Palaeofl ora and climate of lignite bearing lower−middle miocene sediments in the Seyitömer and Tunçbilek Sub-basins, Kütahya province, Northwest Turkey

Lower–Middle Miocene successions from Kütahya province (Seyitömer and Tunçbilek sub-basins) were analyzed to reconstruct the palaeoclimate and vegetation of the period. A rich coniferous forest formed the dominant components among sporomorphs and consisted mainly of undeterminable Pinaceae, Pinus haploxylon type, Pinus diploxylon type, Picea, Cedrus, Cathaya, Keteleeria and Podocarpus.

Palaeofl ora and Climate of Lignite-bearing Lower−Middle Miocene Sediments in the Seyitömer and Tunçbilek Sub-basins, Kütahya Province, Northwest Turkey

Dokuz Eylül University, Department of Geological Engineering, Buca,

Tınaztepe Campus, TR−35160 İzmir, Turkey

3

Institute für Geologie, Nusalle 8, 53115 Bonn, FRG, Germany

4

Senckenberg Research Institute and Natural Museum, Senckenberganlage 25,

D-60325 Frankfurt am Main, Germany

Received 11 August 2011; revised typescripts received 13 November 2010 & 14 January 2011; accepted 15 February 2011

Abstract: Lower–Middle Miocene successions from Kütahya province (Seyitömer and Tunçbilek sub-basins) were analyzed

to reconstruct the palaeoclimate and vegetation of the period A rich coniferous forest formed the dominant components

among sporomorphs and consisted mainly of undeterminable Pinaceae, Pinus haploxylon type, Pinus diploxylon type,

Picea, Cedrus, Cathaya, Keteleeria and Podocarpus Pollen data also indicate the presence of some fern taxa (Osmunda,

Polypodiaceae, Schizeaceae) Selaginella and Lycopodium Cupressaceae frequently occur in all samples from both basins, and form part of the mixed forest assemblage Two woody plants Castanea-Castanopsis and evergreen Quercus

sub-predominate in the assemblages as well Th e reconstruction of palaeoclimate is based on the Coexistence Approach (CA) method and documents subtropical climate with cyclic variation in the sediment sequence Th e vegetation does not show a clear change through the sequences studied Small changes in the quantity of pollen taxa (e.g., thermophilous and arctotertiary) indicate small-scale cyclic temperature and precipitation oscillations during the deposition of the sediments Using CA, seven palaeoclimate parameters were estimated Quantitative palaeoclimate data indicate mean annual temperatures of 17−18°C, with winter temperatures of 8−10°C and summer temperatures of 27−28°C Mean annual precipitation exceeded 1200 mm Th e driest month precipitation was between 20 and 25 mm, the wettest month precipitation most frequently ranged between 200 and 250 mm and the summer precipitation was 100−150 mm Th ese values imply that the Kütahya area was warm and received more precipitation during the Early−Middle Miocene than in modern times.

Key Words: Lower–Middle Miocene, Seyitömer, Tunçbilek, Kütahya, palaeoecology, palynology, sub-basin

Seyitömer ve Tunçbilek Ast-Havzalarında Linyit İçerikli Alt−Orta Miyosen Tortullarının Paleofl orası ve İklimi, Kütahya Alanı, Kuzeybatı TürkiyeÖzet: Paleoiklim ve vejetasyonu yeniden kurmak için, Kütahya alanının (Seyitömer ve Tunçbilek ast-havzaları) Alt−Orta

Miyosen istifl eri incelenmiştir Sporomorfl arın arasında, zengin bir konifer ormanı baskın bileşenleri oluşturmaktadır

ve başlıca ayırtlanmamış Pinaceae, Pinus haploxylon tip, Pinus diploxylon tip, Picea, Cedrus, Cathaya, Keteleeria ve

