Miocene palynofloras of the Tınaz lignite mine, Muğla, southwest Anatolia: Taxonomy, palaeoecology and local vegetation change

Miocene palynofloras of the Tınaz lignite mine, Muğla, southwest Anatolia: Taxonomy, palaeoecology and local vegetation change

    Miocene palynofloras of the Tınaz lignite mine, Mu˘gla, southwest Anatolia: Taxonomy, palaeoecology and local vegetation change Johan...

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    Miocene palynofloras of the Tınaz lignite mine, Mu˘gla, southwest Anatolia: Taxonomy, palaeoecology and local vegetation change Johannes M. Bouchal, Serdar Mayda, Reinhard Zetter, Frigeir Gr´ımsson, Funda Akg¨un, Thomas Denk PII: DOI: Reference:

S0034-6667(16)30145-2 doi:10.1016/j.revpalbo.2017.02.010 PALBO 3845

To appear in:

Review of Palaeobotany and Palynology

Received date: Revised date: Accepted date:

4 August 2016 20 February 2017 28 February 2017

Please cite this article as: Bouchal, Johannes M., Mayda, Serdar, Zetter, Reinhard, Gr´ımsson, Frigeir, Akg¨ un, Funda, Denk, Thomas, Miocene palynofloras of the Tınaz lignite mine, Mu˘gla, southwest Anatolia: Taxonomy, palaeoecology and local vegetation change, Review of Palaeobotany and Palynology (2017), doi: 10.1016/j.revpalbo.2017.02.010

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ACCEPTED MANUSCRIPT Miocene palynofloras of the Tınaz lignite mine, Muğla, southwest Anatolia:

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taxonomy, palaeoecology and local vegetation change

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Johannes M. Bouchal1, 2*, Serdar Mayda3, Reinhard Zetter2, Friðgeir Grímsson2, Funda

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Akgün4, Thomas Denk1

Swedish Museum of Natural History, Department of Palaeobiology, Stockholm,

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Sweden

University of Vienna, Department of Palaeontology, Vienna, Austria

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Natural History Museum, Ege University, 35100 Izmir, Turkey

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Dokuz Eylül University, Department of General Geology, 35210 Izmir, Turkey

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Abstract

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*Corresponding author. Email: [email protected]

Middle Miocene deposits belonging to the Eskihisar Formation exposed at the Tınaz lignite mine, Yatağan Basin, Muğla, southwestern Turkey, were investigated palynologically. Nine spores, aplanospores/zygospores and cysts of fungi and algae, seven moss and fern spores, 12 gymnosperm pollen types, and more than 80 angiosperm pollen taxa were recovered from the Tınaz lignite mine section. Three informal pollen zones were recognized: pollen zone 1 corresponds to the main lignite seam and reflects changes from fluviatile to lacustrine depositional settings; pollen 1

ACCEPTED MANUSCRIPT zones 2 and 3 and a transitional zone 2–3 reflect different stages of lake development and a shift from forested (pollen zones 1 and 2) to possibly more open (transitional

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zone, zone 3) local vegetation. Interpreting changes in regional vegetation from pollen zones 1 to 3 is not straightforward because changes in the pollen spectra may be

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affected by changing contributions of airborne and water-transported pollen and spores to the depositional site. In addition, changes of the water table may have promoted

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temporary dominance of herbaceous plants (Amaranthaceae, Nitraria). Palaeobiogeographic relationships of the palynofloras are generally northern

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hemispheric, with many northern temperate tree taxa having modern disjunctions between East Asia and North America (Tsuga, Carya), East Asia and western Eurasia

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(Zelkova), East Asia, North America and western Eurasia (Liquidambar), or restricted

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to East Asia (Cathaya, Eucommia) or North America (Decodon). A few taxa belong to extinct lineages that have complex biogeographic patterns (Engelhardioideae,

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Cedrelospermum). The presence of Picrasma (Simaroubaceae) in the lower lignite

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layers of pollen zone 1 is remarkable, as the botanical affinities of the enigmatic flower Chaneya, present in lower to middle Miocene deposits of Turkey and Central Europe, have recently been shown to be with Picrasma.

