Acta Bot. Neerl.
25(4), August 1976,
Palynology
Northern
T.A.
Hugo
Wijmstra
de
the
of
the
of
metres)
p. 297-312
120
Greece
and
A.
middle
m
part
(30–78
in
section
deep
(Macedonia)
Smit
Vries-Laboratorium,
Afd.
Palynologie,
Universiteit
van Amsterdam
SUMMARY
In this
bore
the results
paper
hole
in
the
and
steppe phases
14
dated last
C
are
interglacials
that
might
be
the
upper
30
of the
here
given
boring
m,
pollen
an
from
interval
m
as
a
from
reflects
character.
evergreen
steppe)
and the
oaks indicates
of only
model,
the
steppe phases mostly
warmer
(semi)
that
periods
deciduous
van
der
with
oak
grains
of
prominent
pollen
of the
Upper
oak
a
deep
alternating
Taking
oak
as
the
phases
glacials.
might
be called
species
indicates
vegetation
s.l.) and
were
of the
chiefly
obtained
electron
120
and
the last
collected
pollen,
as
by micromanipulator
well
as
from
were
presented
interglacial
observation.
from
samples
showing differences
in
studied with the aid of
microscope (S.E.M.).
Smit
pollen grains
(1973)
of
were
used
to
distinguish
3
major types
within
Quercus:
as
Q. pubes-
Q. robur, Q. petraea.
Quercus
and
in
(1975).
Quercus robur/petraea type, comprising deciduous species such
cens,
(1969),
reported
m was
by light-microscopical
light microscopical observation,
given by
Wijmstra
will be
described and discussed in Wijmstra & Smit
were
values for oak
Criteria
given by
1971,
the
(from
detailed description
a
complete
Zagwijn,
work
pollenanalytical
Earlier results
underlying deposits (78-120 m)
Quercus species,
under
Pleistocene
generalized diagram
was
scanning
the fossil
the
Hammen, Wijmstra &
The present data
Pollen
from the continuation of the
climate and
recent
analysis
from this section
c.
forest and
Tenagi Philippon plain {fig. 1).
representing
separate paper. A
b.
of them with
interstadials
30-78
pollen diagram
interstadials.
are
from the
its
area,
whereas the
a.
some
evergreen
presence
called
The
presented.
INTRODUCTION
on
a
forest,
or warm
from
of pollen
whereas the
The results
in
are
interglacial-glacial cycle (resp.
presence
periods
a
of oak
phases
of the
palynological investigation
interpreted as interglacials
The
1.
of a
Tenagi Philippon
ilex/coccifera
type,
which
includes
the
evergreen
species
Q.
ilex
Q. coccifera (Q. calliprinos).
Quercus
suber type,
comprising Q. cerris, Q. macrolepis, Q. trojana,
mainly semi-evergreen species,
as
well
as
the
evergreen
and other
species Q. suber.
298
1.
Fig.
A
Location
of these types
pollen grain
This division
THE
2.1
proved,
considered
opinion
to
be
collected per
absent,
SMIT
found
being
not
sample.
of the present authors,
of the
interpretation
pollen diagram and
to
be of consider-
the reconstruction
cover.
DIAGRAM
(fig. 2, Appendix
the results of the
diagram
120
m
deep section
with KOH
mainly
and,
after
corroded. The
pollen
types
phaeaceae,
pollen
in
total used
either
pollen
The
gyttja, clay
pollen grains
except
Utricularia,
a
palynological
presented.
pocket
page 3
on
cover)
Peplis
to
sum
open
as
a
of the 30-78
analysis
samples
were
were
to
be
diagram
Cyperaceae,
Typhaceae,
and
since
aquatic vegetation
of 500
pollen
was
these
or
a
to
part of
were
cm
boiled
bromoform-
well-preserved
basis for the main
Rumex
samples
concentrated with
found
m
every 20
analysed
and lake marl. All
acetolysis, subsequently
observed
speaking, belong
samples
are
of peat,
alcohol mixture. The
2.2
in the
vegetation
POLLEN
and consisted
all
A.
General remarks
In the
the
was
Quercus pollen grains
able value for the
of the former
WIJMSTRA AND
Tenagi Philippon.
among 50-100
2.
A.
T.
and
not
consisted of all
Menyanthes,
elements,
hygroseres.
Nym-
generally
In
nearly
taken.
The zonation
The sequence under discussion
can
be subdivided into
a
number of
pollen
zones.
PALYNOLOGY
For the
pollen
120
THE
OF
major pollen
smaller units
cluded also
zones
in
NORTHERN
299
GREECE
the dominance of either the herbaceous
cases
this
In these
(subzones).
variations
Pinus
or
IN
DEEP SECTION
used. In several
was
Quercus
M
major
criteria used for
the
cases
separate pollen
the
or
tree
zonation could be subdivided into
such
curves
subdivision in-
for
as,
the
instance,
curve.
Zone VV
This
zone
in this
is characterized
by
its
high percentages of non
the dominance of grasses, Artemisia and
case
transition from this
to
zone
PP is
in
present
arboreal pollen
Chenopodiaceae.
(NAP),
Only
the
diagram.
our
Zone PP
This
shows
zone
of arboreal
high percentages
very
and Pinus. Identification under S.E.M. observation
Q.
suber type and the
separate
Subzone
excelsior
the
The
oak
first
subzone PP2
can
15,
curve
and
high
the
most
Plantago
is of
Quercus
are
In subzone PP4 the
shows
Quercus
curve
subzone
Ericaceae
also found in this
Quercus pollen
Tilia
and Fraxinus
is worth
mentioning.