Podocarpus’tan meydana gelmektedir Polen verileri, bazı eğreltilerin (Osmunda, Polypodiaceae, Schizeaceae) Selaginella

ve Lycopodium’ların varlığını da belirtir Cupressaceae her iki ast-havzanın tüm örneklerinde sıklıkla bulunur ve karışık orman topluluğuna katılırlar İki odunsu bitki, Castanea-Castanopsis ve her daim yeşil Quercus da topluluklarda

baskındır Paleoiklimi yeniden kurma Birarada Olma Yaklaşımı (KA) yöntemine dayalıdır ve istift e devirsel değişimli ılık-sıcak olduğunu belgelemektedir Çalışılan istifl er boyunca bitki örtüsü belirgin bir değişim göstermemektedir Polenlerin (örnek, termofi l ve aktotersiyer) miktarındaki küçük değişimler, tortulların çökelimi süresince, küçük ölçekli devirsel sıcaklık ve yağış miktarlarındaki dalgalanmaları belirtmektedir KA yöntemine göre, yedi iklimsel değişken elde edilmiştir Sayısal paleoiklimsel veriler, yıllık ortlama sıcaklığın 17−18°C, kış sıcaklıklarının 8−10°C ve yaz sıcaklıklarının 27−28°C olduğunu belirtir Yıllık yağış miktarı 1200 mm’yi aşmaktadır En kurak ay yağış miktarı

Western Turkey is dominated by large

E–W-trending grabens and a number of approximately

N–S-trending grabens In the Kütahya area,

lignite-bearing sediments crop out in two locations, namely

the Seyitömer and Tunçbilek basins that were fi lled

with continental sediments (Figures 1 & 2) Neogene

lignite-bearing lacustrine deposits are also widespread

in western Turkey (e.g., Soma, Çanakkale-Çan,

Konya-Ilgın, and Uşak) Most of the studies in these

areas have been made in order to solve their tectonic,

stratigraphical and sedimentological problems

Initial studies were to establish palynostratigraphic

aspects of these lignites (Akyol 1963, 1964, 1968;

Nakoman 1965; Benda 1971; Arslan 1979; Benda &

Meulenkamp 1979) but later these were extended to

characterize clastic sequences, especially the Miocene

deposits in western Turkey, improving knowledge of

vegetation and climate during this time (e.g., Ediger

1990; Akyol & Akgün 1990, 1992; Yavuz et al 1995;

Ediger et al 1996; Akgün et al 2007; Yavuz-Işık 2007;

Kayseri & Akgün 2008)

Th e Seyitömer and Tunçbilek open pit mines are

in one of the biggest lignite-producing areas Initial

studies to shed light on the geological divisions around

Kütahya were made by Nebert (1960) and Baş (1983)

Th e fi rst palynological studies were carried out by

Nakoman (1968) and Özcan (1987) in the Seyitömer

sub-basin, but it is diffi cult to determine the botanical

affi nity of their pollen since they used morphological

classification for most of the pollen and spores Th e

fi rst comprehensive palaeoecological study in the

Seyitömer sub-basin was made by Yavuz-Işık (2007)

who recognized two diff erent pollen zones indicating

two diff erent periods: late Early Miocene (Zone 1)

and Middle Miocene (Zone 2) Zone 1 is represented

by a high percentage of Pinus and Cedrus Deciduous

Quercus and evergreen Quercus occur abundantly

in Zone 2 Th e percentage of Taxodiaceae decreases

from Zone 1 to Zone 2 Th e author also indicated

that discrepancies between Zone 1 and Zone 2 may

refl ect the global Middle Miocene cooling In this study we enlarge the palynological data by analysis

of new samples from the whole Lower–Middle Miocene sequence of the Seyitömer sub-basin and lower lignite seam of the Tunçbilek sub-basin within

a stratigraphic framework since the palaeofl ora has already been investigated by other researchers Th e present paper provides an analysis of the Early–Middle Miocene vegetation evolution on the basis

of pollen and leaves Th e palynological data have also been analyzed using the Coexistence Approach (CA) in order to obtain quantitative palaeoclimatic data, and to understand the link between the climate changes and vegetation (Mosbrugger & Utescher 1997)

by Neogene lacustrine deposits

In the Seyitömer sub-basin, the Miocene sequence includes, in ascending order: a lower clastic unit, a claystone-mudstone unit, a lower lignite seam, an organic shale unit, an upper lignite seam, a silicifi ed limestone unit, and a clayey limestone unit: all these units belong to the Seyitömer Formation (Sarıyıldız 1987) that is unconformably overlain by an upper clastic unit named the Kocayatak Tepe Formation (Sarıyıldız 1987) (Figure 1a) Th e lower clastic unit mainly comprises greenish unconsolidated