Key words: palynology – palaeoenvironment – vegetation development – basin correlation – plant fossils – mammal fossils

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ACCEPTED MANUSCRIPT 1. Introduction

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During the Neogene, the continents achieved their present positions and modern latitudinal and altitudial vegetation zones appeared. The mode and timing of the latter

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process, however, are far from well-understood. For example, although it is claimed by some authors that the modern summer-dry Mediterranean vegetation was

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established rather recently in the Pliocene (Suc, 1984; Velitzelos et al., 2014), others argue that a Mediterranean climate was established by 11 Ma and possibly long before

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(Tzedakis, 2007).

In general, Aquitanian to Langhian vegetation reflects warm and humid to subhumid

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climatic conditions across the Mediterranean region. For example, littoral Avicennia

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mangrove was present during the early and middle Miocene along the coasts of France, Turkey and Bulgaria (e.g., Bessedik et al., 1984; Kayseri-Özer et al., 2014a).

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Another typical feature of early to late Miocene plant assemblages in the eastern

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Mediterranean was the abundant occurrence of evergreen Fagaceae and Lauraceae (e.g., Velitzelos et al., 2014). In contrast, Jiménez-Moreno and Suc (2007) suggested local arid conditions for southern Spain during the middle Miocene, humid temperate conditions prevailed to the north of the Iberian Peninsula. Neogene plant-bearing strata are very common in western Turkey and have been subjected to numerous investigations concentrating on macrofossils (e.g. Engelhardt, 1903; Mädler and Steffens, 1979; Gregor, 1990; Gemici et al., 1990, 1991, 1993; Erdei et al., 2010; Güner and Denk, 2014; Denk et al., 2014, 2015; Güner, 2016) and on dispersed pollen and spores (e.g. Nakoman, 1967; Benda, 1971; Ediger, 1990; Gemici 3

ACCEPTED MANUSCRIPT et al., 1991; Takahashi and Jux, 1991; Akgün and Akyol, 1999; Akgün et al., 2007; Kayseri and Akgün, 2010; Akkiraz, 2011; Kayseri-Özer et al., 2014a, 2014b; Bouchal

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et al., 2015, 2016a). Miocene strata of the Eskihisar Subbasin of the Yatağan Basin in southwestern Turkey

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were studied by Gemici et al. (1990) and Akgün et al. (2007), and most recently Bouchal et al. (2016a) investigated palynomorphs from the Eskihisar lignite mine

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section, situated about 20 km northwest of the Tınaz lignite mine. Both the Eskihisar and the Tınaz lignite mines belong to isolated subbasins of the Yatağan Basin with

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small mountains separating them from the main basin and from each other (BeckerPlaten, 1970). Bouchal et al. (2016a) suggested that generalized “Pollen Bilder”

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(pollen assemblages) (Benda, 1971) alone cannot resolve Burdigalian to Langhian

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stages, as had been claimed in many previous studies. In addition, they warned against correlating radiometric dates obtained for one basin with other basins that show the

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same “Pollen Bild”. Nevertheless, the thorough documentation of dispersed pollen and

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spores is the prerequisite for a meaningful comparison of different palynofloras, and ultimately could result in a better resolved stratigraphic framework for the numerous Miocene plant assemblages (both macro and micro) in Turkey. In addition, whenever available, the vertebrate fossil record should be taken into account for age determination. This study is the second part of an ongoing project investigating palynofloras from three lignite mines in the Yatağan Basin, Muğla, southwestern Turkey: the Eskihisar lignite mine (Eskihisar subbasin, Bouchal et al., 2016a); the Tınaz lignite mine (Tınaz Subbasin, this study); and the Salihpaşalar lignite mine (Yatağan main basin). The 4

ACCEPTED MANUSCRIPT palynological investigations are complemented by studies of macrofossils preserved within marls overlying the main lignite seams in the three mining areas (Güner, 2016).