Abies is present,
Ulmus
starts
are
of
reaching
pine pollen
and
In
zone
of Bruckenthalia
grains
some
shows
percentage of up
a
conspicuously represented.
rise
to
curve
curve
over
zone
a
the
rise
pine
still
found
pine
diaceae. The
upper part
a
curves
pollen
abundant.
continuous
to
be present
of subzone
curve
percentages
a
high although
the lower part
In
and
Carpinus
In this
zone
the
of this
Fraxinus
are
all three types of
(S.E.M. observation).
PPS
for the
can
curve.
still
are
30%.
the dominance of the
is
sample
at
70.2
m).
Quercus
In the upper half of
be noticed.
Tilia, Ostrya/ Carpinus
curves.
percentages is caused by
a
(except
Abies,
continuous, closed
relatively
with
curve
in the
In this subzone
of Pinus
values between 25 and
are
low
constant
importance.
percentages
zone
were
maxima of Pinus and the
characteristic facts. The continuous
presence of Erica-
The main characteristic
the
for the
maximum of the
a
Ulmus/Zelkova
PP2b the
during part
for Ericaceae
In subzone PP3 the
ceae
curves
found.
were
of
of the
be divided into PP2a and PP2b. In the part PP2a
and
Carpinus
help
Quercus
In the second half of this sub-
important.
more
PP2
Throughout
percentages of
Carpinus pollen
dominant while
while also
PP2b the
the
becoming
of
values.
of oak and
high percentages
subzone
are
With the
of
the presence of the
proved
be subdivided into 5 subzones.
can
occurrence
pollen is
higher
to
this
pollen
next
Q. robur/petraea type.
zone
PPI shows
During
curve.
zone
this
trees
pollen, mainly
orientalis
In the middle of this subzone
are
a
represented
rise
by
in the N.A.P.
greater contribution of Artemisia and of Chenopo-
for Salix and Pistacia show maxima in this interval. In the
marked rise in the
curve
for
Nymphaea
is
noteworthy.
Zone XX
In this
zone
the N.A.P. percentages
are
higher
than the A.P. percentages. This
300
T.
is caused by
rise
a
to
this
of
A division
zone.
some
for Artemisia
curves
the
grains
Tamarix,
be
can
Nerium oleander and
made into
whereas the Thalictrum
excelsior. Ulmus,
curve
is
from XXI
wed
by
Zone
This
a
XX2
to
a
rise in the
maximum of the
restricted
up
to
resp.
and
30%
Myrtus
are
10%.
present,
constantly
high percentages
Cyperaceae
curve
be
can
of monolete fern
curve
curve.
seen,
of Salix
At the transition
and follo-
preceded
spores.
YY
zone
shows
dominant while
Poterium
a
a
Buxus and Nerium
pollen.
tree
for
In
the lower part
Salix
found. The
some
grains
curve
and also
is
some
grains
of
be divided into 2 sub-
zone can
of Hippophaë
were
observed and
of Arceuthobium.
In the upper part of zone YY2 the
of Abies and Ericaceae with
presence
is worth
percentages
of the
oak
Pistacia and
Tilia, Carpinus, Fraxinus,
rise in the
were
At the base of subzone Y Y1
grain
of
high percentage
higher percentages
found. There is
are
Pterocarya,
zones.
one
are
Pistacia and
and shows well-marked maxima of the grass
Typha
and grasses.
are
continuous.
The upper part of subzone XX2 shows
and
Cressa
SMIT
subzones, XXI and XX2. In
two
and oak
percentages of pine
of Fraxinus
WIJMSTRA AND A.
Chenopodiaceae
,
subzone XXI
Also
in the
grains
Some scattered
A.
zone
followed
There is
noting.
by
a
centages of Cyperaceae
a
maximumof
are
maximum of the
pine
curve
this
Quercus. Throughout
higher
in this part
zone
the
per-
high.
Zone ZZ
in the values of Artemisia and
Here the rise
dominance of N.A.P.,
In this
zone,
although
among the
some
pine pollen,
Chenopodiaceae
Quercus
some
and
grains
pine
are
cause
pollen
a
marked
is still found.
very similar
to
those of
Pinus heldreichii.
Zone A
This
pollen
Fraxinus
zone
are
A la may be
In the
shows
present
high percentages
with low
distinguished,
next
interval,
A
in which
lb,
of arboreal
pollen.
percentages throughout
pine pollen
Pinus and oak
is
pollen
this
Abies
Ulmus and
,
zone.
An
interval
dominant.
is present in
approximately
equal percentages.
In subzone A2 the percentages of
this subzone is
a
marked but
pine
are
constantly high.
short rise of the
Quercus
In the middle of
curve.
Zone B
Here the N.A.P. percentages
maximum of the arboreal
the N.A.P.
ages of
Chenopodiaceae.
are
are
caused
in the middle part
generally high again; only
pollen (A.P.)
by
an
can
be observed.
increase
The
a
higher percent-
of the values for Artemisia and
In the lower part the grass percentage is
slightly
lower than
in the upper part.
In the lower
part
the percentage for
Nymphaea
shows
an
increase,
while in
PALYNOLOGY
120
THE
OF
the upper part
M DEEP
attains
Typha
IN
SECTION
NORTHERN
values.
higher
3 subzones. Subzones B1 and B3 show
and
pollen
B3
Pollen
B
zone
curves
values for
higher
pine
increase of the percentages of Cyperaceae
an
be divided into
can
maximum of the
subzone B2 shows
Chenopodiaceae,
zone
a
301
GREECE
of Artemisia
and
oak. In
is present.
