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e Late Miocene Mammal Faunas of the Mytilinii Basin,
Samos Island, Greece: New Collection
3. Palynology
by
Chryssanthi Ioakim1) & George D. Koufos2)
I, Chr. & K, G.D., 2009. e Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece:
New Collection. 3. Palynology. — Beitr. Paläont., 31:27–35, Wien.
1) Dr. Chryssanthi I, IG ME , Ol ym pic Vil la ge , En tranc e
C, GR-13677, Acharnae, Athens, Greece. e-mail: ioakim@
igme.gr
2) Prof. George D. K, Aristotle University of essaloniki,
Department of Geology, Laboratory of Geology & Palaeon-
tology, 54124 essaloniki, Greece, e-mail: koufos@geo.auth.
gr
Abstract
e present article is dealing with the study of the mi-
crofloristic assemblages of the Miocene deposits of the
Mytilinii Basin, Samos Island. e deposits are divided
in five formations (Basal Fm, Mavradzei Fm, Hora Fm,
Mytilinii Fm and Kokkarion Fm) and correspond to the
time interval from middle Miocene to Pliocene. New data
obtained from palynological analysis and chronostrati-
graphic correlations based on field work and sampling
in the Mytilinii Basin, were co-evaluated with already
exist ing da ta co nce rning t he lit ho-bios trat ig rap hy o f ma-
rine and continental deposits of Greece. e qualitative
palynological results were complemented by quantitative
pollen based on climatic reconstructions during Vallesian
and Turolian.
Keywords: Late Miocene, Samos, Greece, Palynology,
Palaeoclimatic Reconstruction.
Zusammenfassung
Die vorliegende Studie beschäftigt sich mit den Mikro-
floren des miozänen Ablagerungen des Mytilinii Beck-
ens der Insel Samos. Die Ablagerungen werden in fünf
Formationen geteilt (die Basal, die Mavradzei-, die
Hora-, die Mytilinii- und die Kokkarion Formation), die
zeitlich vom mittleren Miozän bis zum Pliozän reichen.
Die neuen Daten aus der palynologischen Analyse des
Mytilinii-Becken werden verglichen mit den bereits
existierenden Resultaten aus der Litho- und Biostrati-
graphie der marinen und kontinentalen Ablagerungen
Gr ieche nlands . Die qualit ativ en p al ynologisc hen E rgeb -
nisse werden durch quantitative ergänzt, basierend auf
klimatischen Rekonstruktionen des Vallesiums und
Tur o l iu m s .
Schlüsselworte: Obermiozän, Samos, Griechenland,
Palynologie, paläoklimatische Rekonstruktion.
1. Introduction
e Island of Samos is situated in the eastern Aegean
Sea near the coast of Asia Minor and is well known for
its late Miocene mammal faunas since the middle of the
19th century. e known fossiliferous sites are situated in
the Neogene Mytilinii Basin, the palynological record of
which is poorly known. Some palynological data from the
lowest part of the Neogene deposits are given earlier (I-
& S, 1985). e palynogological analysis
of the deposits is important in order to get information
about the palaeoenvironment and the climatic condi-
tions during the Neogene. Although, there are several
mammal fossil sites on Samos, their chronology and
palaeoecological conditions are not clearly defined, as the
collections are old and lack or have limited stratigraphic
control. In 1993 a team of palaeontologists from the
Laboratory of Geology and Palaeontology, University of
essaloniki, led by G. K., started an extended study of
the Neogene deposits of the Mytilinii Basin. e main
goal of this research was to relocate the fossiliferous
sites, to correlate them with the stratigraphy, to excavate
for new fossils and to date the faunas. e palynological
study of the Neogene deposits of the Mytilinii Basin was
also included in this research. e analysis of the pollen
samples was carried out in the laboratory of IGME (In-
stitute of Geological and Mining Exploration) by C.I.
Beitr. Paläont., 31:27–35, Wien 2009
28 Beitr. Paläont., 31, Wien, 2009
2. Stratigraphy of the Mytilinii Basin
e stratigraphy of the Neogene deposits of the Mytilinii
Basin has been studied by various authors and different
approaches have been formulated. A synopsis of all views
on the stratigraphy of the basin is given in K
et al. (this volume, Fig. 1). According to them the lithos-
tratigraphy of the Mytilinii Basin includes the following
formations from bottom to the top:
Basal Formation: It overlies the basement unconformably
and consists of red-brown sands and conglomerates with
gravels and pebbles from the basement. Its age is estimated
as middle Miocene.