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The aims of the present study are: (i) to investigate palynomorphs of a 75 m stratigraphic section from the Tınaz lignite mine, Tınaz Subbasin of the Yatağan

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Basin, using a combination of light microscopy (LM) and scanning electron microscopy (SEM) to enable high-resolution pollen/spore identification; (ii) to

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combine new mammal data from the Yatağan Basin with the palynological record to constrain the age of the plant bearing Miocene strata in the Yatağan Basin and

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adjacent areas; (iii) to compare pollen zones recognized in the Tınaz lignite mine section with published pollen zones/floras of the Eskihisar subbasin (including the

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palynoflora from the Yenieskihisar mammal locality), and the nearby Çatakbağyaka

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mammal locality; (iv) to explain the cause of fluctuations in the percentages of pollen and spore taxa between the informal pollen zones; and (v) to assess the biogeographic

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affinities of the palynoflora from the Tınaz lignite mine section.

2. Material and methods 2.1. Geological setting The Tınaz lignite mine is located west of the village Kafaça in the province of Muğla, southwestern Turkey and is part of the Yatağan Basin (Fig. 1). This southeast trending graben reaches a length of 50 km and a width of 15 km. The Neogene sediments filling this basin are up to 600 m thick and comprise coarse- to fine-grained siliciclastic alluvial deposits, lignites, lacustrine carbonates and scattered layers of volcanic tephra. The Yatağan Basin is divided by a metamorphic bedrock horst (belonging to the 5

ACCEPTED MANUSCRIPT Menderes Massif) into a larger main basin and a southeast trending, ca 15 km long and 2 km wide side-basin (Tınaz Basin of Becker-Platen, 1970). The Tınaz lignite mine is

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located at the southern end of this side-basin. Becker-Platen (1970) was the first to describe the lithostratigraphic succession of the

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Yatağan Basin and considered it to be a single formation comprising the Turgut, Sekköy, Yatağan and Milet members (”Schichten” in Becker-Platen, 1970). Atalay

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(1980) revised Becker-Platen’s lithostratigraphy of the basin and distinguished three units: the Eskihisar Formation (including Becker-Platen’s Turgut and Sekköy

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members), the Yatağan Formation, and the Milet Formation. In the area of the Tınaz lignite mine, the exposed sedimentary rocks belong to the Eskihisar Formation. The

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Turgut Member has a thickness of 250–350 m in the main basin and up to 99 m in the

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Tınaz lignite mine area (Becker-Platen, 1970, table 7, borehole cores 56A/66 and 56B/66). The lower part of the Turgut Member consists of reddened alluvial-fan

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deposits (immature, matrix-supported poorly stratified conglomerates, intercalated

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with sandstone and silty sandstone) succeeded by fluviatile deposits (clast-supported, channelized conglomerates alternating with sandstones, coal-bearing siltstones and mudstones) (Atalay, 1980; Becker-Platen, 1970). The excavated lignite seam is stiuated in the transition zone between the Turgut and Sekköy members (Fig. 2). The lacustrine sediments of the Sekköy Member, its thickness ranging between 160 and 200 m in the main basin and reaching up to 70 m in the Tınaz lignite mine area (Becker-Platen, 1970, table 7, bore hole cores 56A/66 and 56B/66), consist of alternating clayey limestones, marls, sandstones, siltstones, claystones and thin coal interlayers (Atalay, 1980; Becker-Platen, 1970). 6

ACCEPTED MANUSCRIPT The boundary between the Turgut and Sekköy members, and hence, the position of the main lignite seam, has been variously interpreted in past studies. In his description of

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the Turgut Member type section, Becker-Platen (1970) regarded the thick main lignite seam as the uppermost part of this member. In Atalay’s (1980) revision of the

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lithostratigraphy of the Yatağan Basin, the lignite seam and the following 40 metres of lacustrine sediments were designated as the upper part of the Turgut Member (this

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boundary was adopted in the scheme of Alçiçek, 2010). Inaner and Nakoman (2005, fig. 3) treated the lignite seam as part of neither the overlying nor underlying

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formations/members; Inaner et al. (2008, fig. 2) place it in the lowest part of the Sekköy Member; Querol et al. (1999) placed the lignite seam at the base of the Sekköy

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Member in their figure of the lithostratigraphic section (Querol et al., 1999, fig. 2), but

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stated in their description of the Sekköy Member “…green to gray-colored, generally fine-grained sediments overlying the main coal seam…” .