Zone C
This
zone
zone
Cl
shows again
oak
is
pollen
high
values for the
dominating,
tree
increase in the percentages of Tilia and Abies is worth
for
and Ulmus. Fraxinus is
Carpinus
the
(C2),
centage becomes increasingly higher. At the transition from Cl
case
sub-
pollen percentage. During
while in the second half
to
pine
C2
per-
slight
a
This is also the
noting.
in zone C.
continuously present
Zone D
In this
zone
the dominanceof N.A.P. is caused
centage of Artemisia and Chenopodiaceae.
maximum of the
curve
for
Typha
is observed,
At the end and the
Cyperaceae.
of grasses is
curve
by
a
marked increase in the
In the middle of
pollen
and followed
preceded
of this
beginning
zone
by
a
zone
perD
a
rise in the
the percentage
high.
Zone E
The main features of this
the
pine pollen
psilate
curve.
for
an
higher
is
a
the transition
Typha
be
can
high,
to
ratio of about 1 and
A.P./N.A.P.
monolete spores shows
pollen percentage
near
zone are
The oak percentages attain
whereas
zone
F.
values.
15-20%.
The
for the
the
grass
marked increase of these values takes
place
Throughout
Just after the grass
noticed. Ulmus is
this
rise in
a
curve
increased values
maximum,
this
present throughout
zone
zone.
Zone F
The main characteristics of this
and the presence of Abies,
part of this
zone
preceded by
zone
an
an
the
constantly high percentage
increase in the percentage of grass
the
for
curve
creasing values for Typha,
Cyperaceae
of oak
Salix and Betula. In the
increase of the Alnus percentage
maximum in
a
zone are
Tilia, Juniperus, Ulmus,
can
pollen.
can
be
be
noticed,
upper
which is
In the last part of this
seen
together
with
in-
Filices and Salix.
Zone G
Maxima in
the
predominance
Throughout
curves
for
Artemisia,
Chenopodiaceae
and
grasses
cause
a
of the N.A.P. values.
this
lower part of this
pollen
zone
zone
there is
maxima in the fern
a
maximum in the
curve can
Cyperaceae
be observed. In the
curve.
Zone H
This part of the sequence is characterized by its
mainly
The
from
zone
higher percentage of
tree
pollen,
Quercus.
can
be divided into three
subzones,
HI, H2, H3. During subzone
302
Fig.
A.
T.
3.
Surface
details
of fossil
a.
Quercus robur/petraea type,
b.
Quercus
c.
Quercus ilex/coccifera type,
suber
type,
20.000
x
H1 all three oak types
Carpinus and
higher part
At
the
Salix is
values
Ulmus
of the
to
20.000
47
m).
47
m).
m).
(depth
present
were
H2,
while oak
observed,
Abies
During pollen
followed
by
a
pollen
maximum of Alnus
After this Alnus maximum
At the end of this
period
a
top in the
of N.A.P.
pollen (mainly
upper part of this
Juniperus
curve
Artemisia and
zone.
At the
H2
a
a
remains
we
in the
curves
of
middle and
more
than
10%
marked rise
an
see at
in
and
also
the N.A.P.
important component of
first
a
rise of
up
to
18%
of
(c. 43%).
curves
dominance,
become higher. The subzone H3 shows
N.A.P.
Pistacia. The
as
especially
reaches
zone
the total of arboreal pollen. In this subzone
Salix,
well
as
reaching higher values,
subzone
here.
increasing
be
(depth
47
(depth
(see fig. 3)
are
SMIT
pollen.
20.000
x
A.
zone.
transition
can
oak
x
WIJMSTRA AND
of the ferns and grasses
the
rise in the
percentages
pine
curve,
Chenopodiaceae)
lower half of the
of
occurs.
Cyperaceae
the percentage of
increases towards
the
maximum in
the
sequence
a
is present.
Zone K
The
the
high percentages
N.A.P. of up
maximum of the
a
to
of Artemisia and
90%
Nymphaea
At
the
curve
Chenopodiaceae
boundary
occurs,
cause
between this
a
zone
dominance of
and
zone
L
a
while in the middle part Typha attains
higher percentage.
Zone L
The
zone
can
be divided into 5 subzones.
During
the
first four subzones the
percentage for both pine and oak is somewhat higher, although
Artemisia and
PALYNOLOGY
shows
four maxima
Subzone L5
is
of pine and oak.
an
SECTION
At 35
m
one
by
in the
a
303
GREECE
constituents of the
in each of subzones
very
zone
a
rain. The pine
pollen
LI, 2,
high percentage
shows
spectrum
this
Throughout
increase
IN NORTHERN
major
(one
characterized
Chenopodiaceae.
upper part
M DEEP
remain the
Chenopodiaceae
curve
120
THE
OF
3 and
4).
of Artemisia and of
sudden rise in the percentage
the percentage of grasses is
percentage
of
Typha
be
can
high.
In the
observed.
Zone M
This
is
zone
described earlier in
was
Wijmstra &
1975.
Smit,
subzones. In subzone M1 the percentage of Artemisia and
two
still
The
high.
(semi-evergreen
pollen frequency
major constituent.
Xantium,
There is
a
is much
zone
distinguish
is
Gypsophylla
mainly caused by
an
increase
type).
in subzone
higher
M2, Quercus being
consistent representation of Pistacia,
Castanea. The percentage of Artemisia and of
than in
and
Pleurospermum
the A.P. percentage rises,
of the oak percentage
The A.P.
of
presence
noteworthy. However,
We
Chenopodiaceae
the
Ulmus and
is much lower
Chenopodiaceae
M1.
