A late Miocene leaf assemblage from Vrysses, western Crete, Greece, and its paleoenvironmental and paleoclimatic interpretation
Author
Zidianakis, Giannis
Department of Biology, University of Crete, Voutes, GR- 71305 Irakleio, Crete (Greece) zidang @ in. gr
zidang@in.gr
Author
Mohr, Barbara A. R.
Institute of Palaeontology, Museum of Natural History, Invalidenstr. 43, D- 10115 Berlin (Germany) barbara. mohr @ rz. hu-berlin. de
mohr@rz.hu
Author
Fassoulas, Charalampos
Natural History Museum, University of Crete, GR- 71409 Irakleio, Crete (Greece) fassoulas @ nhmc. uoc. gr.
fassoulas@nhmc.uoc.gr
text
Geodiversitas
2007
29
3
351
377
journal article
10.5281/zenodo.5374994
1638-9395
5374994
Quercus mediterranea
Unger
(
Figs 2
H-K; 5J)
MATERIAL EXAMINED. — Complete leaves nos. 32.1.2.6; 32.1.2.17; 32.1.2.79; 32.1.2.155; 32.1.3.2; fragmentary leaves nos. 32.1.2.24; 32.1.2.43; 32.1.2.147; 32.1.2.229; 32.1.2.233; 32.1.2.248; 32.1.3.4.
MATERIAL EXAMINED. — Complete leaf no. 32.1.3.1; DESCRIPTION
fragmentary leaves nos. 32.1.2.72; 32.1.2.158.
Five complete leaves very variable in shape, 3.3 to
6.9 cm long and
1.8 to 4.7 cm
wide. Shape broadly
DESCRIPTION oval to ovate or obovate. Base acute or rounded, One almost complete and two incomplete leaves; rarely cordate and apex acute, obtuse or slightly broadly oval to elongate, slightly asymmetric in emarginate. Leaf margin simple serrate at least in shape; at least
10 cm
long and
4.5 cm
wide. Mar- the upper half of the lamina, occasionally entire or gin entire, base widely cuneate to rounded, apex undulate. Teeth widely spaced, rather small, more not preserved. Petiole
c
.
0.5 cm
long. Primary vein or less sharply pointed. Venation craspedodromous, stout, slightly bent. Venation brochidodromous, midvein stout, in the upper part more slender and 8-10 pairs of curved secondary veins arising at 35- in some cases slightly sinuous. Secondary veins 60° from the midvein, regularly and widely spaced, distinct, in 5 to 9 pairs, arising at an angle of 30 joined together in a series of arches along the mar- to 80° from the primary vein, irregularly spaced, gin. Tertiary veins percurrent, straight or irregularly rarely forked, running straight or bent towards the branched. Leaf texture possibly leathery. leaf margin entering the teeth.Tertiary veins poorly visible, irregular, simple or branched. Leaf texture
REMARKS rather chartaceous or coriaceous.Also, 7 fragmentary Size, shape and venation pattern of these specimens leaves are assigned to this fossil species.
seems to fit the description of the fossil species
Juglans acuminata
.
This taxon is present in many European REMARKS
Neogene deposits. Leaflets of
Juglans
sect.
Juglans
Quercus mediterranea
is a characteristic common show entire margins (e.g.,
J. regia
L.). Therefore, element in Neogene deposits especially in southern
Juglans acuminata
may be related to modern
Jug-
Europe. In
Greece
, this species is represented in
lans
regia
that grows in mixed mesophytic forests numerous Miocene and Pliocene plant assemblages in southeast Europe and North Asia. However, of mainland and islands (Knobloch & Velitzelos Knobloch & Kvaček (1976) had studied the epi- 1987; Velitzelos & Gregor 1990). Leaf remains of dermal anatomy, and considered
Juglans acuminata
Quercus mediterranea
from the Vrysses outcrops to be related to the North American species
Juglans
correspond much better to the records from Vegora
rupestris
Engelm. (late Miocene, N
Greece
) and Kymi (early Miocene Since several species of
Pterocarya
Kunth
also Evia Island,
central Greece
) on the basis of grosshave leaves with entire margins, it can not be com- morphology (Velitzelos
et al.
2002; Kvaček
et al.
pletely ruled out that the Vrysses leaves belong to 2002; Kvaček & Walter 1989) while leaves assigned this taxon. In the Makrilia plant assemblages from to
Q. mediterranea
from Makrilia (late Miocene,
Miocene leaf assemblage from Vrysses, western
Crete
eastern
Crete
, S
Greece
) seem to differ slightly. At dulate or simply serrate with small widely spaced the latter locality oaks are represented by a few teeth, serration starting in the lower third of the leaves, which are rather small. They hardly reach a lamina. Base cordate, apex acute maybe acuminate. length of
2.5 cm
(
Sachse 1997
). Venation craspedodromous. Midvein prominent Based on studies of leaf epidermis tissue from more or less curved, slightly zig-zag at the upper Lava (North
Greece
) that indicated an extremely part. Secondary veins more than 10 pairs, regularly thin cuticle, it was suggested that
Q. mediterranea
disposed, quite dense, diverging at angles of 40 to was a mesophytic deciduous oak (Kvaček & Walter 80° and running straight, parallel to each other, 1989). However newly obtained epidermal charac- across lamina, entering marginal teeth. Higher teristics show that in spite of the thin cuticle, the order venation poorly preserved. epidermis of either leaf side is sclerenchymatous underlain by a hypodermis (Kvaček
et al.
2002). REMARKS Thus
Q. mediterranea
probably was a sclerophyllous Such leaves may be assigned to the genus
Quercus
, element able to survive short dry periods. representing probably an evergreen oak quite dif-
Quercus mediterranea
is perhaps ancestral to the ferent from
Q. mediterranea
especially in leaf size modern evergreen species
Quercus ilex
L. and
Quer-
and shape and secondary veins pattern. Leaf remains
cus
coccifera
L., although both do not correspond from early Miocene plant assemblage of Kymi figured in all details of leaf anatomy and morphology to by
Unger (1867
: taf. VI, figs 23-27) and identified the fossils. Today these oaks are present through- as
Quercus drymeja
Unger
by Velitzelos
et al.
2002 out the Mediterranean basin and on coastal slopes (revised list of Kymi material), are very close in mortowards the Atlantic (South
France
) and Black Sea, phology to Vrysses specimens. However more and on relatively dry soils. better preserved material is necessary to elucidate According to Barbero
et al
. (1992),
Q. ilex
thrives the systematic relationships of these specimens. in a wide climatic range (from warm-temperate to subtropical and from medium humid to semi-arid) due to numerous morphological adaptations. In Family
ULMACEAE
the northern Mediterranean realm,
Q. ilex
forms woodland with several laurophyllus elements such
Ulmaceae
gen. et sp. indet. as
Arbutus
L.,
Hedera
L.,
Ilex
L. and
Viburnum
L., (
Fig. 3B
) while in south it thrives together with more xerothermic taxa such as
Juniperus
L.,
Phillyrea
Tourn.
