communities of Juncus maritimus and J. acutus (J u n c e t u m
maritimi)-Salicornietum fruticosae- Salicornietum radi-
cantis-Arthrocnemetum-Salicornio-Halocnemetum stro-
b i l a c e i as figured in fig. 2.
The only instances of true allogenic succession seem to be the suc-
cession from S a l o c o r n i e t u m f r u t i c o s a e t y p i c u m into Sali-
c o r n i e t u m f r u t i c o s a e h a l i m i o n e t o s u m (page 112) and from
Arthrocnemetum into B e t o - M a t t h i o l e t u m tricuspidatae
(page I27).
VEGETATION MAPPING
The major reason for the mapping of the vegetation of the salt-
marshes near Mesolonghi was to give an objective impression of
the importance of the area for nature-conservancy. For this reason
the vegetation map with comments on it is not comprised in this
botanical study. Except for the aquatic vegetations all communities
mentioned in this study are mapped. The map is filed in the
library of the Hydrobiologisch Instituut afd. Delta-onderzoek,
Yerseke, Holland.
REFERENCES
BEEFTINK, W. G. 1962 - - Conspectus of the phanerogamic salt plant commu-
nities in the Netherlands. Biol. Jaarb. Dodonaea 3o: 325--362.
BEEFTINK, W. G. 1965 - - De zoutvegetaties van ZW-Nederland beschouwd in
Europees verband. Dutch with English summary. Meded. Landbouwhoge-
school Wageningen65--1 : I67 pp.
BRAUN-BLANQUET,J., N. ROOSSlNE& R. Ns 1952 - - Les groupements v6g6-
taux de la France M6diterran6enne. Centre Nat. Rech. Scient. 297 PP-
B~AuN-BLANQUET,J. i95I - - Pflanzensoziologie. Grundztige der Vegetations-
kunde. 2. Aufl. Springer, Wien.
EIG, A. 1946 - - Synopsis of the phytosociological units of Palestine. Palestine J.
Bot., Jerusalem Set. 3: 183--248.
ELL~NBERG, H. 1956 - - Aufgaben und Methoden der Vegetationskunde. Ulmer,
Stuttgart. I36 pp.
134 W.J. WOLFF
FREI, 1VI. 1937 - - Studi fitosociologici su alcune associazione littorali in Sicilia

(Ammophiletalia e Salicornietalia) Nuove Giorn. Bot. ital. NS 44: 273--294.
HARTOO, C. DEN & S. SEOAL 1 9 6 4 - A new classification of the waterplant com-
munities. Acta Bot. Neerl. 13:367--393 .
HATZIKAKIDES, A. D. I952a - - Recherches saisonni6res hydrologiques des la-
gunes de Messolonghi et d'Aitolikon. Greek with French summary. Prak-
tika Hellenic Hydrobiol. Inst. 6: 85--144.
HATZlKAKIDES,A. D. I952b - - Rougissement periodique des eaux de la lagune
d'Aitolikon. Greek with French summary. Praktika Hellenic Hydrobiol. Inst.
6: 21--54.
HAYEK, A. I927--I933 - - Prodromus florae peninsulae Balcanicae. Dahlem.
HOgVATIC, S. 1934 - - Flora und Vegetation der nord-adriatischen Insel Pag.
Bull. int. Acad. Yougoslave Sei. Arts 28:
OBERDORFER, E. 1952 --Beitrag zur Kenntnis der nord~ig~iisehen Kiistenvege-
tafion. Vegetatio 3: 329--349.
PIGNATTI S. 1953 - - Introduzione allo studio fitosociologico della pianm'e vene-
ta orientale. Atti Ist. Bot. Univ. Lab. Crittogamico Pavia Serie 5, 9:92--258.
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ser. 7:
R~.Nz, C. I 9 5 5 - Die vorneogene Stratigraphie der normalsediment~iren For-
mationen Griechenlands. Athens.
S~GAL, S. I965 - - Een vegetatieonderzoek van de hogere waterplanten in
Nederland. Wetensch. Meded. K N N V 57. 80 pp. Dutch with English sum-
mary,
SI~GAL, S. & J. J. BARICMANI960 - - Enige opmerkingen over abundantie en do-
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Wolff - The Halophylous Vegetation

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THE HALOPHILOUS VEGETATION OF THE LAGOONS

Salt-marshes cover extensive areas of the mediterranean region.