Mavradzei Formation: e formation consists of fos-
siliferous limestones with intercalations of lignitic clays
rich in Planorbiidae. In the upper part of the Mavradzei
Fm there is a basalt flow and a series of lahar type vol-
canoclastic sediments. e age of the formation is middle
Miocene-lowermost late Miocene.
Hora Formation: e formation consists of a series of
thick-to-laminated, lacustrine limestones with intercala-
tions of tuffaceous clays. It is dated to Vallesian.
Mytilinii Formation: Fluvio-terrestrial deposits, consist-
ing of reddish-brown volcanoclastic sediments, mainly
sands, sandstones, tuffaceous sands, lenses of conglom-
erates, marly limestones and silty marls. All the known
mammal fossiliferous sites of Samos Island are situated
in the Mytilinii Fm. It is dated to Turolian.
Kokkarion Formation: e younger formation of the
basin represents shallow lacustrine deposits. It consists of
alternated beds of white-yellowish limestones, travertine
limestones with green-brown clays and tuffaceous sands.
e limestones are occasionally fossiliferous with plant
remains, Phragmites and the gastropod Brotia cf. graeca.
It is dated to latest Miocene-Pliocene.
A composite lithostratigraphic column of the Mytilinii
Basin with the different formations and their age according
to K et al. (this volume) is given in Fig. 1.
Figu re 1: Synoptic stratigraphic
column of the Mytilinii Basin
with the lithology and age of the
various formations, indicating
the pollen sampling horizons.
The stratigraphic column was
taken from K et al.
(this volume).
I, Chr. & K, G.D., Palynology.29
3. Methodology
As a great number of the available sediments are not ap-
propriate for the preservation of the pollen grains, the
sampling of the pollen bearing sediments focus on the
Vallesian and the Turolian deposits of the basin. us, 46
samples were collected along the Neogene sedimentary
sequence (Fig. 1). e collected material was treated ac-
cording to the standard palynological method with HCL
and HF acides and sieved through a 6µm nylon mesh.
Slides were prepared using glycerine jelly as a mounting
medium. e palynomorphs were analysed per sample un-
der a binocular transmission light Nikon microscope. ey
were processed according to standard procedures (C,
1974). For each sample at least 200 pollen grains were
counted besides the dominant taxons, because the pollen
concentrations were very low and 85 taxa were identified.
e palynological results are presented as percentages in a
synthetic diagram (Fig. 2). e pollen taxa were organized
in ten groups on the basis of the ecological and climatic
requirements of the present correlatives (S, 1984, 1989).
e floral list is plotted in Table 1. In the pollen spectra
the groups from left to right are:
• Megathermic(-tropical) elements: Mimosaceae, Ru-
biaceae, Euphorbiaceae (SUM A ).
• Mega-mesothermic (subtropical) elements, requir-
Figure 2: Synthetic Pollen-
diagrams of the Mytilinii Basin
deposits correlated with the
stratigraphy. e stratigraphic
column was taken from K-
et al., this volume.
30 Beitr. Paläont., 31, Wien, 2009
ing high (climatic or edaphic) humidity: Taxodiaceae
(Taxodium- type and rare pollen grains of Sequoia-type)
principally and subordinated Nyssa, Palmae, Sapotaceae,
Engelhardia, Myrica (SUM B).
• Cathaya plus Pinus haplostelle type: Cathaya is an Asiatic
gymnosperm tolerating low temperature but requiring
Pinaceae Betulaceae Juglandaceae Plumbaginaceae
Pinaceae sp. Betula Juglandaceae sp. Armeria
Pinus Carpinus Carya Ranunculaceae
P. haplostelle-type Alnus Engelhardia Ranunculaceae sp.
Cathaya Ostrya Juglans Platanaceae
Cedrus Buxaceae Platycarya Platanus
Tsu ga Buxus Pterocarya Polygonaceae
Picea Caprifolliaceae Liliaceae Rumex
Abies Lonicera Liliaceae sp. Rubiaceae
Cupressaceae Caryophyllaceae Loranthaceae Rubiaceae sp.