MATERIAL EXAMINED. — Complete leaf no. 32.1.2.70.
ex Adans.,
Genista
,
Ephedra
L. and
Buxus
. In the eastern Mediteranean basin
Q. ilex
is replaced by DESCRIPTION
Q. coccifera
. One small, broadly elliptic leaf, about
1.8 cm
long and
1 cm
wide. Apex incomplete, probably acute to acuminate, base rounded, strongly asymmetrical.
Quercus
sp.
Leaf margin serrate at the upper half part of lamina
(
Figs 3A
;
5Q
) with several wide, simple teeth. Venation craspedodromous. Midvein prominent, quite thick and
MATERIAL EXAMINED. — Fragmentary leaves nos. strongly curved. From the primary vein arise four 32.1.2.19; 32.1.2.95; 32.1.2.142.
pairs of secondaries at angles of 35 to 55°, rather strong, more or less curved, running into the tooth
DESCRIPTION apices. No other details of the venation preserved. Three leaf fragments, long petiolate (up to
1.5 cm
). Complete leaf length estimated at
7 to 9 cm
or REMARKS perhaps more, width about
2.5 to 3.2 cm
. Shape Such a leaf shape (leaf margin, asymmetrical base, of lamina ovate to broadly elliptic, leaf margin un- laminar shape) suggests a relationship with the
Zidianakis G.
et al.
family
Ulmaceae
. However this specimen lacks margin dentate with obtuse teeth. Petiole not prefurther diagnostic features for a determination to served. Venation actinodromous with three prigenus level. The
Ulmaceae
is a family of tropical mary veins. Midvein slightly bent, laterals curved and mainly temperate trees and shrubs, with 16 and forked. Four or five pairs of curved secondary genera (Heywood
et al.
1993). Due to very small veins arising from the midvein at 30-55°, looping size (nanophyll) and simple dentate margin, the with the laterals and the adjacent secondaries near Vrysses specimen may represent a slightly xero- the margin.Tertiary veins oblique forming meshes morphic element. between secondaries.
REMARKS Family
SALICACEAE Shape
, teeth at the leaf margin and venation of Genus
Salix
L. these specimens are similar to lamina of the modern species
Populus tremula
. Leaf remains of this cf.
Salix
sp.
poplar have been found in several European Neo- (
Fig. 3C
) gene deposits. Today
Populus tremula
trees grow in open mesophilous forests up to
1000 m
of altitude
MATERIAL EXAMINED. — Fragmentary leaf no. in Europe, North Africa,
Asia Minor
and Siberia
32.1.2.39. (
Leroy & Roiron 1996
).
DESCRIPTION
Basal half of a probably narrow elliptic, slightly Family
BUXACEAE
asymmetrical leaf (
3.9 cm
long and
1.7 cm
wide); Genus
Buxus
L.
entire leaf possibly twice that long. Leaf base acute,
margin finely serrate with simple acuminate teeth,
Buxus pliocenica
Saporta & Marion
regularly spaced. Midvein medium, thick, well (
Figs 3E
;
5M
)
developed and curved. Higher order venation not visible. M ATERIAL EXAMINED. — Complete leaf no.
32.1.2.140. REMARKS
Shape and margin of this leaf show resemblance to DESCRIPTION those of extant
Salix
L. However
, because the upper One complete leaf, narrowly elliptic, with short part of the leaf is missing, as well as higher order petiole,
3.6 cm
long and
1.2 cm
wide. Leaf margins venation, a definite identification is not possible. entire, apex acute, base cuneate. Primary vein thin and almost straight. Secondary veins fine, arising at an angle of 30 to 70°, closely spaced, parallel
Genus
Populus
L. to each other, branched once to many times in acute angles without higher order veins rejoining
Populus tremula
L., foss. secondaries.
(
Fig. 3D
)
REMARKS MATERIAL EXAMINED. — Complete leaf no. 32.1.3.7;
Buxus pliocenica
fossils have been known in Mifragmentary leaf no. 32.1.2.160.
ocene and Pliocene deposits of Eurasia, mostly in the form of leaf impression and compression, rarely DESCRIPTION as pollen or fossil fruits. Saporta & Marion (1876) One complete and one fragmentary leaf. Lamina treat
Buxus pliocenica
as an ancestor of
Buxus sembroadly
ovate,
4.5 to 4.8 cm
long and about 5.5 to
pervirens
L. However, Kvaček
et al.
(1982) consider
5.7 cm
wide. Apex probably obtuse, base rounded,
Buxus pliocenica
closely allied to the
B. sempervirens
-
Miocene leaf assemblage from Vrysses, western
Crete
FIG. 3. — Photographs of fossil leaves from the Vrysses outcrops (western Crete):
A
,
Quercus
sp.
, no. 32.1.2.142;
B
,
Ulmaceae
gen. et sp. indet., no. 32.1.2.70;
C
, cf.
Salix
sp.
, no. 32.1.2.39;
D
,
Populus tremula
L., foss., no. 32.1.3.7;
E
,
Buxus pliocenica
Saporta & Marion
, no. 32.1.2.140;
F
,
Rosaceae
gen. et sp. indet., no. 32.1.2.55;
G
,
Fabaceae
type 1, no. 32.1.2.161;
H
,
Fabaceae
type 2, no. 32.1.2.78;
I -L
,
Acer pseudomonspessulanum
Unger
, nos. 32.1.2.61, 32.1.3.8, 32.1.2.46 and 32.1.2.67;
M
,
Acer
sp.
, no. 32.1.2.12;
N -P
, cf.
Ziziphus ziziphoides
(Unger) Weyland
, nos. 32.1.2.3, 32.1.2.180 and 32.1.2.5. Scale bars: 1 cm.
Zidianakis G.
et al.
B. colchica
-
B. hyrcana
group. According to these DESCRIPTION authors this group replaced the large-leaved
Buxus
One
complete, very well preserved leaflet;
3.3 cm
lineage which became extinct because of the climatic long and
1.5 cm
wide, with oval outline. Margin of deterioration during the late Miocene. Extant
Buxus
lamina entire, rounded at the apex and quite
asymsempervirens
is a typical element of sclerophyllous metrical at the base. Texture chartaceous, petiolule vegetation in Mediterranean basin while
Buxus
missing. Venation brochidodromous. Primary vein
hyrcana
Pojark.
and
Buxus colchica
Pojark.
occur in thin, running irregularly (zig-zag) to the apex of pure stands or in underwood in lowland deciduous leaflet. Five pairs of secondaries arising at an angle broad-leaved mesophilous forests in the Hyrcanian of 30 to 80° from the midvein, running more or region (between the southern shores of the Caspian less sinuous, joining each other by loops near the Sea and the Elburz Mountains) and in the
Colchis
margin. The lateral secondary at one leaflet side is (Western
Georgia
). prominent, extending to the upper third of the middle of the leaflet. Intersecondary veins two or three,
weakly developed. Higher order venation branching
Family
ROSACEAE
out and joining the secondaries to form a network of rather large, irregularly shaped areoles.