Zone O
This
of
Ericaceae
of
high
a
arboreal
occurs.
content
pollen
a
1975. A continuous
Smit,
presence
by
a
subzone 02
the
curves
of
Carpinus
Abies and
Ostrya
of
start to
of
curves
by
the
rise.
with
Carpinus
curve
zone
maximum in the
and
continuous representation
this
throughout
Tilia, by the continuous high percentage of
low percentage of Ericaceae. The
In the
and
The subzone Ol is characterized
and of
Quercus
in Wijmstra &
already described
zone was
Abies,
a
high
percentage and the presence of Rhododendronare worth noticing. In the interval
02a there is
a
lowering
Tilia and Fraxinus
pollen
The
Quercus
zone
is apparent. Here also the
curve
Zone
of the
In the second half of
curve.
in
zone
O
ornus are
belongs
to
no
O
curve
accompanied by
(interval 2b)
curves
a
of Betula and Alnus
longer represented.
a
rise in the Pinus
maximum in the
The
the deciduous type ( Quercus
start
majority
Quercus
rising
whereas
of the
Quercus
robur/petraea type).
Q
beginning
Quercus,
the
percentage
of this
latter
of the
increasing again.
is marked
by
zone
is marked
mainly consisting
by
rise of
a
of the
herbaceous elements
The
of this
upper part
like
zone
the presence of the evergreen
as
pine pollen
and
semi-evergreen
Artemisia
(see also
well
as
a
decrease of
type. The pollen
and
Gramineae
Wijmstra &
are
Smit,
1975),
oak
pollen
semi-evergreen
type.
3.
THE
Using
on
a
DEVELOPMENT
the
A.P./N.A.P.
physiognomic
OF
VEGETATION
relation
a
basis may be
(table 1)
reconstruction of the local
given.
As
discussed in
vegetation types
Wijmstra
(1969),
a
Table
1.
Reconstruction
of the
Middle
Pleistocene
vegetation
donia.
mtr
Q
30-
O
oak
a
oak
o
1
onlower
forest
z
oaks
with
slopes.more upwards Abes forest
forest with
open oak
2
M
evergreen
Carpmus betulus
with
oak forest
o
2
and
forest with Carpmus betulus
oak forest
b
open
dry steppe w.th
very
Pistacia
Artemisia.
Chenopodiaceae
35-
5
Artemisia,
steppe with
Chenopodiaceae
4
L
P-
NEAKTERIN
2
alternating steppe and
and
pine).
AO-
K
steppe with
3
oak
forest
locally
O
_j
2
H
>
.Chenopodiaceae
Artemisia
and
forest steppe with pine
z
O
oak
forest steppe (with
1
of
45
as in
shore
a
m theupperpart
oak
forest
, but
zone
stands
is present.
of this
sub
replaced by
is
with
vegetation
alder and willcw
Jumperus.
the
zone
open forest type
a more
with
to
with
forest
1
and
Carpmus
and deciduous
evergreen
Abies,
continental forest
theupper
in
with only
part
at
oaks,
of
deciduous
higher parts
this sub zone a more
oak
species
steppe with Artemisia .Chenopodiaceae
G
50
oak
deciduous
forest
with
pine
some
F
(Quercus
AXIOS
like
cerns.confertae
E
forest steppe with stands of
D
steppe with
association)
oak
and pine
I
Artemisia
.Chenopodiaceae
and
grasses
<
2
t
pine forest with
oak
deciduous
_5C
1
(Quercus
oaks
some
with
forest
some
pine
cerns.confertae like association)
3_
!
B
stands
forest steppe with
2
of
pine and scattered oaks
1
to
2
UJ
A
LU
mixed
60
b
oak
a
pine
pine. oak
forest
forest.
with deciduous
oak species and
pine
?
cc
z
zz
forest.
steppe
1
$
Art Chenop
decid
oak
and
grasses
species and
Salix
Abies
wet
on
sites
at
the higher locations
deciduous submediterean
1
t
with
with
forest
2
65
Buxus
oak
forest
steppe with Artemisia, Chenop
2
with
and Pistacia
,
shore vegetation
with willow and Tamarix
—
XX
■
steppe
elements
with
forest
5
higher
oak
with
in
the
composition
deciduous
oaks
and Acer
lower
and in
species
Fraxinus
z
oaks
b
with
the
at lower
sites
Pistacia
parts evergreen
upper
part
a
mixed
arboreous
in
Quercus, Carpmus betulus,
and Ulmus
dominance of Pmus and
some
and an
forest
Pmus,
exelsior
with
forest
3
undergrowth
which
of
scattered
Ericaceae
oak.Tilia.Carpmus.Ostrya
and
Fraxinus
are
75-
replaced by
at
a
forest
with
location
steppe
locally
an
Abies
of
forest
Ericaceae is
developing,
is
present
(submed ) deciduous oaks,Carpmus betulus,
Fraxinus,Tilia.Ulmus
78-
undergrowth
pine, an
the higher
1
vv
and some
■
to
2
a mixed
location Abies and
vegetation
_J
pine and oak
Artemisia,Chenopodiaceae and grasses
Tilia.Ostrya, R-axmus
forest
<
with
70at
4
like
:
forest
1
with
Salix
and
Pistacia.