Cupressaceae sp. Caryophyllaceae sp. cf. Loranthaceae sp. Salicaceae
Ephedraceae Chenopodiaceae - Amaran-
thaceae
Mimosaceae Salix
Ephedra Cyperaceae Mimosaceae sp. Symplocaceae
Taxodiaceae Cyperaceae sp. Myricaceae Symplocos
Taxodium t. Cyrillaceae - Chlethraceae Myrica Tiliaceae
Sequoia t. Cyrillaceae - Chlethraceae sp. Nymphaeaceae Tilia
Aceraceae Ericaceae Nymphaeaceae sp. Typhaceae
Acer Ericaceae sp. Nyssaceae Typha
Araliaceae Nyssa
Araliaceae sp.
Asteraceae - Asteroideae
Artemisia
Asteraceae - Cichoroideae
Asteraceae - Cichoroideae sp.
Euphorbiaceae
Euphorbiaceae sp.
Fagaceae
Castanea
Quercus
Fagus
Quercus ilex t.
Hamamelidaceae
Hamamelidaceae sp.
Liquidambar
Parrotia
Oleaceae
Olea
Fraxinus
Palmae
Palmae sp.
Platanginaceae
Platango
Ulmaceae
Ulmus
Celtis
Vitaceae
Vitis
Osmundaceae
Osmunda
Polypodiaceae
Polypodiaceae sp.
Ta bl e 1: Stratigraphic comparison of previous studies complied
in the Mytilinii Basin, Samos, Greece.
high humidity during all the year (SUM C).
• Mesothermic elements (warm temperate): requiring a
humid climate but tolerating seasonal contrast in humidity
and temperature. e deciduous Quercus is the principal
component with Carya, Juglans, Platycarya, Pterocarya,
Ulmus/Zelkova, Liquidambar, Carpinus, Castanea, Acer,
Fraxinus, Araliaceae etc. (SUM D).
• P inu s and indeterminable pollen grains of Pinaceae, char-
acteristic of various and different ecological and climatic
requirement (SUM E).
Meso-microthermic elements (i.e. mid to high altitude
trees) Ts uga, Cedrus (SUM ST).
• e altitudinal elements Abies and Picea (SUM Z).
Cupressaceae and Alnus, Salix elements with local signifi-
cance (SUM H).
• Mediterranean thermophyllous evergreen elements: Olea,
Quercus ilex-type, Cistus, Buxus (SUM TH).
• Herbaceous elements (Amaranthaceae-Chenopodiaceae,
Compositae, Graminae, Ericaceae, Plantago, Polygo-
naceae, Umbelliferae, Plumbaginaceae, Caryophyllaceae),
including steppe elements (Artemisia, Ephedra, requiring
an arid and sometimes a cooler period, and water depend-
ing plants (Typhaceae, Nympheaceae, etc.) (SUM I).
4. Pollen Flora: Vegetational and Climatic
Inferences
e pollen flora is dominated by arboreal pollen grains
(Fig. 2): Taxodiaceae (Taxodium-type) are principally
followed by Pinus, sometimes Abies, Tsuga and Cedrus,
also associated with the deciduous Quercus taxa, Carya,
Pterocarya and the evergreen Mediterranean taxa (Olea,
Quercus ilex-type etc). Herbs and shrubs (e.g. Poaceae,
I, Chr. & K, G.D., Palynology.31
Amaranthacea, Chenopodiaceae, Gramineae, Ephedra,
Artemisia and other Compositae) are constantly present
but they show a major increase. e sclerophyllous Quer-
cus ilex-type and the deciduous Quercus-type are used as
climatic indicators, based on their climatic requirements.
e first species (Quercus ilex-type) require substantial
winter precipitation and low winter temperatures. On
the contrary, the deciduous taxa of Quercus-type (Quercus
pubescens o r Q. robur) require higher soil water availability
in summer and support also cold and dry winters. e
Mytilinii Basin data set are divided in three pollen as-
semblages.
Mavradzei Assemblage: e palynological associations are
included in five dark clay horizons sampled from an out-
crop that is located very close to the village of Mavradzei.
e thickness of each clay horizon ranges from 1 - 2 cm
and they are intercalated by grey-brown marly beds, which
are rich in fresh-water gastropods (mainly Planorbiidae)
and sandy layers with amorphous leaf impressions (Fig.