Rosaceae
gen. et sp. indet.
(
Figs 3F
;
5G, S
) REMARKS
This
type
of leaflets is related to the subfamily Cae- MATERIAL EXAMINED. — Complete leaf no. 32.1.2.41; salpinioideae and especially to the artificial genera fragmentary leaves nos. 32.1.2.8; 32.1.2.55.
Cassiophyllum
and
Caesalpinites
by the asymmetrical form, the size and the venation pattern. These DESCRIPTION genera occur in European deposits from the Oli- One complete and two fragmentary leaves with short gocene to Miocene. The determination of modern petiole. Leaf length
2.7 to 3.5 cm
(may be more) equivalents is difficult because the leaflets of many and width
1.3 to 2 cm
. Shape of lamina ovate or species have a similar morphology, as for instance oval, rounded at the base acute at the apex. Margin
Scohtia humboltioides
Oliv.,
Wagetia spicata
Daez
finely serrate with small, simple and sharp teeth. and some
Cassia
species (Shakryl 1992)
. Primary vein well developed and straight. Higher order veins not visible.
Fabaceae
type
2 REMARKS (
Fig. 3H
) The features observed in these fossils and especially leaf margin suggest an affinity to the family MATERIAL EXAMINED. — Almost complete leaflet no.
32.1.2.78.
Rosaceae
. Today this large family comprises about 3300 species. It is cosmopolitan with maximum diversity in the northern temperate area (Heywood DESCRIPTION
et al.
1993). One leaflet
3.8 cm
long (complete about
4 cm
) and
1.5 cm
wide, narrow elliptic to lanceolate, slightly asymmetrical,entire-margined.Apex of lamina acute,
Family
FABACEAE
base obtuse or rounded, asymmetrical (part of the base is missing). Primary vein strong, S-like curved.
Fabaceae
type
1 Higher order venation fine, hardly visible.
(
Fig. 3G
)
REMARKS MATERIAL EXAMINED. — Complete leaflet no. Entire margins and asymmetry of this specimen 32.1.2.161. are features that suggest a relationship with the
Miocene leaf assemblage from Vrysses, western
Crete
Fabaceae
. Unfortunately, poor preservation of the Weber. The name
Acer pseudomonspessulanum
is leaflet does not allow a more precise determination. used as the Neogene ancestors of the modern spe-
Fabaceae
is a very large plant family (approximately cies of the series Monspessulana Pojark., especially 17000 species) with great variety of habitat and a
A. monspessulanum
and
A. sempervirens
and their cosmopolitan distribution in tropical, subtropical intermediate forms (
Leroy & Roiron 1996
). Leaf and temperate zones (Heywood
et al.
1993). The and fruit remains of
Acer pseudomonspessulanum
are small-sized leaflet of Vrysses may belong to a more known from several Neogene deposits in Europe. or less xeric element. Today
Acer monspessulanum
extends from central,
south Europe and northwest Africa to north
Iran
(south shores of the Caspian Sea), while
A. sem-
Family
ACERACEAE
pervirens
occurs at the east Mediterranean basin.
Genus
Acer
L.
Acer monspessulanum
is a deciduous element while
A. sempervirens
is considered as deciduous or semi
-
Acer pseudomonspessulanum
Unger
evergreen element. Both of these maples are trees
(
Figs 3
I-L; 5L, P, R) or shrubs and common elements of more or less xerothermic mixed evergreen and deciduous forests. MATERIAL EXAMINED. — Complete leaves nos. 32.1.2.46; Today
Acer sempervirens
grows on
Crete
at moun- 32.1.2.61; 32.1.2.67; 32.1.3.8; fragmentary leaves nos. tain slopes between 300 and
1700 m
(Turland
et
32.1.2.28; 32.1.2.29; 32.1.2.31; 32.1.2.37; 32.1.2.40; 32.1.2.45; 32.1.2.58; 32.1.2.62; 32.1.2.91; 32.1.2.244;
al
. 1993). 32.1.3.9; 32.1.3.10.
DESCRIPTION
Acer
sp.
Four complete and 12 incomplete leaves, are assigned (
Figs 3M
;
5K
) to this fossil species. Lamina deeply trilobate,
2.1 to 4 cm
(or somewhat more) long and
3 to 6.3 cm
MATERIAL EXAMINED. — Almost complete leaves nos.
32.1.2.12; 32.1.2.56;
fragmentary leaves nos. 32.1.2.14;
wide, with short petiole. Leaf texture coriaceous. 32.1.2.49; 32.1.2.75.
Base of lamina rounded or cordate, margin entire. The median lobe more or less broader than the lat- DESCRIPTION eral lobes. Apices of lobes acute to obtuse. Venation Two almost complete leaves and 3 fragments,5-lobed, actinodromous with three primary veins, straight
3.1 cm
to
3.9 cm
(possibly more) long and 4.4 to to slightly bent. Secondary veins fine, dense, aris-
5.8 cm
wide. Margins entire with acuminate apex. ing at an angle of 60 to 80° from the primaries, Leaf base cordate, venation actinodromous with branching near the margin. Higher order veins five primary veins. Lateral primaries arising under poorly preserved. angles of 40 to 55° from the midvein, while basal primaries arise under angles of 80 to 110°. Five to
REMARKS seven pairs of hardly visible secondary veins, arise Most of these fossil leaves (the larger specimens) are at 30 to 55° from primary veins and run in curves most likely related to
Acer monspessulanum
L. but towards the leaf margin. some of them, with sharper sinuses and less broad lobes with acuminate apices which stretch upwards, REMARKS are related to
Acer sempervirens
L. In
fact, remains of Some of these specimens are comparable to
Acer
inte-
Acer pseudomonspessulanum
from Vrysses do not differ
gerrinum
, a common element of European Neogene in gross-morphology from the records in Makrilia, deposits. In
Greece
,
A. integerrinum
is represented from where this leaf
type
was described as
Acer
series in the late Miocene fossiliferous deposits of Licudi, Monspessulana Pojark. Material from Vegora that
central Greece
(Velitzelos & Gregor 1990). is also similar in its variability has been determined
Acer laetum
C.A.Mey.
and
Acer pictum
Thunb.
as
A. pseudomonspessulanum
and
A. integrilobum
are considered to be the possible analogue among
Zidianakis G.
et al.
extant maples on the basis of their gross-morphology
Dicotylophyllum
sp. 1
(
Leroy & Roiron 1996
).
Acer laetum
thrives from (
Fig. 4A
) the Caucasus to the Himalayas and the mountains of central
China
while
A. pictum
grows on the slopes
MATERIAL EXAMINED. — Complete leaf no. 32.1.2.1.
of northern
China
and
Japan
.
DESCRIPTION
One entire leaf, reniform,
2.4 cm
long and about
Family
RHAMNACEAE
3.2 cm
wide, stalked, entire-margined. Leaf texture
Genus
Ziziphus
Mill.
evidently fine.Venation actinodromous with seven primary veins. Midvein straight, stronger than the
cf.