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For the present study, the type profiles of both the Turgut and Sekköy members

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designated by Becker-Platen (1970, p. 22 ff. and pp. 26–27) were taken as reference points (see Fig. 2), with the change from fluviatile (non-carbonate siliciclastic) facies to lacustrine (carbonate) facies indicating the boundary between the two members.

2.2. Sampled stratigraphic section A stratigraphic section comprising 75 m of sediment (hereafter Tınaz section; Fig. 2) located near Becker-Platen’s (1970) borehole core locality 56B/66 was investigated. In total, 28 sedimentary rock samples were collected of which 15 were suitable for palynological analysis (detailed sample information is provided in Supporting 7

ACCEPTED MANUSCRIPT Information, Appendix S 1). The Tınaz lignite mine section (Fig. 2) starts below the excavated lignite seam (uppermost part of the Turgut Member) with weakly

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compacted, blueish-grey to dark grey, micaceous clayey siltstones intercalated with thin lignite layers; indeterminable plant debris is common in these layers. The

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following 10 to 12 metres thick unit consists of coal seams intercalated with thin claystone to siltstone layers (1–5 cm) in its lower part, replaced in the upper part by

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light-grey to blueish-grey marls and clayey limestone (5–50 cm). This change from siliciclastic to carbonate-rich sedimentary rock indicates the lower boundary of the

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Sekköy Member. Above the uppermost coal seam follows a series of alternating lightgrey clayey limestone and blueish-grey marlstone reaching a thickness of 24 m. Leaf

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fossils, mollusc shell debris, and concretions are present in this part but are rare or

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restricted to single beds. Above this unit, there follows a 20 cm thick layer of laminated clayey limestone, including a thin layer (0.5–1 cm) of dark grey to black

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iron-rich sandstone (S153611). Similar beds have been reported from the Eskihisar

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lignite mine (Bouchal et al., 2016a). The laminated clayey limestone is succeeded by 39 m of beige to light-grey limestones interbedded with clayey limestones. In the upper third of this unit are three humic lignite horizons (2–10 cm thick). Sedimentary rocks higher in the section have been removed by mining activity and were not available for study. This 75 m thick sequence of the Tınaz lignite mine section corresponds closely to Becker-Platen’s (1970, table 7) profile of the borehole core 56B/66, differing merely by the presence of thin humic lignite beds in the upper part of the Sekköy Member.

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ACCEPTED MANUSCRIPT 2.3. Age of the Tınaz lignite mine section No vertebrate fossils or datable volcanic layers have yet been reported from the Tınaz

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Subbasin, but lithostratigraphic and palynological data strongly suggest that deposition of the lacustrine sediments overlying the main coal seam at the Eskihisar lignite mine

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and at the Tınaz lignite mine occured during the same time interval (Becker-Platen, 1970; Atalay, 1980; Bouchal et al., 2016a; 2016b; this study). Sickenberg et al. (1975)

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reported the proboscid Gomphotherium angustidens Cuvier, 1817 [specimen PV12402, EUNHM, Appendix SI 2(1)] from the main lignite seam (upper part of the

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Turgut Member) of the Eskihisar lignite mine, ca 20 km northwest of the Tınaz lignite mine. It is noteworthy that the lower molar of the Eskihisar specimen shares several

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morphological and metric features with that of Kultak (MN5–6), described as G.

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angustidens (Kayseri-Özer et al., 2014a; Kaya et al, 2001) and differs from younger (late Miocene) forms in having smaller dimensions. The ongoing study of mammalian

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fossils collected from the Eskihisar main coal seam has revealed the presence of the

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middle Miocene carnivore Percrocuta miocenica Pavlov et Thenius, 1965 [specimen PV-12400, EUNHM, Appendix SI 2(1)]. This species is a peculiar element of middle Miocene faunas of the southeast Mediterranean region and is mostly known from the vast amount of material from the hominid locality Paşalar (early MN6); the youngest record of this taxon comes from the Prebreza fauna (late MN6, Serbia). The Eskihisar material is very similar to that of Paşalar and more evolutionarily advanced than that of Percrocuta sp. from the Antonios faunas (MN4–5, Greece) (Mayda et al., 2015; Koufos, 2008). In the light of these new faunal data, it seems probable that the Eskihisar fauna belongs to MN6, thus providing an age constraint for the main lignites 9

ACCEPTED MANUSCRIPT of the Eskihisar and Tınaz mines. Therefore, a late Langhian to early Serravallian age is suggested for the lower parts (lignite seam and overlying sediments of the Sekköy

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Member, pollen zones 1 and 2) of the Tınaz lignite mine section.