Artemisia, Grammeae,
bushes
on
wetter
Chenopodiaceae
sites
and
sequence
in
Eastern
Mace-
PALYNOLOGY
120
OF THE
M
subdivision in
major
DEEP
and
steppe”
following values,
305
GREECE
as
“steppe”
be
can
made.
derived from Frenzel (1968),
used:
are
AP
forest
55%:
30%
AP
AP
55%:
forest steppe
30%: steppe
This physiognomic
Mediterranean
Amor
the
subdivision
proposed
area
Bonatti
(1962),
is comparable
Beug
by
of
presence
Using
have
from
pollen
in the
as
criteria,
zones
IIX, B, E and L had, accordingly,
zones
(“Waldsteppe”)
the characteristic
as
XX2, ZZ, D, G, K and L5
must
Henendez
Niklewski &
evergreen type
an
during
area
Florschutz &
and
(1974)
can
be
Zeist
van
a
closed forest
a
vegetation
seems
PP, YY, A, C, F, H and O/N. The
rather
open
forest
have been characterized
by
forest steppe
or a
the
vegetation type. Finally,
an
steppe
open
of each
zone are
registered.
A
detailed
more
description
VV,
zones
tation. In table I these major types and the variations in the composition
vegetation
due
distinguished
evergreen oaks.
the indicative percentages
prevailed
with other divisions for the
(1967a),
Bottema
(1966),
Within the forest formations
(1970).
to
NORTHERN
“forest
“forest”,
As indicative percentages the
to
IN
SECTION
is
vegeof the
presented
here.
3.1
Steppe vegetation
During
zone
ceae must
VV
have
local stands of
X5,
zones
Wijmstra
vegetation might
X3
(1969).
D, G,
zone
14
C
comparable
dating
Rosenberg
a
a
growing
regular
at
the
Tamarix and Pistacia
ported by
of the
a
1969),
as
contents
well
vegetation type
cold and
same
as
the
to
a
occurrence
points
the fact that
a
to
cold
zones
an
open vege-
is caused
high
very
mainly by
percentages,
lacustrine environment with
lower and
wetter
dry
ZZ,
dry phases.
of Tamarix and Pistacia. This may
desiccating
remaining
attains
by
(Verg-
cold and
but also of
dominance, however,
Salix
cores
of Ceratoides
presence
and
described
as
deep-sea
only VV,
The
area.
described
climate phase
dry
boring
1970) clearly point
of not
Chenopodia-
parts of the
dominancein the AP percentages suggests
probably seasonally
Tamarix
found in
Smit & Wijmstra
grass percentages, whereas
higher
the
is, therefore, indicative of similar
XX2
combinedwith the
to
was
to
wetter
and correlation with
tation of the usual steppe type. This
the
with Artemisia and
occupied
higher parts
The similar pollen
K and L5
During
in the
There
(= Krascheninnikovia)
period.
be
This steppe type
etc.)
naud-Grazzini &
climate
whereas Salix
occurred,
by Wang (1961:203).
(the
vegetation
open steppe
an
sites.
The
point
Salix
and
occurrence
of
somewhat raised temperature which is
sup-
a
steppe element
as
Ceratoides could
not
be
esta-
blished.
The
pollen
rain of the
rain that fell
during parts
“interpleniglacial”
1969). Therefore,
the
vegetation
must
scattered stands of Pinus. In these
occurrence,
which
points
to
of zone L is
comparable
of the Last Glacial of the
less
also
patches
dry
have
of
same
been of
forest,
oak
with the
section
an
was
pollen
(Wijmstra
open type
of
more
with
frequent
and somewhat warmer circumstances.
306
T.
3.2
PP the
zone
with
corresponds
forest
WIJMSTRA
AND
A.
SMIT
vegetation
Forest
During
A.
composed
allow
pollen assemblages
a
subdivision of which each
part
type of forest. Zone
particular
a
of oaks of the
PP1
Quercus robur/petraea
and other non-mediterraneanspecies) and also of
is
characterized
group
(probably Q.
of the
species
suber
Q.
by
a
robur
group
(probably Q. cerris).
In this forest also
Carpinus
can
betulus
Ulmus and Fraxinus occurred. Above this forest
Tilia,
belt
developed
therefore be considered
In
PP2a
zone
the
on
have been
to
As
slopes.
a more
further differentiationtook
a
compared
humid and
the altitudes of the forest belts. On the lower parts of the slopes
was
an
gradually replaced
Abies of the
A.
and
with
type,
raise
a
fores types
Quezel
stands of oaks
in the
probably
of
subzone PP4
In the
Subzone PP5 has
a
our
of
of the Pinus
increasing
discussion of
dryness
comparable
(1973).
but
area,
scattered
some
evergreen oak
an
not
increasing dryness.
present
that time,
at
at
A clear
least
not
site.
upper part
a
mixed
forest invaded the lower belts of the
forest
pubescens,
as
developed
well
pollen
Carpinus,
in
which deciduous
Ulmus and
as
zone an
Abies belt
can
being
was
Carpinus betulus
deciduous oak forest
(probably
interpreted
more
can
pollen
in the lower
important
be
the
to
tree
accepted
with
Q. pubescens)
the
as
Fraxinus and Acer in
present up
of the next forest phase YY
content
be
Tilia, Ostrya,
evergreen oaks and Pistacia
Above the oak
regions.
The
belts,
that
pollen composition
a
mixed oak forest with
the upper
a
a
An
zone
found higher up the slopes.
rain of
of
For
because of the
probably
was
oaks, probably Q. cerris and Q.
were
accepted.
forest established itself in the
in altitudinalbelts
forest.
up
a
the timber line.
may have been present. The deciduous elements in this forest
neighbourhood
During
higher parts
to
to
the oak forest
pine forest, probably
undergrowth.