20 ve 25 mm arasındadır Nemli ay yağış miktarı büyük çoğunlukla 200 ve 250 mm ve yaz yağış miktarı 100−150 mm arasındadır Bu veriler, Kütahya alanının Erken−Orta Miyosen süresince sıcak ve günümüzdeki iklimsel kayıtlardan daha fazla yağış aldığını göstermektedir.

Anahtar Sözcükler: Alt–Orta Miyosen, Seyitömer, Tunçbilek, Kütahya, paleoekoloji, palinoloji, ast-havza

conglomerates and yellowish planar and cross-bedded

sandstones that are overlain by a greenish medium to

poorly bedded claystone-mudstone unit, up to 50 to

60 m thick, over lignite seam, ranging from 2 to 15 m

thick, is medium to thick bedded with claystone and

silicifi ed interlayers Th e organic shale unit generally

consists of grey shale and siltstone, containing leaves, mammals and ostracoda Th e upper lignite seam, which is non-commercial, is made up of an alternation of lignite, silicifi ed limestone and clayey lignite (Figure 1) Th e clayey limestone unit consists

of consolidated tuffi te and clayey limestone

upper lignite seam

organic shale with leaf and ostracod fossils

lower lignite seam

greyish mudstone - claystone

lower clastics ultramafics, metamorphics, olistostroms

lower lignite seam

upper lignite seam

Figure 3a

upper ligniteseam siliceouslimes

tone,clayeylimestone

lower lignite seam

Figure 1 (a) Generalized stratigraphic column of the Seyitömer sub-basin (Sarıyıldız 1987) (b) Field photograph showing the

Aslanlı section (c) Field photograph showing the Tumulus section Open circles show beginning of the sections.

In the Tunçbilek sub-basin, terrestrial and

lacustrine sediments contain half-cemented

conglomerates, claystones of fl uviatile origin

showing a fi ning-upward sequence, marls, siltstones,

sandstones, lacustrine limestones, and lignite,

located in the lower and upper parts of the sequence

(Figure 2a, d) Th e main lignite seam, from 7 to 15

m thick, includes claystone interlayers as well Th e

overlying claystone-marl alternations, over 300 m in

total thickness (Figure 2c), are generally horizontal

bedded but locally folded, and include poorly

preserved leaves Th e sequence ends with lacustrine

limestones containing chert nodules and thin lignite

seams (Figure 2a) All these lithologies belong to the

Tunçbilek Formation described by Baş (1983) Upper

Miocene to Pliocene volcanic rocks, lava, tuff and

agglomerate of the Domaniç Series (Nebert 1960)

unconformably overlie the Tunçbilek Formation

(Figure 2b)

Özcan (1987) studied the palynology of

lignite-bearing sediments exposed in the Seyitömer

sub-basin, and also identifi ed a vertebrate fossil,

Aceratherium tetradactylum (Lartert) Hooijer which

suggested a Vallesian age (Late Miocene) Later

Kaya (1993) described mammalian fauna including

Begertherium grimmi (Heissig)

(Rhinocerotidae-Perissodactyla) and Moropus elatus (Marsh)