Ziziphus ziziphoides
(Unger) Weyland
lateral primaries. All primary veins forked before
(
Fig. 3
N-P) reaching leaf margin. Secondary veins on the me-
dial primary vein widely and irregularly spaced
MATERIAL EXAMINED. — Almost complete leaves nos. in four pairs, also forked. Higher order venation 32.1.2.3; 32.1.2.5; 32.1.2.22; 32.1.2.180; fragmen- not visible. tary leaves nos. 32.1.2.10; 32.1.2.32; 32.1.2.106; 32.1.2.209.
REMARKS
DESCRIPTION A very interesting leaf form. In morphology (ve- Four almost complete leaves and four fragments. nation, margin, shape), it is comparable with the Lamina broad oval to ovate
3.2-5.8 cm
long (maybe foliage of modern
Cercis silliquastrum
L. (
Fabaceae
), more) and
1.9-4.1 cm
wide, simply dentate at the differing mainly by having smaller leaf size than upper two thirds of the leaf margin. Apex acute to living element.
Cercis silliquastrum
is a Mediterobtuse, base partly asymmetric. Venation acrodro- ranean, deciduous tree. mous with three stout primary veins running in convergent arches toward the leaf apex.Higher order veins very fine, hardly visible, vertical to midvein.
Dicotylophyllum
sp. 2
(
Figs 4B
;
5H
)
REMARKS The specimens described above probably belong to
MATERIAL EXAMINED. — Almost complete leaf no.
the species
Ziziphus ziziphoides
, a common element
32.1.2.92.
in the European Paleogene and Neogene. The genus
Ziziphus
consists of about 100 species of decidu- DESCRIPTION ous or evergreen trees and shrubs distributed in One almost complete (only a small part of its apex the tropical and subtropical regions of the world is missing) leaf, narrow oblanceolate
6.2 cm
long (Johnston 1963). This leaf
form may
be related to (complete leaf somewhat more) and
2.2 cm
wide. a non-sclerophyllous species of
Ziziphus
not iden- Lamina with entire margin, gradually narrowing tical with recent species of the Mediterranean area towards the base. Primary vein straight, very strong and the Near East. The nearest extant relatives of and thick. Leaf texture coriaceous. Secondary veins
Ziziphus ziziphoides
considered to be the decidu- fine, hardly visible, arising at 30 to 50° from the ous
Z. sinensis
and
Z. glabrata
both thrive in Asia midvein. (Givulescu 1962). However, this leaf
type
may also represent a completely different family (see Sachse REMARKS 1997;
Dicotylophyllum
type
3). This leaf with poorly preserved venation cannot be reliably assigned to any specific genus. The general shape of this specimen suggests that it
Family
incertae sedis
represented a broad-leaved humid or mesic ele-
Genus
Dicotylophyllum
Saporta
ment.
Miocene leaf assemblage from Vrysses, western
Crete
FIG. 4. — Photographs of fossil leaves identified in Vrysses outcrops (western Crete):
A
,
Dicotylophyllum
sp.1
, no.32.1.2.1;
B
,
Dicotylophyllum
sp.2
, no.32.1.2.92;
C
,
Dicotylophyllum
sp.3
,no.32.1.2.16;
D
,
Dicotylophyllum
sp.4
,no.32.1.2.33;
E
,
Dicotylophyllum
sp.5
,no.32.1.3.6;
F
,
Dicotylophyllum
sp. 6
, no. 32.1.2.9;
G
,
Dicotylophyllum
sp. 8
, no. 32.1.2.223;
H
,
Dicotylophyllum
sp. 9
, no. 32.1.2.30;
I
,
Dicotylophyllum
sp.
10, no. 32.1.2.93;
J
,
Dicotylophyllum
sp.
11, no. 32.1.2.159;
K
,
Poaceae
/
Cyperaceae
type 1, no. 32.1.2.191;
L
,
Poaceae
/
Cyperaceae
type 2, no. 32.1.2.54. Scale bars: 1 cm.
Zidianakis G.
et al.
Dicotylophyllum
sp. 3
broadly lanceolate to obovate, apex acute to obtuse
(
Fig. 4C
) and base obtuse to slightly cordate. Margin slightly wavy or dentate with small simple teeth. Venation MATERIAL EXAMINED. — Complete leaf no. 32.1.2.16. semicraspedodromous. Primary vein prominent,
somewhat straight. Secondary veins thin, densely DESCRIPTION spaced at angles of 45 to 80°, running almost straight, One complete leaf (only petiole missing), narrowly parallel to each other across the lamina, dichotomised elliptic, slightly asymmetrical leaf;
5 cm
long and near the margin before entering teeth.Tertiary veins
1.2 cm
wide. Apex rounded, base obtuse, somewhat percurrent or simply forked forming irregular meshes asymmetrical, leaf margin entire.Primary vein stout, with quaternary veins. Also two fragmentary leaves S-like curved. Secondary veins fine, rather dense, are assigned to this form.
hardly visible.
REMARKS
REMARKS This fossil leaf taxon is similar in gross-morphol- It has not been possible to determine the affinity ogy to numerous genera in different families such of this leaf due to poor preservation. as
Rhamnaceae
and
Fagaceae
.
Dicotylophyllum
sp. 4
(
Figs 4D
;
5O
)
Dicotylophyllum
sp. 6
(
Figs 4F
;
5I
)
MATERIAL EXAMINED. — Complete leaves nos. 32.1.2.27; MATERIAL EXAMINED. — Fragmentary leaves nos. 32.1.2.9; 32.1.2.33; 32.1.2.60; 32.1.2.144. 32.1.2.107.
DESCRIPTION DESCRIPTION Four complete leaves, small, long petiolate, oblan- Two fragmentary leaves with broad obovate outceolate to obovate leaves,
4 to 5.5 cm
long and line,
5 to 5.7 cm
long (perhaps more) and 2.2 to
1 to 1.5 cm
wide. Leaf apex obtuse or rounded,
3.2 cm
wide. Petiole rather slender, up to
1.2 cm
base attenuate and leaf margin entire. Texture co- long. Base of lamina heart-shaped, apex absent, riaceous. Primary vein strong, more or less curved. with undulate or finely, simply dentate margin. No other details of the venation visible. Midvein fine, hardly visible, straight or slightly bend. Higher order venation invisible. Leaf texture REMARKS coriaceous. The affinity of these specimens is unknown and much better preserved material is necessary for a REMARKS reliable identification. The poor preservation makes it difficult to clarify the systematic relationship of these specimens.
Dicotylophyllum
sp. 5
(
Figs 4E
;
5B, N
)
Dicotylophyllum
sp. 7
(
Fig. 5T
)
MATERIAL EXAMINED. — Complete leaves no. 32.1.3.4;
32.1.3.6; fragmentary leaves nos. 32.1.2.34; 32.1.2.35. MATERIAL EXAMINED. — Complete leaf no. 32.1.2.44.