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2.4. Age of the Çatakbağyaka locality deposits, Sekköy Member

The fossil mammal assemblage found in the Sekköy Member of Çatakbağyaka (Muğla

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province, site Çatakbağyaka, ca 10 km south of the Tınaz lignite mine; Fortelius, 2016; Sickenberg et al., 1975) has been assigned to the Mammal Neogene (MN) zones

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(5–)6–8 [Supporting Information, Appendix SI 2(2)] by Fortelius (2016). Previous palynological studies (Jiménez-Moreno , 2005; Jiménez-Moreno et al., 2007; Biltekin,

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2010) also referred the Çatakbağyaka deposits to MN5 (late Burdigalian to Langhian).

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However, a thorough re-evaluation of the mammal fauna established an age for this assemblage of about 12.5–11.2 Ma (MN7/8, late Serravallian) [Supporting

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Information, Appendix SI 2 (3)]. These age determinations are important when

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comparing pollen zones of the Tınaz lignite mine section with adjacent areas.

2.5. Sample processing and data presentation Sediment samples were processed following the protocol described by Grímsson et al. (2008). The same pollen grains were investigated using LM and SEM (single grain method, Zetter, 1989). Light microscopy photographs of dispersed fossil pollen were taken with an Olympus BX51 microscope equipped with an Olympus DP71 camera. Specimens were sputter coated with gold and photographed using a Hitachi S-4300 10

ACCEPTED MANUSCRIPT cold field emission scanning electron microscope (NRM, Stockholm). For the quantitative analysis, 400 palynomorphs per sample were counted. The terminology

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for pollen morphology follows Punt et al. (2007) and Hesse et al. (2009). The pollen diagram (Fig. 3) was generated in C2 vers. 1.7.6; maps and sections were drawn in

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Adobe Illustrator 15.0.0 and photographs were cropped in Adobe Photoshop 12.0. Sediment samples, processed samples, and SEM stubs are stored at the Swedish

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Museum of Natural History, Stockholm, under accession numbers S153594 to S153621.

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To avoid redundancy with a previously published paper (Bouchal et al., 2016a) dealing with a coeval palynoflora from the neighboring Eskihisar Subbasin, full

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documentation (description, LM and SEM images) was only provided for those taxa

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that are new for the Yatağan Basin or difficult to discern based on LM (e.g. genera of Fagaceae, Oleaceae). Common types were documented using LM and for their

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detailed descriptions and taxonomic remarks reference was made to the work by

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Bouchal et al. (2016a).

3. Results/Systematic palynology In general, we employed the botanical nomenclature when describing palynomorphs. In some cases, when artificial names have been widely used to denote particular palynomorphs, the botanical name is followed by the relevant artificial (fossil-taxon) name.

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ACCEPTED MANUSCRIPT 3.1. Fungal spores

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Fungal spores were encountered but not included in the pollen count.

Pucciniaceae Chevallier

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Genus Puccinia Persoon Puccinia sp. (Plate II, 1–3)

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Description: Uredospore, shape spheroidal, outline circular, diameter 25–35 µm (LM, SEM); uredospore wall 1.5–2 µm thick (LM, SEM); no germ pores observed; psilate

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(LM), microechinate (SEM), microechinus base width 0.4–0.6 µm, microechinus length 0.5–1 µm (SEM), microechini surrounded by a torus, at a distance of 0.2–0.5

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µm (SEM).

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Ocurrence: Present in sample S153610. Remarks: This uredospore has ornamentation (microechini surrounded by a torus)

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diagnostic of Puccinia (Käärrik et al., 1983). Extant Puccinia spp. are pathogenic

3.2. Algae

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parasites mainly on angiosperms as principal hosts (Webster and Weber, 2007).