(Chalicotheriidae-Perissodactyla) in the lower

lignite seam, and suggested a late Middle Miocene

age of the Seyitömer sub-basin In contrast, a late

Early Miocene–Middle Miocene (MN 4-8) age was

suggested by Saraç (2003) based on mammalian data

Recent observations indicate that sediments in the

Seyitömer sub-basin were deposited during the early

Langhian (Middle Miocene) based on the mammal

data (T Kaya, personal communication) But

although comprehensive mammal data have been

obtained from the Seyitömer sub-basin to date, the

age is still debated Also, as no age data are available

for the Tunçbilek sub-basin, better age control of the

sedimentary sequences in the basin is needed

Material and Methods

Th e study was carried out at three sections in two

diff erent lignite open pit mines Two stratigraphic

sections were measured from the Seyitömer

sub-basin (Figure 1a, b) One, north of Aslanlı village,

located on the northern side of the mine, is named here the Aslanlı section, up to 33 m in total thickness (Figures 1b & 3a) Th e other, measured east of the archaeological excavation area, is named here the Tumulus section (total thickness around 37.40 m) (Figures 1c & 3b) During the fi eld studies, palynological samples of the Tumulus section were collected from the lower and upper lignite seams, since the organic shale includes sandy and coarse-grained clastics which are not suitable lithologies for palynological analysis (Figures 3b & 4b) Additionally,

a section from the lower lignite seam, approximately

17 m thick, was sampled in the Tunçbilek sub-basin (Figure 3c) A total of 171 samples were collected from the 3 outcrop sections, where palynologically productive fi ne-grained sediments and lignites are well exposed 139 samples were collected from the Seyitömer sub-basin (Figure 3a, b) 32 samples were taken from the lower lignite seam of the Tunçbilek sub-basin (Figure 3c) Samples which were barren or contained very few pollen grains were not included

in the analysis Quantitative data are thus confi ned

to 73 productive samples, of which 59 came from the Seyitömer sub-basin (Figure 4) and 14 samples from the Tunçbilek sub-basin (Figure 5) Distinguishable genera and families were identifi ed In most cases, the spore and pollen sums were based on more than

200 grains (Figure 5) TILIA soft ware was used to calculate the sporomorphs, and TILIAGRAPH was used to plot the sporomorph diagrams (Grimm 1994) Selected palynomorphs are illustrated on Plates 1–4

For the recovery of palynofossils, mineral components were fi rst removed with specifi c acids: the silicates were removed with 40–60% HF; carbonates, commonly found as calcite, were dissolved in HCl; sulphates, sulphides and carbon contents are soluble in concentrated HNO3 Slides prepared from the productive samples were investigated under the microscope for qualitative and quantitative assessment of palynofossils Th e

CA method has been used to reconstruct the palaeoclimate (Mosbrugger & Utescher 1997) Th e application of the CA was facilitated by the computer program ClimStat and the database Palaeofl ora that includes nearest living relatives of more than 3000 Tertiary plant taxa, together with their climatic requirements, which are derived from meteorological stations located within the distribution areas of the

whitish clayey limestone, limestone

reddish - brownish basalt flows

Figure 2 (a) Generalized stratigraphic column of the Tunçbilek sub-basin (Nebert 1960) (b) Field photograph showing the

boundary between the tuff and basalt (c) Field photograph showing marl-dominated lithologies and volcanics (d)

Field photograph showing lower lignite seam including claystone interlayers and also syn-sedimentary fault.

Seyitömer Formation

Organicshale Upperlignite

Lower-claystone lignite Lower-claystone marl silicified

Middle Miocene

P inaceae

Picea P inus

Pinus

haploxylon type

P inus

type

C upressaceae

Sequoia

Sapotaceae

R hus

ev re en

Q u rc u

deciduous

Q uercus

Q uercus

C arpinus C orylus Betula

O lea Fagus

P hillyrea

M oraceae

Ilex

E ricaceae

O strya

T ilia

A cer

C ycadaceae

A lnus S alix/P latanus U lm

Z elkova Liq uidam bar

Pterocarya C arya

T axodium N yssa M yrica

P otam ogeto n

S parga nium yperaceae C

Brassicaceae A steraceae Po lygalaceae

Artem isia

P oaceae

C henopodiaceae

E phedra

O sm unda

S chizaceae

ngelhardia R eveesia

Araliaceae A recaceae

Castanea/

Castanopsis Castanea/

Castanopsis

Lowerlignite seam Upperlignite seam

taxa (see also information given on the web site

www.palaeofl ora.de) Th e resolution increases with

the number of taxa included in the analysis and it

is in fl oras with 10 or more taxa for which climate

parameters are known (see Mosbrugger & Utescher

1997 for detailed discussion) Hence, the following

seven palaeoclimate climate parameters have been

considered: MAT– mean annual temperature (°C);