DESCRIPTION DESCRIPTION
Two almost complete petiolate leaves. Length of the One complete, petiolate, slightly asymmetrical, lamina
2.5 to 3.9 cm
and width
1.7 to 1.9 cm
. Leaf broadly oval leaf with entire margin. Leaf length
Miocene leaf assemblage from Vrysses, western
Crete
FIG. 5. — Drawings of selected fossil leaves from Vrysses (western Crete):
A
,
Dicotylophyllum
sp.
12, no. 32.1.2.42;
B
,
N
,
Dicotylophyllum
sp. 5
, nos. 32.1.3.4 and 32.1.2.34;
C
,
Dicotylophyllum
sp.
10, no. 32.1.2.4;
D
,
Daphnogene polymorpha
(A.Braun) Ettingsh.
, no. 32.1.2.26;
E
,
Dicotylophyllum
sp.
11, no. 32.1.2.98;
F
, cf.
Juglans acuminata
Braun ex Unger
, no. 32.1.3.1;
G
,
S
,
Rosaceae
gen. et sp. indet., nos. 32.1.2.41 and 32.1.2.8;
H
,
Dicotylophyllum
sp. 2
, no. 32.1.2.92;
I
,
Dicotylophyllum
sp. 6
, no. 32.1.2.107;
J
,
Quercus mediterranea
Unger
, no. 32.1.2.147;
K
,
Acer
sp.
, no. 32.1.2.56;
L
,
P
,
R
,
Acer pseudomonspessulanum
Unger
,nos.32.1.3.9,32.1.2.37 and 32.1.2.31;
M
,
Buxus pliocenica
Saporta & Marion
, no. 32.1.2.140;
O
,
Dicotylophyllum
sp. 4
, no. 32.1.2.144;
Q
,
Quercus
sp.
, no. 32.1.2.95;
T
,
Dicotylophyllum
sp. 7
, no. 32.1.2.44. Scale bar: 1 cm.
5.9 cm
, width
2.2 cm
. Leaf base obtuse, apex round- veins at least seven pairs, hardly visible, very deli- ed. Venation probably semicraspedodromous. Pri- cate, rather widely spaced, arising at angles of 55 mary vein quite strong, S-like curved. Secondary to 80° from the midvein, running nearly straight
Zidianakis G.
et al.
to somewhat bent towards the margin. Probably 3.6 long and
3 cm
wide). Leaf apex not preserved, secondaries forking near the margin. Further vena- base obtuse, margin hardly visible, undulate. Leaf tion pattern not preserved. texture coriaceous. Venation craspedodromous, primary vein stout, S-like curved. Secondaries at REMARKS least seven pairs, strong, parallel, regularly spaced, The systematic affinity of this leaf taxon remains more or less straight, directed towards the margin. unclear. Higher order veins not visible.
REMARKS
Dicotylophyllum
sp. 8
Such leaves may be related to the genus
Fagus
L., (
Fig. 4G
) but such affinity cannot be proved. More and much better preserved material is necessary to elucidate MATERIAL EXAMINED. — Almost complete leaf no. the position of this specimen. 32.1.2.223.
DESCRIPTION
Dicotylophyllum
sp.
10 One leaf almost complete (only a small part of apex (
Figs 4I
;
5C
) missing), broadly ovate, quite asymmetrical,
5.7 cm
long (complete leaf about
6 cm
) and
4.3 cm
wide.
MATERIAL EXAMINED. — Complete leaf no. 32.1.2.93;
fragmentary leaves nos. 32.1.2.4; 32.1.2.94. Apex of lamina acute or acuminate and base heartshaped. Margin simply dentate with moderately large and acute teeth, irregularly spaced. Venation DESCRIPTION actinodromous with five primary veins.Primary vein One complete and two fragmentary leaves, petirather stout, S-like. Basal primaries quite weak, aris- olate, with elliptic outline. Their length is 5 to ing from the midvein at angles of 100 to 120° while
6.2 cm
(maybe somewhat more) and their width lateral primaries stronger, arising at angles of 60 to
1.7 to 1.9 cm
. Leaf margin entire, acute at the apex 80°. Four pairs of curved secondary veins arising from and cuneate at the base. Primary vein prominent the midvein, widely spaced, dichotomised near the and straight. Secondary veins fine, hardly visible. margin.Tertiary veins usually simple forked forming Higher order venation not preserved. polygonal meshes with quaternary veins. REMARKS REMARKS The systematic affinity of these
three specimens
Crucial characters are missing and thus it is im- is unclear. possible to determine the systematic affinity of this specimen.
Dicotylophyllum
sp.
11 (
Figs 4J
;
5E
)
Dicotylophyllum
sp. 9
(
Fig. 4H
)
MATERIAL EXAMINED. — Complete leaf no. 32.1.2.159;
fragmentary leaves nos. 32.1.2.98; 32.1.2.131.
MATERIAL EXAMINED. — Fragmentary leaf no. DESCRIPTION
32.1.2.30. One complete leaf, the largest in the Vrysses assemblage,
17 cm
long and
4 cm
wide (size of lamina
DESCRIPTION more than
4.500 mm
2). Also two fragmentary leaves,
One fragmentary leaf, broadly elliptic or obovate, smaller, up to
12 cm
long and
2 to 2.5 cm
wide.
2.6 cm
long and
2.5 cm
wide (entire leaf about Leaf apex obtuse or acute, base cuneate, entire-
Miocene leaf assemblage from Vrysses, western
Crete
margined, with broad elliptic outline. Primary
Cyperaceae
. The
Poaceae
and
Cyperaceae
are two vein characteristically thick and strong, straight large families of monocotyledons, cosmopolitan. or slightly bent. Higher order venation not visible Only few ecological plant formations lack
Poaceae
, probably duo to thick coriaceous texture. while many formations are dominated by them (Heywood
et al.
1993).
REMARKS
Such leaf specimens are usually considered to belong to magnoliid families, such as
Magnoliaceae
Poaceae
/
Cyperaceae
type
2
or
Lauraceae
. However a relationship with other (
Fig. 4L
)
families cannot be excluded.
MATERIAL EXAMINED. — Fragmentary leaves nos. 32.1.2.54; 32.1.2.198.
Dicotylophyllum
sp.
12
(
Fig. 5A
) DESCRIPTION
Two fragments of broad linear leaves, wide, more MATERIAL EXAMINED. — Complete leaf no. 32.1.2.42. than
2 cm
wide, with parallel venation.
DESCRIPTION REMARKS A small complete leaf, short petiolate,
3.2 cm
long These specimens probably also represent leaves of and
1 cm
wide. Lamina narrow obovate, with apex
Poaceae
or
Cyperaceae
. rounded, base acute and entire margin. Primary vein quite strong and thick, S-like curved. Secondary veins poorly preserved, arise from the midvein at FLORISTIC COMPOSITION angles of 30 to 50°. Leaf texture chartaceous.