& Panukcuouglu
pine
of minor importance,
separation
pine
a
the
forming
in temperature may be
see
whereas in the
pine forest,
further extension of the
a
Ericaceae
In subzone PP3
were
a
nordmannianatype came into existence
Zone PP2b shows
nigra
by
interval.
warmer
in the forest in relation
place
a
VV, PP1
to zone
line.
indicative
as
Pistacia and
Buxus,
Pterocarya.
From subzone YY1
a
change
were
into
a more
to
YY2 there is
found and other decidous oaks
association
elevations
described
Carpinus
Forests
during
as
(sub)
The forest of
by
the
zones
The
Horvat
were
present
(1954)
zone
H
can
and
was
as
cerra-like
well,
a
probable,
clearly
first
association
be differentiated in
(zone
HI
lower
part)
pollen grains
Q. cerris-confertae
whereas
formed
by
an
must
at
higher
have occurred
a
lower oak belt and
oak forest with
species (Q. ilex)
elevated part of this forest contained
timber line
pollen content, indicating
Q.
F.
but later also mediterraneanQuercus
more
seems
Q. cerris-confertae
A lb. Cl
upper coniferous belt. At
found,
in the
change
and Abies dominated.
resembling
the
a
continental forest type. Since
Abies forest. Some
Q.
an
cerris is
became established.
Carpinus betulus,
whereas the
Fagus pollen grains
indicated
PALYNOLOGY
OF
120
THE
M DEEP SECTION
the presence of this tree, also
forest
but the
found,
was
willows and alder
In
zone
became
of
rather
a
drought.
more
Juniperus
Chenopodiaceae
first
more
of zone
was
O,
forest
a
found in
may
climate
became
zone
eastern
a
and
increasing
the
by
with
in
a
period
zones
forest and the
frequent
more
of Artemisia and
this
during
drier
development
in situ
pine
a
amount
warmer,
colder and
interval
marked
at
was
summer
started.
O and
Q
dealt
were
resemblance to the Colchic forest types
a
of the
part
deciduous
semi-evergreen Quercus species
(zone Q),
replaced by
prevailing
kind of
area.
concluded from the
rapidly
same
reflected in the
as
pollen produced
lake
Mediterranean
from the presence of Rhododendron,
chiefly
of the Colchic type of
Pinus
and the
established with
be inferred
be
H2 the
detail (Wijmstra & Smit 1975). During the deposition
the
This forest type with
of
The
but
the
became
can
of the forest
phases
with elsewhere in
rest
open
as
by
zone
regional forest,
swampy,
gradually
At the end of this
nowadays
then
the,
pollen
pollen.
cool one,
The different
This
in
trees
H3 the oak forest
occurrence
of the
307
GREECE
elevations. In
higher
composition
is somewhat obscured
pollen samples,
vegetation
at
IN NORTHERN
zone
O is
Quercus species replaced
a
(Anatolia).
Nerium,
first
by
a
more
open
vegetation
forest
steppe vegetation (zone R,
etc.
the earlier fo-
of zone M2. The closed forest
replaced
ultimately by
area
Ilex,
see
type with
Wijmstra,
1969).
3.3
Forest steppe
This type of
vice
or
versa,
several
vegetation
vegetation
but in
occurred
most
times, however,
forest-steppe phase.
Salix and
scattered
some
cases
regularly
the transition from steppe
was
climatical circumstances
from
Apart
locally
Fraxinus, the principal
In this open
patches.
at
its presence
permitted
developed
tree was
vegetation type
to
forest
of rather short duration. At
Pinus,
some
a
prolonged
more
stands of
with
vegetation
probably occurring
Pistacia and
Juniperus
in
were
present.
A rather similar situation
3.4
Local
aquatic
and marsh
We shall end the discussion of the
in situ. In
vegetation present
algal types,
with
as
a
more
such
open
Nymphaea
as
curve
Botryococcus
and
Pediastrum,
be
were
stands
hardly
of
decrease in
must
found.
it is
Typha
Sometimes
positively
forest phases, Equisetum
spores
a
E and
L.
vegetations
with
vegetational history
place
Menyanthes.
to
XXI, B,
first
Typha phase, suggesting
Cyperaceae
zones
the
vegetation,
and
found in
was
a
water
were
that
depth.
stems
hardly
found. In the
frequently found,
were
There is
Typha
been present in
Remains of
short review of the
Nymphaea phase
correlated with the
have
a
noteworthy
were
the
a
any
phases
well
as
is followed
tendency
curve.
by
for the
During
the
hydrosere although
frequently
encountered,
however.
For
some
only
an
indication of the temperature the
of
presence
importance,
in
alpine
since
at
present this plant
mountainbogs
at an
altitude of
occurs
1,800
Menyanthes might
be of
in the Mediterranean
m
and
over
area
(Rikli 1948).
308
4.
DISCUSSION
At first
an
history
OF
THE
sequence
correlations with the
Interglacials
Wijmstra
to
distinguish
from interstadials. In the
the climatic
(1969)
WIJMSTRA
AND
A.
SMIT
RESULTS
attempt will be made
interglacials
4.1
A.