CMT– mean temperature of the coldest month (°C);

WMT– mean temperature of the warmest month

(°C); MAP– mean annual precipitation (mm); HMP–

precipitation of the wettest month (mm); LMP–

precipitation of the driest month (mm); WMP–

precipitation of the warmest month (mm)

Floral Analysis

Leaf Data

Although both plant megafossils and microfossils

occur in the sediments of the Seyitömer and

Tunçbilek sub-basins, this work focuses mainly

on the sporomorph assemblages Moreover, the

lignite deposits of Seyitömer and Tunçbilek open pit

mines include those poor in leaf assemblages In the

Seyitömer and Tunçbilek localities open pit mines

have been found fossil plant remains of Lauraceae,

Taxodiaceae, Fraxinus, Glyptostrobus, Engelhardia

and Zelkova Glyptostrobus are oft en dominant One

specimen includes a shoot of Pinaceae Additionally,

Nebert (1962) determined a poor leaf fl ora including

Taxodium distichum (L.) Rich.), Myrica lignitum

(Unger) Saporta., Nerium sp., Apocynophyllum

sp and cf Phragmites sp from the Tunçbilek

sub-basin Later, Mädler & Steff ens (1979) described the following leaf fl ora from the Tunçbilek sub-basin:

Taxodium dubium (Sternberg) Heer, Sciadopits tertiaria (Menzel), Fagus attenuata (Goeppert), Diospyros brachysepala Al Braun and Laurophyllum pirimigenium (Unger).

Palynological Data

palynological data obtained from the Aslanlı and Tumulus sections are presented separately (Figure 4a, b)

total of 53 diff erent spore and pollen taxa belonging

to pteridophytes, gymnosperms and angiosperms (Figure 4a, b) Based on changes in certain sporomorphs of the samples, the pollen diagram can be interpreted in three parts: lower lignite seam, organic shale and upper lignite seam

Lower Lignite Seam (0–3.80 m of the Measured Section; Samples 07/103–119 from the Lower part

percentages of the fern spore Polypodiaceae (range

5 to 52%), undeterminable Pinaceae (range 6 to 17.4%) and Cupressaceae (range 4.2 to 21.1%)

Castanea-Castanopsis is present in comparatively

lower percentages (range 1.1 to 4.9%) Also, lower

Figure 6 Coexistence intervals derived from the samples of the Aslanlı section (a), Tumulus section (b) of the Seyitömer sub-basin,

and lower lignite of the Tunçbilek sub-basin (c) MAT– mean annual temperature; CMT– mean temperature of the coldest

month; WMT– mean temperature of the warmest month; MAP– mean annual precipitation; HMP– precipitation of the wettest month; LMP– precipitation of the driest month; WMP– precipitation of the warmest month.

CMT [C]

WMT [C]

MAP [mm]

800 1100 1400 1700

HMP [mm]

300

50

LMP [mm]

30

WMP [mm]

CMT [C]

WMT [C]

MAP [mm]

HMP [mm]

LMP [mm]

WMP [mm]

MAT

[C]

CMT [C]

WMT [C]

MAP [mm]

HMP [mm]

LMP [mm]

WMP [mm]

percentages were reported of Osmunda (range

0.2 to 5%), Cyrillaceae-Clethraceae (range 1 to

4%), evergreen Quercus (range 3.2 to 13.8%) and

Potamogeton (range 0 to 5.8%) (Figure 4a).

