Académique Documents
Professionnel Documents
Culture Documents
OF MESOLONGHI, GREECE
by
W. J. WOLFF
Yerseke
INTRODUCTION
PREVIOUS STUDIES ON MEDITERRANEAN SALT VEGETATIONS
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
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
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
CLIMATE
TABLE I
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
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
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
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
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
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.
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
1.3 . . . . 2a.3 --
+.2 -- -- +.2 -- -- --
I°.5 I~.5
+.2 - - 2m. 5 -- --
- - 3 . 4 - - + . 2 - -
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 - - - - - -
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 ?
Hordeum marlnum m
Juneus ambiguus
Centaurium spicatum m u
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 ° ~. ?
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 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
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
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
-- -- +.2 -- -- +.2 . . . . . . . . . .
+.2 +.2 +.~ +.2 -- +.x -- -- +.~ i.o :t.2 z,~ 1.2 z.2 +.2 x.2
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
Companions:
Salieornia herbacea s.I. +,I JF.I l,I -- I.I
Juncus aeutus
Phragmites eommunis
TABLE 6.
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
! ,,:> 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.
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
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
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.
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
Local characteristicspecies:
Triglochin bulbosum . . . . . . I. x -- -- -- 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
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
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
-- -- 1.2 . . . . . m -- +.2
. . . . . . . . +.2 -- +.2
m -- +,I +.2
m -- +.I
-- +.2
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
Characteristic species of the
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 - -
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
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 - -
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 .
HALOPHILOUS VEGETATION OF MESOLONGHI 123
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-
Companions:
Sallcornia radleans r.~
Salicornia herbacea s.1. +. I
Halimione portulacoides 3.5 x.2 - -
Statice limoniurn ssp. serotinum + . I - - - -
Phragmites communis +. I
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
HALOPHILOUS VEGETATION OF MESOLONGHI 125
TABLE I O.
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.
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.
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
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
Characteristic species of the
Cakiletea maritimae:
Cakile m a r i t i m a . . . . . . . - - - - + . 2 - - - -
Nitrophitous spades:
Atriplex cf. rosea . . . . . . . +.i +.t +.x -- - -
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 . . . .
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.
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
HALOPHILOUS VEGETATION OF MESOLONGHI 131
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 - - - - - -
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 . . . . . . . + __ __
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 .
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
HALOPHILOUS VEGETATION OF MESOLONGHI 133
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