1). e total thickness of the sampled section is about 12
m, from where seven samples were collected for pollen
analysis. e megathermic elements, living under moist or
dry conditions, such as the pollen grains of Mimosaceae
are abundant, while the Euphorbiaceae, Rubiaceae are
very rare. e mega-mesothermic and mesothermic ele-
ments requiring high (climatic or edaphic) humidity, like
Taxodiaceae (Taxodium-type) in particular, Engelhardia,
Myrica are well represented. e taxa demanding a warm
temperate climate and relatively high humidity, like the
deciduous Quercus, Carya, Eucommia, Ulmus/Zelkova,
Juglans are also well expressed. e Pinaceae, Pinus, Abies
and Tsuga are abundant. e altitudinal elements are very
poor, while pollen grains of the riparian plants, like Alnus,
Salix a nd C upr essac eae are al so r ecor ded. O n the c ont ra ry,
mo st of the thermo philo us e vergree n ta xa ( Buxus, Quercus
ilex-type, Cistus) are rare. e herb assemblages are domi-
nated by Amaranthaceae-Chenopodiaceae, Gramineae,
Compositae. Other common elements are those of water
depended plants, such as Nymphaeaceae and Typhaceae,
while some fern spores are also recorded. e presence
of this microflora in the Mavradzei Fm suggests warm
and humid climatic conditions during middle Miocene.
Equivalent warm and humid conditions are also assumed
for the Mediterranean during middle Miocene with the
development of mixed mesophytic forests (A et al.,
2003).
Hora Assemblage: e overlying Hora Fm consists of
lacustrine limestones intercalated by tuffaceous clays in
the upper part. Twenty eight samples were collected from
the type section of the Hora Fm, across the road from
Hora to Mytilinii, from the clayey intercalations covering
a thickness of about 120 m (Fig. 1).
e pollen assemblage obtained from this section reveals
a homogeneous vegetation of a more or less widespread
forest consisted mainly of mega-mesothermic Engelhardia,
Taxodiaceae (Taxodium-type principally and sporadically
Sequoia-type), Hamamelidaceae, Nyssa, Myrica, Sapota-
ceae etc.) and mesothermic Carya, Pterocarya, Cathaya,
Ulmus/Zelkova, Acer, Carpinus, Symplocos, Betulaceae,
Fagaceae (Quercus, Castanea, Fagus) elements. e altitu-
dina l b elt s a re cha ra cteriz ed by e nr ichme nt o f mes ot her mi c
trees (Cedrus, Abies, Tsuga), (Fig. 2). Herbs occur in lower
frequencies with respect to the previous palynological
assemblages.
Mytilinii Assemblage: e general lithologic features
of the Mytilinii Fm correspond to volcanoclastic fluvio-
lacustrine sediments which are not favorable for the
preservation of pollen grains. e total thickness of the
formation is ~230 m but only a short part of it (~30 m) was
sampled. is part covers the main mammal fossiliferous
horizons (Fig. 1) from which 14 samples were collected
from the most profitable clayey horizons.
e pollen spectra is characterized by the presence of Cal-
ligonum, Lygeum, as well as the high abundance of more
thermophilic and xerophytic elements like Olea, Cistus,
Quercus ilex-type. Moreover, it includes a high percent-
age of open vegetation plants, like Compositae, Ama-
ranthaceae-Chenopodiaceae, as well as steppe elements
like Artemisia and Ephedra. e absence of aquatic plants
is characteristic (Fig. 2), indicating drier conditions in
comparison to the previous assemblages. is pollen flora
suggests an open vegetation rich in herbs, growing under
dry / warm-temperate climatic conditions. e presence
of these taxa is characteristic of southern Mediterranean
vegetation and they are found today in Southern Spain,
Southern Italy, Sicily, Crete and North Africa
Mo reo ver, du ri ng the beginn in g of lat e Mio cene , the con -
ditions changed, becoming more and more arid. According
to A et al., 1999, 2003; A & A, 2002;
B e t al., 1992; K, 20 06a) this a ri dit y wa s g radu -
ally extended to the Western Mediterranean regions.