Organic Shale (3.80 − 22.00 m of the Measured

Section; Samples 07/122-192 from the Middle Part

dominated by undeterminable Pinaceae (range 2.2

to 21 2%), Picea (range 1 to 20.1%), Pinus (range

0 to 15.2%), and Ostrya (range 1 to 22.3%) A

considerable increase of evergreen Quercus (range 4.1

to 41.3%) is recorded here (Figure 4a) Cupressaceae

show a reduction trend from the lower part of the

organic shale side of it Potamogeton is recorded as

single grains Additionally, a few deciduous Quercus,

Moraceae, Asteraceae, Poaceae, Caryophyllaceae

appear (Figure 4a)

Upper Lignite Seam (22.00 − 33.10 m of the

Measured Section; Samples 07/193-218 from the Upper

percentages of Polypodiaceae (range 0.8 to 80.8%),

Picea (range 3 to 18.2%), Pinus (range 1 to 14.8%),

Cyrillaceae-Clethraceae (range 0.2 to 17.3%), and

evergreen Quercus (range 9.8 to 25 3%) Deciduous

Quercus also occurs, but in minor amounts (Figure

4a) Relatively high percentages of Fagus (range 0.3

to 8.8%) have been reported Additionally, Cedrus,

Quercus spp., Corylus, Carpinus, Carya, Olea,

Pterocarya and Ulmus are constantly present along

the Aslanlı section It is also clear that Polypodiaceae

are abundant in the lower and upper lignite seams,

but are scarce in the organic shale part

Tumulus Section

Lower Lignite Seam (0−7.42 m of the Measured

Section; Samples 07/06-20 from the Lower part of

the Seyitömer Formation) – Palynological data of

the Tumulus section are similar to the palynological

data of the Aslanlı section Th ey show abundant

Polypodiaceae (range 4.2 to 32.1%), Osmunda (0

to 29.8%), undeterminable Pinaceae (range 3.2 to

18.1%), Cathaya (range 0.3 to 11.8%), evergreen

Quercus (range 15.1 to 19.7%), Potamogeton (range

1 to 31.3%) and Sparganium (range 0.8 to 13.2%)

Cupressaceae, Alnus, Salix/Platanus and Nyssa

are constantly present, but in minor proportions

Deciduous Quercus and Zelkova are temporarily

missing between 0 to 1.5 m of the section (Figure 4b)

Upper Lignite Seam (20−29.8 m of the Measured Section; Samples 07/23-31 from the Upper part of

percentages of Polypodiaceae (range 2.1–71.3%),

undeterminable Pinaceae (range 0.8 to 18.2%),

Cedrus (range 0.3–19.6%), Cyrillaceae-Clethraceae

(range 0.4–19.8%), and evergreen Quercus (range 3.1–22.8%), Cupressaceae and Taxodiaceae slightly

increase (Figure 4b) Th roughout the Tumulus

section, Cedrus occurs constantly, but never above 15% Pollen grains of Cathaya, Engelhardia, deciduous

Quercus, Corylus, Olea, Ulmus and Quercus spp are

52.3%), Osmunda (range 0.3–34.5%), decreasing in

percentage upwards, undeterminable Pinaceae (range

1.2–17.4%) and Cupressaceae (range 0.8–40.3%), Arecaceae (range 0.8–21.1%) and Alnus (range 30.4– 70.3%) Castanea-Castanopsis, evergreen Quercus,

Engelhardia, Ostrya, Taxodiaceae and Pterocarya are

constantly present but scarce (Figure 5)

Palaeovegetation

Th e palaeovegetation reconstruction is based on both the palynological data and some macrofl ora During the deposition of sediments in both sub-basins, there was a heterogeneous mixture of plants based

on diff ering autecologies and the present ecological requirements of the nearest living relatives of the fossil taxa (e.g., Hofmann & Zetter 2005; Kovar-Eder

vegetation profi le consists of four varied types, beginning near the lake:

communities are typifi ed by Sparganium,

Cyperaceae and Potamogeton Th e presence

of Potamogeton, common in the lower and

upper lignite seams, indicates a water depth

of approximately 2–3 metres (Cook 1996) In

contrast to their occurrences in the Seyitömer

sub-basin, they are scarcer in the lower lignite

seam of the Tunçbilek sub-basin Freshwater

marshes formed adjacent to standing – and

fl owing – water herbaceous plants and grasses

Th e thin-walled pollen of Cyperaceae occur

in minor amounts Herbs like Asteraceae,

Brassiaceae and Polygalaceae also existed, but

probably over limited areas as there are only

low percentages of these taxa in the pollen

diagrams (Figures 4 & 5)