T.
for the
stratigraphy
in
next
area
in the
sequence
part, with the
is discussed
N. W.
Europe
help
under discussion
of the
vegetational
(fig. 4). Finally
are
tentative
suggested.
and interstadials
concluded that
Fig.
the forest
4. Climate
derived
phases
sequence
from the
in
the
in
the
area
corresponded
Tenagi Philippon
pollen assemblages.
area,
PALYNOLOGY
with
120
THE
OF
M
The
with
interglacials
to a
sion combined with
dinal belts
of
(in preparation)
such
proposed,
are
leading
the forest
a
to
criteria, only
the
mediterranean
O/H,
zones
Abies,
indicating
occurring
a
species,
dominated. We
interstadials for
which
the
we
an
an
two
proposed.
also marked
upper
belt,
these
interglacials, for
as
are
in
succes-
Abies
apply
and Lekanis are
as
a
series of altitu-
complete
a
C and F
by
mountainous
the
tree
submediterranean type of oak forest in
with semi-
species, perhaps together
consider these
provisionally
names
Such
vegetation.
be considered
phases YY, A,
probably
distinguishing
the presence of
Pangaion, Symvolon
the forest
most
can
cerris and other deciduous
Quercus
Strimon
Lithochoris,
phases
as
war-
and
Kavalla
Krimines,
proposed.
are
Phases with forest steppe
a
H and PP,
the final local stage will have been
mer
with
succession”
“complete
a
represent optimum climatic conditions. If
Although, however,
evergreen
further criteria for
of climax
type
pollen assemblage
which respectively the names
of
some
the incidence of
as
vegetation, including especially
is considered
which
phases corresponded
“interstadials”, during phases
minor amelioration of the climate.
only
belt,
309
GREECE
in
“Waldsteppe” appeared
In Wijmstra & Smit
presence
NORTHERN
and that the steppe
interglacials and interstadials
glacial periods.
IN
DEEP SECTION
represented
the
by
in
conditions, present
minor increase of forest elements,
Quercus
mainly
E and L and
showing
lesser extend
Quercus,
XXI, B,
Pinus and
roburj petraea type,
to a
may
be
considered
cool
as
interstadials.
4.2
Major
The
climatic
occurrence
is
assemblages
roundings
changes
of the different
given
in the
curve
of the drill hole
were
vegetation types
the
the deciduous oak forest and the
ranean area
is
mainly governed by
two
derived from their
vegetation types
pollen
in the
sur-
the forest
steppe,
open Artemisia steppe,
mediterranean oak forest. For the
it is assumed that the transition of one
vegetation type
mediter-
into another
parameters: temperature and precipitation. There-
fore the combined effect of these
on
as
of fig. 4. The main
two
factors
on
the
vegetation
cover
is reflected
the horizontal axis.
The vertical
the
being
From
the
Pangaion
zone
axis,
unknown
curve
and
the
depth,
can
can
be
read that
stages had
Symvolon
X in the upper part of the
cold climate (Wijmstra
parable
to
the
be used
as
a
relative time
base,
the parameter
of sedimentation.
rate
and Lekanis
green oak forest in those
climate
extreme
of the
zones.
This
the
points
vide the
to a
between
comparable
to a
Symvolon stage
phases,
period
character
Tenagi diagram pointing
1969). During
Pangaion
the
an
dry
and
to
climate with
the
relatively
the climate was
development
the
of
summer
com-
an ever-
drought
and winter rainfall.
Between the
Symvolon
and Strimon forest
conditions and, consequently,
of the
Kavalla,
to a
dry and
Krimenes and Lithochoris
mediterraneanclimate with
a more even
phases
a return
cold climate took
phase
spread
must
to extreme
place.
steppe
The forests
have grown under
a
of the annual precipitation
subcom-
310
T.
Table 2.
Tentative
correlations
1st
interglacial,
zone
of the
TenagiPhilippon
correlation
2nd
and N.W.
A. WOMSTRA AND A. SMIT
European sequences
correlation
interstadial
M/0
L
Eemian
Pangaion
3
Eemian
Bantega
Saalian
L
1
Hoogeveen
H
Symvolon
F
Strymon
C
Kavalla
Holsteinian
Bantega
Sa lian
«
M
0)
■U
A1
Krimenes
M
YY
Lithochoris
Cromer
III
PP
Lekanis
Cromer
II
Hoogeveen
Holsteinian
c
•H
U
Q)
4-1
in
u
to
parable
that found
the
during
mate
this
phases;
nowadays
interim
may
as
not
Lekanis
presence
intervals,
stratigraphical
correlation between the
sequence is
European
Europe.
seems to
hampered by
The correlation between the
other
steppe
grass steppes.
a severe
forest
wijn
Wijmstra
(see
When
we
(1973)
try
and
to
is
too
Therefore,
steppe
cli-
interpreted
one
sequence and the classi-
of absolute
dating
of
the
last
and
too
cannot
interglacials.
be
to
be
used
correlation in which the
leaves
Weichselian
during
a
to
two
in the
se-
Eemian
cases
glacial
is
reviewed
by
the
con-
1975).
sequence
as
zone
YY
(the
can
Zag-
be made.
Lithochoris
reliable criterium for correlation.
determine
and the Lekanis
This
in both
the following remarks
Nerium
scarce
Symvolon
European
sequence,
Pterocarya and
their presence
as
before the
dating
of the type sections in N. W.
interglacial and the
Pangaion
1969, Wijmstra & Smit
Macedonian
irregular
discussed above,
cycle
correlate the N.W.
our
occurrence
phase)
Tenagi
be rather well established because of the fact that
fully developed
cerned
viz.,
the
principally
however,
the lack
discussion and the uncertain
quence under
The
of
in
as
correlation
Stratigraphical
first
The steppe cli-
Yugoslavia.
extreme
prevailed again.