Botryococcaceae Wille Genus Botryococcus Kützing Botryococcus cf. B. brauni Kützing (Plate I, 1) Remarks: Colonies of Botryococcus are present in most samples of the Tınaz section (Fig. 3) and have been reported by others from Miocene palynofloras of western Turkey (e.g. Akgün et al., 2007; Bouchal et al., 2016a). 12

ACCEPTED MANUSCRIPT Zygnemataceae Kützing Genus Spirogyra Link/Ovoidites R. Potonié

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Remarks: Zygospores/aplanospores of Spirogyra are commonly assigned to the fossilgenus Ovoidites (R. Potonié) Krutzsch (e.g. Krutzsch and Pacltová, 1990; Van Geel

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and Grenfell, 1996; Worobiec, 2014). Ovoidites has been associated with shallow, stagnant, oxygen-rich, open fresh waters, lake margins, and marsh habitats (Rich et al.,

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1982; Van Geel and Van der Hammen, 1978). In the pollen diagram (Fig. 3) the four types were combined under the category Zygnemataceae. Highest diversity (four

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morphotypes) was encountered in sample S153601.

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Spirogyra sp. 1/Ovoidites lanceolatus Takahashi et Jux (Plate I, 2)

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Ocurrence: Most abundant zygospores/aplanospores encountered in the Tınaz section, present in samples S153594 to S153610 and S153621.

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Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 20, figs 3A–C, as

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Zygnemataceae gen. indet./Ovoidites lanceolatus).

Spirogyra sp. 2/Ovoidites microfoveolatus Krutzsch (Plate II, 4–6) Description: Zygospore or aplanospore, outline elliptic, length of axis perpendicular to fissure 90–110 µm (LM, SEM), length of axis parallel to fissure 100–130 µm (LM, SEM); mesospore 2–2.5 µm thick (SEM); psilate, perforate (LM, SEM). Ocurrence: Present in samples S153595, S153598 and S153601.

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ACCEPTED MANUSCRIPT Remarks: Similar perforate zygospores/aplanospores have been reported by Takahashi and Jux (1991, Schizosporis cf. parvus Cookson et Dettman, pl. 23, fig. 4) from the

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Spirogyra sp. 3/Ovoidites pococki Nakoman (Plate II, 7–9)

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Soma Basin.

Description: Zygospore or aplanospore, outline elliptic, length of axis perpendicular to

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fissure 50–65 µm (LM, SEM), length of axis parallel to fissure 95–115 µm (LM, SEM); mesospore 1.5–2 µm thick (SEM); weakly rugulate, rugulae forming a

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polygonal mesh-like pattern, scattered perforate (LM, SEM), rugulae form a reticulum-like pattern.

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Ocurrence: Present in sample S153601.

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Remarks: Similar rugulate zygospores/aplanospores have been reported from the Thrace Basin (Nakoman, 1966, pl. 11, figs 34–35) and the Soma Basin by Takahashi

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and Jux (1991, Ovoidites pseudoligneolus Krutzsch pl. 23, only fig. 8).

Spirogyra sp. 4/Ovoidites raatzi Nakoman (Plate II, 10–12) Description: Zygospore or aplanospore, outline nearly circular, length of axis perpendicular to fissure 70–85 µm (LM, SEM), length of axis parallel to fissure 70–80 µm (LM, SEM); mesospore 0.5–1 µm thick (SEM); sparsely rugulate (LM, SEM), rugula length 1–3 µm, rugula width 0.2–0.5 µm (SEM), predetermined breaking fissure indicated by slightly raised, ca 0.5–1 µm broad, band expanding from pole to pole.

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ACCEPTED MANUSCRIPT Ocurrence: Present in sample S153601. Remarks: Similar rugulate zygospores/aplanospores have been reported from the

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Thrace Basin (Nakoman, 1966, pl. 11, figs 24) and the Soma Basin by Takahashi and

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Jux (1991, pl. 23, fig. 5).

Algal cysts of uncertain affinity

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Ocurrence: Both types of algal cysts are abundant in samples S153594 to S153610 and

Algal cyst indet. 1 (Plate I, 3–4)

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were combined for the pollen diagram (Fig. 3).

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Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 20, figs 3G–I).

Algal cyst indet. 2/Sigmopollis pseudosetarius (Weyland et Pflug) Krutzsch et

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Pacltová (Plate I, 5)

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Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 20, figs 3G–I).