The Vrysses plant assemblage is documented by REMARKS foliage. More than
260 specimens
of leaf mega- This leaf lacks diagnostic features. fossils, in many cases very fragmentary, were examined. A total of 31 taxa of vascular plants have been described based on their macro morpho-
Class
MONOCOTYLEDONAE
logical features (
Table 1
). Nineteen of them were
Families
POACEAE
/
CYPERACEAE
relatively well preserved and identified at least to family level.Two genera of conifers and 29 taxa of
Poaceae
/
Cyperaceae
type
1 angiosperms, shrubs, trees and, rarely herbs have
(
Fig. 4K
) been documented.
Conifers are represented in the assemblage by MATERIAL EXAMINED. — Fragmentary leaves nos.
Tetraclinis
and
Pinus
shoots and leaves. Fertile 32.1.2.57; 32.1.2.63; 32.1.2.151; 32.1.2.169; 32.1.2.186; parts, such as cones, cone scales or seeds were not 32.1.2.189; 32.1.2.190; 32.1.2.191; 32.1.2.192; 32.1.2.199; 32.1.2.203; 32.1.2.208; 32.1.2.224; found. 32.1.2.228; 32.1.2.236; 32.1.2.245. Angiosperm taxa make up approximately 95%
of the total flora. Among the angiosperms, Acer- DESCRIPTION aceae and
Fagaceae
are most common.
Fagaceae
is Sixteen fragments of narrow strap-like leaves about represented by the genus
Quercus
, mainly by the
1 to 1.5 cm
wide, that bear parallel venation. probably evergreen oak
Quercus mediterranea
, which predominates and
Quercus
sp.
considered also to REMARKS belong to evergreen oaks. Two leaf-types belong to These fossil leaves are similar in gross-morphol-
Aceraceae
; a small 3-lobed leaf identified as
Acer
ogy to numerous genera mainly in
Poaceae
and
pseudomonspessulanum
, considered to be more or
Zidianakis G.
et al.
TABLE 1. — Summary of the taxa of the Vrysses flora (western Crete),their dispersal mechanisms and physiognomic signatures.Note:
a
, taxa which cannot be restricted to one single category,are included in all assigned categories.Percentage of entire-margined taxa:48.1% (13 taxa).Percentages of notophyll,microphill and nanophyll taxa:notophyll 7.4% (2 taxa),microphyll 85.2% (23 taxa) and nanophyll 7.4% (2 taxa). Percentages related to deciduous, laurophyllous and sclerophyllous taxa: deciduous 37-59% (10-16 taxa), laurophyllous 11-30% (3-8 taxa) and sclerophyllous 15-52% (4-14 taxa). Abbreviations:
D
, deciduous;
L
, laurophyllous;
S
, sclerophyllous.
Family Genus Species/ Leaves/ Leaf mar- Maximum leaf D, L, S a Dispersal
type
series needles shoots gin entire size class mechanism
GYMNOSPERMAE
Pinaceae
Pinus
sp.
17 needles - - - wind
Cupressaceae
Tetraclinis
sp. 4
twig fragments - - - wind
ANGIOSPERMAE
Dicotyledonae
Lauraceae
Daphnogene polymorpha
2 yes microphyll L endozoochory
Hamamel-?“
Parrotia
”
pristina
1 no microphyll D autochory idaceae
Juglandaceae
?
Juglans acuminata
3 leaflets yes notophyll D dyschory
Fagaceae
Quercus mediterranea
12 no microphyll S dyschory
Quercus
sp. 3
no microphyll S dyschory
Ulmaceae
gen. indet. - 1 no nanophyll D/S wind
Salicaceae
?
Salix
sp. 1
no microphyll D wind
Populus tremula
, foss. 2 no microphyll D wind
Buxaceae
Buxus pliocenica
1 yes microphyll L/S autochory/ myrmecochory
Rosaceae
gen. indet. - 3 no microphyll D wind/ exozoo-/endozoochory
Fabaceae
gen. indet.
type
1 1 leaflet yes microphyll D/S -
gen. indet.
type
2 1 leaflet yes microphyll D/S -
Aceraceae
Acer
pseudomons-
16 no microphyll D/S wind
pessulanum
Acer
sp. 5
no microphyll D wind
Rhamnaceae
?
Ziziphus ziziphoides
8 no microphyll D endozoochory
Family
incertae
Dicotylophyllum
sp. 1
1 yes microphyll D/L/S -
sedis
Dicotylophyllum
sp. 2
1 yes microphyll L -
Dicotylophyllum
sp. 3
1 yes microphyll S -
Dicotylophyllum
sp. 4
4 yes microphyll L/S -
Dicotylophyllum
sp. 5
4 no microphyll D/S -
Dicotylophyllum
sp. 6
2 no microphyll D -
Dicotylophyllum
sp. 7
1 yes microphyll L/S -
Dicotylophyllum
sp. 8
1 no microphyll D -
Dicotylophyllum
sp. 9
1 no microphyll D -
Dicotylophyllum
sp.
10 3 yes microphyll L/S -
Dicotylophyllum
sp.
11 3 yes notophyll L -
Dicotylophyllum
sp.
12 1 yes nanophyll S -
Monocotyledonae
Poaceae
/ gen. indet.
type
1 16 - - - wind/
Cyperaceae
nautochory/ exozoo-/ endozoochory gen. indet.
type
2 2 - - - wind/ nautochory/ exozoo-/ endozoochory less xeromorphic and a 5-lobed leaf represents most naceae may be represented by irregularly dentate, likely a mesophytic deciduous species. 3-veined leaves that are close in appearance to the
Fabaceae
are represented by a few leaflets that have Neogene species
Ziziphus ziziphoides
. Leaves of been subdivided into two morphotypes (
Fabaceae
Lauraceae
seem to have been present in the assemtype 1 and
type
2).
Juglandaceae
seem to be also blage as well by the narrow leaf
type
of
Daphnogene
present. The leaflets of the Vrysses plant assemblage
polymorpha
. Imprints of
Populus tremula
, foss.,
Salix
may belong to
Juglans acuminata
. The family Rham- sp. and possibly “
Parrotia
”
pristina
occur rarely.One
Miocene leaf assemblage from Vrysses, western Crete leaf of
Buxus
sp.
has been identified.
Rosaceae
and from different habitats and the high percentage of
Ulmaceae
may have been present as well. Mono- small-sized leaves (Ferguson 1985; Spicer 1991). cots are represented by numerous specimens of the More than 85% of the taxa identified in Vrysses families
Poaceae
and
Cyperaceae
. However many outcrops are classified as microphyll (225-2.025 leaf impressions of the Vrysses plant assemblage mm2), while 7.4% as nanophyll (
25-225 mm
2) have doubtful affinities. They are considered to (
Table 1
). The allochthonous nature of the assembelong to 12 taxa of angiosperms, and classified as blage is also reflected by the absence of different
Dicotylophyllum
. organs, such as seeds and fruits of the same taxon Considering the number of specimens, four woody in the assemblage and by the high percentage of taxa are most common in the assemblage:
Pinus
sp.