A direct
cal
was
be concluded from the
Between the Lithochoris and
mate
4.3
in Middle and W.
periods
warm
a
Holsteinian age. As
forest
phases
ways of correlation
Symvolon phase
is
(see
may be
table
2).
correlated with the Hoi-
PALYNOLOGY
OF
120
THE
M
DEEP
steinian and another in which the
Holsteinian. When
we
stadials of
and
pollen
Bantega
L3 and
zone
the
apply
NORTHERN
311
GREECE
Lekanis forest
is
phase
Symvolon-Holsteinian
Hoogeveen
In the
LI.
interstadials would
IN
SECTION
(Zagwijn
correlated with the
1973) might
be correlated with
“longer” correlation the Bantega and Hoogeveen
with the
correspond
and the
Symvolon
Lithochoris of the
Macedonia core. Within the “short” correlation the Lekanis forest
be correlated with the Cromer III
possible
decide
to
“the short”
one
to
seems
of Zagwijn.
phase
which of the
definitely
slightly
us
the inter-
correlation,
two
more
For the time
correlations is
phase
being
correct,
may
it is
not
although
probable.
ACKNOWLEDGEMENTS
The
authors
wish
thank
to
the
Netherlands
They
are
to Mrs.
M.
indebed
Posthumus who
of the
A. D. J.
the
Meeuse and Dr. T.
nee Besselink
who
prepared the samples
typed
the
Advancement
of
Pure
drilling possible.
the
der
van
Hammen,
manuscript,
and to Mr. H. J. Koerts
to Mrs.
Meijer
who
R.
vetted
and
W. Bakker
nee
who made the final draft
figures.
The
assistance
of Amsterdam
ples
to Prof.
J, Content
for
Organisation
Research (Z.W.O.) for the grant which made
of the research
and
for S.E.M,
of Mrs.
for
group
E. Thomassen
observation,
is
scanning
electron
of the
microscopy
Beglinger during the preparation
nee
University
of the
sam-
gratefully acknowledged.
REFERENCES
H.
Beug,
Ges.
J.
Probleme
(1967a):
der
in
Vegetationsgeschichte
Ber.
Siid-Europa.
Deut.
Bot.
80:682-689.
E.
Bonatti,
North Mediterranean
(1966):
climate
the last Wiirm
during
glaciation.
Nature
209:984-985.
S.
Bottema,
(1974):
Late
North-Western
Quaternary vegetation history of
Greece. Thesis
Groningen.
Florschutz,
F. &
J. Menendez
Amor
(1962): Beitrage
tionsgeschichte Nordspaniens. Veroff.
B.
Frenzel,
(1968)
der
Grundziige
Geob.
Inst.
Pleistozdnen
zur
Kenntnis der
Riibel
quartaren Vegeta-
37:68-73,
Vegetationsgeschichte
Nord
Eurasiens.
Wiesbaden.
Hammen,
Late
Th.
van
der,
Cenozoic
Horvat,
I.
of
T.
H. Wijmstra & W.
Europe.
In:
The Late
H. Zagwijn
Cenozoic
(1971):
Glacial
ages.
The floral
Yale
Univ.
record
of the
Press.
(1954). Pflanzengeographische Gliederung Siidosteuropas. Vegetatio
5-6:434-
447.
Niklewski,
Syria.
Quezel,
J. & W.
Acta Bot.
P.
&
A.
van
Zeist
Neerl.
(1970):
Panukcuouglu
bioclimatologique
de
A late
Quaternary pollen diagram
from northwestern
19:737-755.
(1973):
Contribution
a
1’etude
quelques groupements forestiers du
phytosociologique
Taurus.
Feddes Reperl.
et
84:
185-220.
Rikli,
Smit,
M.
A.
genus
—
Das
(1948):
(1973):
A
Quercus.
Acta
& T. A. Wijmstra
reconstruction
Mittelmeerldnder. Hiber
der
electron
Bot. Neerl.
microscopical study
vegetation.
C. & H.
oriental.
Rev.
of the
(Bonn)
1418
pp.
pollen morphology
in
the
22:655-665.
(1970): Application
of former
Vergnaud-Grazzini,
Mediterannee
Pflanzenkleid
scanning
of transmission electron
Acta Bot. Neerl.
Rosenberg
(1969):
Geogr. Phys.
et
de
Etude
Geol.
microscope analysis
in the
19:867-876.
paleoclimatique
Dyam.
d’une carotte de
XI:279-292.
312
T.
Wang,
Chi-Wu
Wijmstra,
Greece.
&
A.
T. A.
Acta Bol.
Smit
(Greece).
—
&
thern
W.
Neerl.
1st Int.
Symp.
on
The
Balkan
Med.
flora
and
mediterranean
(1973): Pollenanalytic
Netherlands.
press,
120
m
WIJMSTRA
313
AND
SMIT
pp.
section in
deep
A.
Northern
18:511-527.
(in preparation):
H.
Univ.
of the first 30 metres of a
(1975): Vegetation development during
Proc.
A. Smit
Zagwijn,
(1961): The forests of China. Harvard
(1969): Palynology
A.
studies
the
Eemian
vegetation,
in
Eastern
Macedonia
Sofia.
glacialinterglacial cycle.
of Holsteinian
Rijksgeologische Dienst,
and Saalian
N.S. 20:139-156.
beds
in
the Nor-
Tenagi
Philp on
II
Fig.
T.
A.
2.
Ap endix
Pol en Wijmstra Acta
diagram and Bot.
A.
Tengi Smit,
Neerl.
25(4),
Philp on Palynol gy August
11. of
the
mid le
part
(30-78
metres)
of
the
120
m
deep
section
in
Northern
Gre ce
(Macedonia).
1976,
p.
297-312.