(2) Th e azonal riparian-swampy forest along

the lake borders and discharging streams,

consisting of Taxodium, Myrica, which thrived

with Nyssa and hygrophytic broad-leaved

deciduous forest (Salix/Platanus, Liquidambar,

Carya, Pterocarya and Alnus) and helophytic

herbs such as Osmunda, Polypodiaceae,

Corylus, Araliaceae, Arecaceae, Ericaceae

and Ilex in the undergrowth, was widespread

Azonal woody taxon, representing possibly

riparian and swampy forest, is indicated by

Taxodium dubium (Sternberg) Heer, Myrica

lignitum (Unger) Saporta and Glyptostrobus

(macrofl oral record) Some Pinaceae (Cathaya,

Pinus diploxylon type and probably Sequoia)

grew at the swamp margin Components with

local ecological and edaphic requirements are

present, especially the cosmopolitan family

Cupressaceae

(3) Mesophytic woody taxa occurred on the

lake margins and the lower mountain slopes,

as did Alnus, Betula, deciduous Quercus,

Fagus attenuata (Goeppert), Ulmus, Zelkova

(pollen and leaf), Fagus, Carya, Engelhardia

(pollen and leaf), Rhus, Acer and Tilia

Species of Castanea, Corylus, Ostrya and

Carpinus also dominated in these forests

Within this vegetation belt, thermophilous

elements were common and consisted mainly

of Engelhardia and Cyrillaceae-Clethraceae

In contrast, Castanea-Castanopsis, Reveesia,

Sapotaceae, Schizeaceae and Cycadaceae occurred sporadically There are some

Zelkova, Betulaceae, Juglandaceae, Lauraceae

(leaf), Nyssa and Engelhardia and only a

few Mediterranean taxa, such as evergreen

Quercus, Phillyrea, Olea and Nerium (leaf)

Conifers also played an important role in the

mid-high altitude forest cover, with Sequoia, many undeterminable Pinaceae, Pinus,

Cedrus, Keteeleria, Picea, Cupressaceae and

some Podocarpus and Larix (Larsson et al

2010) Among the conifers, undeterminable

Pinaceae, Pinus diploxylon and haploxylon types and Cathaya were most abundant,

but unfortunately they do not allow an interpretation of the palaeoecology since they can travel long distances by wind or water (Traverse 1988) Th e presence of Sciadopits tertiaria (Menzel) (leaf) indicates upland vegetation Th e understorey may have included abundant Polypodiaceae

(4) Evergreen sclerophyllous oaks, Olea and

Ephedra, formed subxeric woodland on drier

ground

Generally, mixed coniferous and deciduous broadleaved forests covered larger areas; grass and herbaceous assemblages including Asteraceae, Poaceae, Chenopodiaceae, Caryophyllaceae,

Brassicaceae and Artemisia became diverse, but

in low percentages Th ere is no evidence for the spread of herbaceous vegetation in the area as the assemblages include very low contents of herbs and ferns (Figures 4 & 5)

Additionally, though the palaeovegetation types in the Seyitömer and Tunçbilek sub-basins have almost similar aspects, there are some minor discrepancies between them For instance, Palm

(Arecaceae) (range 0.8 to 21.1%) and alder (Alnus)

(range 30.4 to 70.3%) were abundant in the Tunçbilek sub-basin (Figure 5), but were either relatively scarce

or totally absent in the Seyitömer sub-basin As a detail, the Tunçbilek sporomorph community is comparable to the sporomorph community of the lower lignite seam in the Seyitömer sub-basin, as

there were abundant Polypodiaceae and Osmunda in both assemblages (Figures 4 & 5) Evergreen Quercus

occurs abundantly in the Seyitömer samples but is