THE PRESENT INVESTIGATION

To obtain a clearer picture of the east-mediterranean halophi-

saly, Macedonia and Thrace. The lagoons of Mesolonghi were

Hordeum marinum HuDs. Puccinelliaf estucif ormis

;" / - HAG ~.G. "~IA~IA

SITUATION AND TOPOGRAPHY OF THE AREA

The lagoons of Mesolonghi are situated in the province of Aito-

The climate of the studied areas is typically mediterranean with

GEOLOGY AND GENESIS

The Pindos range and some neighbouring chains form a large

Meteorological data from Mesolonghi, derived from HATZmAKm~S I952a.

Mean number of days Mean temperature of the

January 12.6 9.9

likon (North of Aitolikon) differs considerably from the rest (HAT-

The vegetations of the L i m o n i e t a l i a , the S a l i c o r n i e t a l i a

DESCRIPTION AND CLASSIFICATIONOF THE VEGETATION

Zosteretea (PIGNATTI I953) DEN HARTIG @g SEGAL I964

Zosterion (CHRISTL~NSEN I934) DEN HARTOG & SEGAL

Ruppietea (J. T/dXEN I960 ) DEN HARTOG & SEGAL I964

Ruppietum spiralis IVERSEN 1936

Potametea Tx. & PRSO. 1942

Characteristic species combination

Characteristic species combination Ranunadetum baudotii:

Characteristic species of the Parvopotamion:

Well developed vegetations occur in stagnant or slowly running

-- -- 2m.3--4 2a,2 +.t 1.2 • -- +.~ I°-2 ~m.5 5"5 5-5

xoo% 95% xoo% ~oo% 5o% 3o% 5% 45% 75% 2o% ? . . . .

5.5 1.2 1.5 -- +.2

3.5 -- -- 2b.3 4.5 2a'3 ] . . . . .

1,2 +.o -- 5,5 I ~-.I

ditches (lO--3o cm) with in summer slowly running fresh water

Thero-Salicornietea (Tx. 1954) Tx. & OBERD. 1958

and the Ruppia-vegetations a vegetation occurs consisting nearly ex-

Cyanophgceae 5.5 . . . . 5.5 . . . . 5.5 -- --

Charaeterlsti¢ species Salieornietum radicantis:

Differential species of the dry variant:

Characteristic species of the Limonietalia:

Characteristic species combination

O c c u r r i n g i n r e c o r d 78: P u c c i n e I l i a c o n v o l u t a t . ~ ; 79: t M c c i n e l l i a c o n v o l u t a + . 2 , A r t h r o c n e m u m g l a u c u m + . 2 ; 82: S c i r p u s m a r l t i m u s

+.2 1.2 3"3 1.2 +o.i +.2 . . . . . . . . .

-- -- x.t -- +.x x.t +.1 2a,2 1.I +.I +.~,

- - - - - - r -- +.i 1.2 . . . . . . +.i --

+.2 -- +.2 +.1 ~ -- +.t +.t +.x +,2 +.i -- -- +.1 --

+,i -- -- i,i +.I -- -- -- +.x . . . . +.x --

-- -- -- +.2 2a.3 +.2 --

-- +.~ -- +.x 1.2 2a.3 3.4 -- x.~ --

. . . . . . . . . . . . . 2a.2 2b.3 i2b.3

-- -- +.r . . . . . . r.~ -- +.2 . . . .

Characteristic species Sallcornietum radicantls:

Characteristic species of the Salicornietalia fruticosae:

Aeluropus litoralis 1,2 I,I 1,2 ~*I --

getation in which only a few individuals of Puccinelliafestuciformis,

is the fact that Puccinelliafestuciformis is optimally developed on and

Limonietalia BR-BL. & DE BOLOS 1957

Nr. 96 97 98 99 ioo IOi IO2 IO3 Io4 zo5 xo6 lO7

Characteristic species Salicornio-

LAGOON' BARE ISALICORNIO- ,IJ SALICORNIE-{'JUNCUS PHRAG

nemetum are lacking in the S a l i c o r n i o - H a l o c n e m e t u m

discover one instance of an A r t h r o c n e m e t u m near Arta and none

Characteristic species Salieorn:eto-