3.3. Ferns and mosses Sphagnaceae Dumortier Genus Sphagnum Linneus Sphagnum sp./Stereisporites stereoides Pflug (Plate III, 1–3) Description: Spore, amb circular to convex triangular, equatorial diameter 30–35 µm (LM, SEM); exospore 1–1.5 µm thick (LM); trilete, labrum, laesurae spanning entire radius (LM, SEM); psilate (LM, SEM), line of perforations parallel to laesura present. 15

ACCEPTED MANUSCRIPT Ocurrence: Present in samples S153594 to S153610.

Genus Osmunda Linnaeus

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Osmunda sp./Baculatisporites spp. (Plate I, 6–7)

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Osmundaceae Berchthold et J. Presl

Ocurrence: Spores of Osmundaceae are common in samples S153594 to S153609

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(Fig. 3), with a mass occurrence in sample S153598 (<50 %).

Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 20, figs 4A–F, as

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Osmunda spp.).

Pteris sp. (Pl. I, 8–9)

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Genus Pteris Linnaeus

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Pteridaceae E. D. M. Kirchner

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Occurrence: Single occurrences in samples S153595, S153598, and S153604,

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Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 2, figs 5D–G, as Pteris sp.).

Pteridaceae gen. indet./Cingulatisporites macrospeciosus (R. Potonié et Gelletich) Nakoman (Plate III, 4–6) Description: Spore, spheroidal, amb convex triangular, length of polar axis 28–32 µm (SEM), equatorial diameter 50–65 µm (LM, SEM); exospore (including sculpture elements) 2–2.5 µm thick (LM); trilete, laesurae 1/2 to 2/3 of radius, (equatorial flange) cingulum (LM, SEM); distal sculpturing rugulate, proximal sculpturing 16

ACCEPTED MANUSCRIPT rugulate and verrucate (LM, SEM), two main types of rugulae present, long rugulae, situated in equatorial area adjacent to cingulum, smaller irregularly shaped

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rugulae/verrucae in polar areas. Ocurrence: A single specimen was encountered in sample S153602.

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Remarks: Similar exospore ornamentation (long rugulae adjacent to cingulum) occurs in extant Pteridaceae, e.g. Anogramma Link, Cheilanthes Swarts (Tyron and

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Lugardon, 1991). Similar spores have been reported from Neogene palynofloras of western Turkey: Cingulatisporites macrospeciosus (R. Potonié et Gelletich) Nakoman

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(Nakoman, 1967, pl. 1. fig. 12, possibly fig. 3; Akgün and Akyol, 1999, fig. 11.7;

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Akyol and Akgün, 1990, pl. 1., fig. 5).

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Trilete and monolete spores of uncertain botanical affinitiy

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Fossil-genus Laevigatosporites Ibrahim

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Monolete spore fam. indet./Laevigatosporites haardti (R. Potonié et Venitz) P. W. Thomson et Pflug (Plate I, 10) Occurrence: Present in nearly all samples of the Tınaz section. Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 23, figs 6D–F, as Monolete spore fam. et gen. indet./Laevigatosporites haardti). Monolete spores of Aspleniaceae, Davalliaceae, Dryopteridaceae, Lomariopsidaceae, Oleandraceae, Polypodiaceae, Pteridaceae, Thelypteridaceae, and Vittariaceae lacking their perispor can be psilate (cf. Manchester and Zavada, 1987; Tyron and Lugardon, 1991, Stuchlik

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ACCEPTED MANUSCRIPT et al., 2001); these can only be unambigously differentiated by their exine architecture

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using TEM (cf. Tyron and Lugardon, 1991).

Fossil-genus Convolutispora Hoffmeister, Staplin et Malloy

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Trilete spore fam. indet. 1/cf. Convolutispora ampla Hoffmeister, Staplin et Malloy (Plate III, 7–9)

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Description: Spore, oblate to spheroidal, amb circular, diameter 60–70 µm (LM, SEM); exospore (including sculpture elements) 3–4 µm thick (LM); trilete (indistinct),

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laesurae 1/3 to 1/2 of radius (LM); rugulate, fossulate (LM, SEM). Ocurrence: A single specimen in sample S153621.