, heavily fragmented leaves that reach 64% of the
Acer pseudomonspessulanum
,
Quercus mediterranea
examined specimens. and
Ziziphus ziziphoides
(
Table 1
).
Tetraclinis
sp.
Plant debris have probably been transported by and
Acer
5-lobed are found also more frequently. river floods into their offshore burial site. Alterna- More than one species may have existed among tively, plant remains may have drifted on the sea
Aceraceae
,
Fagaceae
Salicaceae
and
Fabaceae
(all surface and reached the locality of sedimentation two taxa). after having been blown by strong winds from high The majority of the genera identified at Vrysses coastal cliffs. are related to elements now growing in the North- According to its composition, the Vrysses plant ern Hemisphere. The modern living relatives of assemblage is a heterogeneous mixture of higher
Buxus pliocenica
,
Populus tremula
, foss.,
Acer
pseudo-
plants with different ecology. Considering the
monspessulanum
and
Juglans acuminata
thrives in specific features of the modern associations (Wolfe Eurasia while
Daphnogene polymorpha
has its clos- 1979; Klötzli 1988;
Song 1983
, 1988) that can est counterparts in Asia and America and
Ziziphus
be related to the Vrysses fossil leaves, the outline
ziziphoides
in Asia. A few taxa have affinities to of a regional vegetation reconstruction has been living Mediterranean species such as
Tetraclinis
and developed, that is characterized by a floristic dif-
Quercus mediterranea
while extant
Parrotia
is a relic ferentiation based on water efficiency and altitude found in
Iran
(Zohary 1947; Wang 1961; Zohary (Gastaldo & Ferguson 1998). & Orshan 1966;
Meusel
et al.
1965
,
1978
; Strahler – a) Riparian paleocoenoses composed of representa- & Strahler 1992; Achhal
et al.
1980;
Quezel 1988
; tives of the wood elements
Salix
sp.
and perhaps Barbero
et al.
1992; Turland
et al.
1993; Arabatzis
Populus tremula
and
Juglandaceae
, as well as
Poaceae 1998
, 2001). The core of the nearest living relatives and
Cyperaceae
herbs. These elements were forming of the fossil taxa grows today in the temperate to a belt of a riparian forest along the banks of inflowsubtropical zone, whereas only a few taxa extend ing streams. Subtropical, partly evergreen elements, to the tropical zone (
Lauraceae
) (Walter & Lieth such as
Daphnogene polymorpha
may have played 1967; Browics 1982; Klötzli 1988). also a certain role in these lowland forests. – b) Sclerophyll forest paleocoenoses consisted in general of microphyllous, broad-leaved taxa TAPHONOMY AND VEGETATION (such as
Quercus mediterranea
,
Quercus
sp.
, possi- ANALYSIS bly
Buxus pliocenica
,
Acer pseudomonspessulanum
,
Ulmaceae
and several
Fabaceae
) and conifers (
Pi-
The Vrysses Formation was most likely deposited in a
nus
, possibly
Tetraclinis
) forming woodland at dry shallow, semi-enclosed marine embayment, bordered lowland areas. by islands and shoals (
Meulenkamp
et al.
1979
). – c) Mixed-Mesophytic forest coenoses. Sclerophyl- The fossil assemblage contains autochthonous and a lous evergreen forests of lowlands were gradually large proportion of allochthonous elements. This is replaced by mixed mesophytic forests of higher elevaconfirmed mainly by the composition of the taphoc- tions along an altitude gradient. The canopy of these oenosis that consists of a mixture of plant remains forests was dominated by deciduous representatives
Zidianakis G.
et al.
)
C
°
(30 of sclerophyllous elements (about 15-52%) reduces temperature
20
the – modern b accuracy) Co-existence taxon of analog this approach correlation of the species. Table. recognized 2 shows the in
annual 10
temperature the Vrysses plant requirements assemblage. A and range their of mean 15-16 annual °C for Mean
0
mean Several annual of the temperature Vrysses taxa is also seem estimated to have. their
0 20 40 60 80
natural habitat today in the Euxino-Hyrcanian Prov-
Percentage of entire-margined taxa ince. These taxa comprise
Parrotia
,
Buxus
,
Juglans
FIG. 6. — Correlation of percentage of species with entire-mar-
and
Acer
. The average annual temperature of the gined leaves in the humid and mesic broad-leaved forests of Hyrcanian region varies from 15°C in the west to eastern Asia with mean annual temperature, after Wolfe (1979). 17.5°C in the east. The temperature of the warmest
---
, percentage of entire-margined taxa in the Vrysses (western month ranges between 28-35°C while that of the
Crete
) plant assemblage.
coldest month ranges from 1.5 to 4°C.
Rainfall varies substantially from
530 mm
in of
Acer
5-lobed,
Populus tremula
,
Ziziphus ziziphoides
, the east to
1350 mm
in the west. In most areas
Juglandaceae
and probably
Acer
pseudomonspessula-
of this region precipitation is all year round with
num
and
Ulmaceae
. In the stratum of lower trees maxima in spring, late fall and winter, the eastern and shrubs a mixture of evergreen (
Buxus pliocenica
parts experience three months of summer drought and
Daphnogene polymorpha
), as well as deciduous (see www.rifr-ac.ir/index_eng.htm). If we take this elements (“
Parrotia
”
pristina
,
Rosaceae
) and conifers region as model for the Neogene climate, a range of (
Tetraclinis
sp.
) seem to have existed. 15-17.5°C for mean annual temperature (MAT), It is also interesting to consider dispersal mecha- and more than
800 mm
for mean annual precipitanisms of the recorded taxa. About 23% of the tion (MAP) is estimated.
identified plant taxa were dispersed either endo- or The average small leaf size of the assemblage exozoochorously by birds or mammals (see
Table 1
) (microphyllous flora) may be not only caused by that implies also a diverse paleofauna. taphonomic processes but also by climatic factors,
such as periodic drought. The relatively high proportion of sclerophyllous xeromorphic elements PALEOCLIMATE RECONSTRUCTION in the plant assemblage may confirm the latter hypothesis.
Analyses of Neogene plant assemblages are in gen- On the basis of physiognomic features, leaf reeral a very useful tool to extract proxy data for mains of Vrysses are divided in three main catpaleoclimatic reconstructions. We tried to apply egories: 1) a broad-leaved deciduous component two different methods in order to estimate climatic which predominates with approximately 37-59%; parameters of the broader Vrysses area during the 2) broad-leaved evergreens (laurophyllous taxa) reach late Miocene: the leaf margin analysis and the co- about 11-30%; and 3) the sclerophyllous compoexistence approach. However because of the lim- nent with the relatively high proportion of 15-52% ited number of taxa and the unresolved taxonomic (
Table 1
). The broad-leaved deciduous elements problems due to poor preservation of the material are characteristic for humid, temperate climatic only rough estimations were possible. conditions, while the broad-leaved evergreen taxa – a) Leaf margin analysis. The proportion of taxa for humid, subtropical and sclerophyllous taxa for with entire-margined species in the Vrysses plant sub-humid, subtropical conditions (Berger 1954; assemblage reaches 48% (
Table 1
), suggesting a Kovar-Eder
et al.