5. Conclusions
e palynological study of the Neogene deposits from My-
tilinii Basin allows the recognition of different vegetational
en vi ronme nts cor res po ndin g to t he cl imat ic evo lut io n of the
wider Samos area during Miocene. In the Middle Miocene
the main vegetational features are characterized by mixed
mesophytic forests. is view fits quite well with the known
palaeoenvironmental conditions of the Mediterranean dur-
ing middle Miocene (A et al., 2003). During Valle-
sian, the pollen flora of the Mytilinii Basin was developed
near a deep lacustrine environment under warm-temperate
clima tic c ondit ion s. e f orma tion of t hi n-l am inat ed ma rl y
limestones depends on the water depth and suggests that
the sedimentary sequence of the Hora Fm was deposited
in a deeper lacustrine environment, followed by the clastic
se di ment s of t he My tilini i Fm d ur in g Tur olia n. is m ea ns
a desiccation of the Vallesian lake of the Mytilinii Basin.
In fact, during the beginning of Vallesian, the conditions
in Eastern Mediterranean started to change, being more
arid, as is indicated from the study of the mammal fau-
nas (B et al., 1992; K; 2006a). In spite of the
relatively drier conditions, the lacustrine environment of
the Mytilinii Basin seems to be preserved till the end of
Vallesian. However, in the wider Eastern Mediterranean
32 Beitr. Paläont., 31, Wien, 2009
the Vallesian mammal assemblages suggest a relatively
open environment with bushes, shrubs and grass (B
et al., 1992, 1999; B & K, 1994; M et
al., 2005, 2007; K, 2006a).
e My ti li ni i Fm cor re spon ds to th e Tur olia n t ime-i nte rv al
and its palynological composition (low abundance of Taxo-
diaceae and Pinus, high percentage of herbaceous plants,
including steppe elements and the fairly continuous pres-
en ce of the M edit er rane an scle roph yllo us pl ants), indic ates
a warm-temperate climate, where an open vegetation was
developed. Similar conditions were also proposed from the
study of the dental wear and the analysis of the mammal
fauna found in the Mytilinii Fm, suggesting an open bush-
land with rich grass floor (K et al., this volume-a).
Similar conditions were recorded in the Turolian of Axios
Valley, Macedonia, Greece (B et al ., 1992 ; M
et al., 2005, K, 2006b), as well as in the Turolian
of essaly, Greece (K et al., 2006). e Turolian
palynological record of the Serres Basin (Macedonia,
Greece) indicates the existence of sub-tropical vegetational
development associated with well represented Poaceae. e
latter suggests a more open environment growing under
sub-tropical climate with moderately moist to dry condi-
tions (B et al., 1990 ; K & I,
1989). According to B et al. (1994), also in Turkey an
open environment under arid climatic conditions existed
during the Turolian period. More precisely, the study of
the Turolian mammal faunal assemblages of Asia Mi-
nor indicate that there is a trend to more open and arid
conditions than those of Continental Greece from Early
Turolian (MN 11) to the end of Middle Turolian, MN 12
(K et al., this volume-a). e study of the phytolith
assemblages from the Samos mammal fossiliferous hori-
zons confirms the open character of the landscape during
Turolian (S et al., 2008)
6. Acknowledgements
e excavations on Samos have been supported by the Pre-
fecture of Samos and the “Konstantinos and Maria Zimalis”
Foundation. e Municipality of Mytilinii provided generous
help too. e Natural History Museum of the Aegean offered
us the premises for the preparation and storage of the fossils.
I also want to thank Dr. R. Zetter (Vienna) for his remarks
in order to improve the original manuscript.
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34 Beitr. Paläont., 31, Wien, 2009
PLATE 1
Fig. 1. Pinus diplostelle-type Fig. 7. Alnus sp.
Fig. 2. Pinus haplostelle-type Fig. 8. Corylus sp.
Fig. 3. Tsuga diversifolia Fig. 9. Alnus sp.
Fig. 4. Ulmus/Zelkova Fig. 10 Betula sp., Myrica sp.
Fig. 5. Taxodium sp. Fig. 11. Quercus pubescens
Fig. 6. Alnus sp. Fig. 12. Betula sp.
I, Chr. & K, G.D., Palynology.35
PLATE 1
I, Chr. & K, G.D., Palynology.36