2006).
mean annual temperature of 15-16°C (Zidianakis Thus, the Vrysses plant assemblage may indicate
et al.
2004) (Fig. 6). The relatively high percentage a warm-temperate, humid climate for the late
Miocene leaf assemblage from Vrysses, western
Crete
TABLE 2. — Mean annual temperature (MAT) requirements for nearest living relatives (NLR) of taxa of the Vrysses (western Crete) assemblage. Data are taken from Palaeoflora Database (Utescher & Mosbrugger 1999).
Taxa NLR MAT of NLR (°C)
Tetraclinis
sp.
Tetraclinis articulata
15.60-19.90
“
Parrotia
”
pristina
Parrotia persica
13.60-15.80
Juglans acuminata
Juglans regia
10.10-17.30
Quercus mediterranea
Quercus ilex
,
Quercus coccifera
11.70-19.50
Populus tremula
, foss.
Populus tremula
1.60-19.00
Buxus pliocenica
Buxus sempervirens
9.70-17.30
Acer pseudomonspessulamum
Acer pseudomonspessulanum
, 9.60-20.50
Acer sempervirens
Ziziphus ziziphoides
Ziziphus sinensis
7.30-27.60
Miocene. The existence of a dry period during the flora was determined using nannoplankton and year is probable, although this seasonality was clearly dinoflagellate cysts as late Miocene (7.7-8.6 Ma; not yet as pronounced as today. Using the climatic
Sachse & Mohr 1996
) and is thus slightly older than classification of Köppen (1931), the Vrysses paleo- the Vrysses flora. As it was expected, the Makrilia climate is most likely ranging between Cfa (warm assemblage shares many arboreal taxa with the temperate with warm, humid summers) and Csa Vrysses flora. More than 70% of the taxa identified (warm temperate with warm, arid summers) cli- in Vrysses exist also in Makrilia. Subtropical lauromates. The climatic change from pre-late Miocene phyllous elements, such as
Daphnogene polymorpha
, summer wet climates to summer dry climates was deciduous elements, such as
Acer
5-lobed,
Populus
, possibly caused by the successive westward retreat of
Salix
and
Ziziphus
, conifers, such as
Tetraclinis
,
Pinus
the Paratethys between the Oligocene and the mid- and xeromorphic taxa, such as
Quercus mediterranea
, dle Miocene (Popov
et al.
2004) and the changing
Acer pseudomonspessulanum
and
Buxus pliocenica
are topography of eastern Asia (
Sachse 1997
). common in both floras. However, the Makrilia assemblage is more diverse than the one from Vrysses.
Considering the composition of the Makrilia flora
DISCUSSION it clearly contains relatively more deciduous taxa than the Vrysses plant assemblage. However, it is
The discovery of a new Miocene leaf assemblage in not clear whether this is a real signal or the result of the eastern Mediterranean realm offers the opportu- taphonomic processes or in the nearby hinterland nity for a better understanding of the floristic and of the Vrysses area floras of high elevations might climatic development of this region. Neogene floras have simply not existed.
occur in central and northern
Greece
, as well as in The late Miocene floras of continental
Greece
several
Aegean
islands including
Crete
(Knobloch & (Vegora, Likudi, Prosilion, Lava) and other islands Velitzelos 1987; Velitzelos & Gregor 1990; Sachse (Chios, Kerkyra) consist mainly of conifers and & Mohr 1996, Velitzelos
et al.
2001). If the Vrysses deciduous broad-leaved elements, such as
Ginkgo
, plant assemblage is compared with the known Greek
Taxodiaceae
,
Pinaceae
,
Sassafras
,
Alnus
,
Fagus
, deplant assemblages on the basis of physiognomy and ciduous
Quercus
,
Populus
,
Zelkova
,
Ulmus
and floristic composition, the closest match seems to
Acer
(Velitzelos
et al.
2001)
. Obviously, not only be with the late Miocene assemblage of Makrilia, on
Crete
but in entire
Greece
, the exotic elements eastern
Crete
(
Sachse 1997
, 2004). of the early Miocene were gradually replaced by The plant macro-remains of Makrilia occur in deciduous broad-leaved trees and shrubs, forming semi-pelagic silty layers and represent a plant assem- forests that resembled modern mixed mesophytic blage of which 94% of the leaf taxa are microphyll woods of Eastern Asia, the
Colchis
and the Caspian or nanophyll (
Sachse 1997
). The age of the Makrilia area. This change most likely indicates a decrease
Zidianakis G.
et al.
of mean annual temperature during the Miocene
pliocenica
and
Daphnogene polymorpha
), deciduous in
Greece
, as it seems to have happened in all the (“
Parrotia
”
pristina
,
Rosaceae
) and conifers (
Tetraother
Tethys and Paratethys areas (Gregor 1990;
clinis
sp.) occurred in the lower strata.
Kovar-Eder 1987). The floristic composition and physiognomy of the Furthermore, the relative increase of xeromorphic assemblage, as well as a comparison with published taxa, such as evergreen oaks (
Quercus drymeja
Unger, Miocene Greek
assemblages indicate a warm-tem-
Q. mediterranea
and
Q. sosnowskyi
Kolakovskii
), as perate, humid climate for this period, possibly with well as
Acer pseudomonspessulanum
and
Chamaerops
not very pronounced dry periods for the end of the
humilis
(Velitzelos & Schneider 1978; Velitzelos Tortonian in the eastern Mediterranean area.
et al.
2001; Kvaček
et al.
2002) may indicate the beginning of weak drought periods during the last
Miocene in
Greece
and the onset of the Csa climate
Acknowledgements
type
sensu
Köppen (1931) (warm temperate with We would like to thank Prof. M. Mylonas, director arid summers). The existence of summer drought of the Natural History Museum of
Crete
, for his may be more obvious in the plant assemblage of helpful suggestions, and Dr. A. Nikolaidis, direc- Vrysses indicating already the beginning Messinian tor of the Mediterranean Agronomic Institute of salinity crisis (Hsu
et al.
1973). Chania, for providing samples from this institution. For the free access to the Berlin Herbarium of the Botanic Garden for comparative studies we thank CONCLUSIONS Dr. R. Vogt. This work would not have been possible without the help of the staff of the Natural The upper Tortonian (6-7.5 Ma) Vrysses plant as- History Museum of
Crete
, A. Kagiambaki, M. Trisemblage consists of a mixture of leaf taxa that had kali, G. Iliopoulos, E. Nikolakakis and P. Lyberakis. most likely originally grown in riparian to possibly Also, comments of Dr. D. Franceschii, Dr. S. Sen mid altitude forests. and of an anonymous reviewer greatly improved The Vrysses assemblage represents a microphyll this paper.
flora consisting of 31 taxa. Remains of
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