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Remarks: Trilete spore fam. indet. 1 has closest morphological similarity [indistinct

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trilete mark, size range, rugulate (=vermiculate) sculpturing] with Convolutispora ampla (Hoffmeister, Staplin and Malloy, 1955, p. 38, fig. 12); Krutzsch (1967, p. 132,

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pl. 46, 8–11) also compared similar spores to this fossil-specis. It is noteworthy that

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extant Osmunda regalis Linnaeus can produce spores with fused bacula forming rugulae (Stafford, 2003; Tyron and Lugardon, 1991, fig. 10.6) similar to the specimen encounered here. The corroded surface of this spore (Pl. III, 6) may indicate long distance dipersal (exposure to UV or transport) before burial or that the fossil spore had been reworked.

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ACCEPTED MANUSCRIPT Fossil-genus Stereisporites Pflug et P. W. Thomson Trilete spore fam. indet. 2/cf. Stereisporites (Stereigranisporis) granulus Krutzsch

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(Plate III, 10–12) Description: Spore, amb circular to weakly convex triangular, equatorial diameter 25–

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30 µm (LM, SEM); exospore (including sculpture elements) 1.5–2 µm thick (LM); trilete, laesurae 2/3 to entire radius (LM); rugulate, verrucate (LM), fossulae between

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sculptural elements perforate.

Occurrence: A single specimen in sample S153599.

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Remarks: This spore corresponds to Stereisporites (Stereigranisporis) granulus of Krutzsch (1963) based on its rugulate to verrucate sculpturing, but differs by the

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absence of a distinct curvatura imperfecta (exospore thickening in apical area of

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laesura). Similar exospore ornamentation is found in the extant Pteridaceae Cryptogramma R. Brown [e.g. C. crispa (Linnaeus) R. Brown] and Grammitidaceae

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Grammitis Swarts (Tyron and Lugardon, 1991; Vaganov et al., 2010).

3.4. Gymnosperms

Ephedraceae Dumortier Genus Ephedra Linnaeus The three Ephedra types were combined in the pollen diagram (Fig. 3)

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ACCEPTED MANUSCRIPT Ephedra sp. 1/Ephedripites (Ephedripites) regularis Hoeken-Klinkensberg (Plate IV, 1–3)

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Description: Pollen, shape oblate, equatorial outline elliptic, length of polar axis 18–30 µm (LM, SEM), equatorial diameter 45–60 µm (LM, SEM); exine ca 1 µm thick,

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tectate; inaperturate; plicate, psilate, fossulate (LM, SEM), 10–15 plicae present, pseudosulci running parallel to plicae (LM, SEM).

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Occurrence: Single occurrences in S153610 and S153617.

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Ephedra sp. 2/Ephedripites (Distachyapites) tertiarius Krutzsch (Plate IV, 4–6) Description: Pollen, shape oblate, equatorial outline elliptic, length of polar axis 20–35

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µm (LM, SEM), equatorial diameter 55–75 µm (LM, SEM); exine ca 1 µm thick,

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tectate; inaperturate; plicate, psilate, fossulate (LM, SEM), 8–12 plicae present,

SEM).

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pseudosulci running parallel to plicae, pseudosulci showing first order branching (LM,

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Occurrence: Present through the entire Tınaz section.

Ephedra sp. 3/Ephedripites (Distachyapites) lusaticus Krutzsch (Plate IV, 7–9) Description: Pollen, shape oblate, equatorial outline elliptic, length of polar axis 25–35 µm (LM, SEM), equatorial diameter 30–60 µm (LM, SEM); exine ca 1 µm thick, tectate; inaperturate; plicate, psilate, fossulate (LM, SEM), 6–9 plicae present, pseudosulci running parallel to plicae, pseudosulci show first and second order branching (LM, SEM).

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ACCEPTED MANUSCRIPT Occurrence: Present through the entire Tınaz section. Remarks: Extant and fossil Ephedra pollen have two main types of pseudosulci;

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unbranched (ancestral) and branched (derived) (Bolinder et al., 2016). Unbranched pseudosulci, as encountered in Ephedra sp. 1, are found in extant Mediterranean and

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South American species of Ephedra. Ephedra sp. 2 has branches of first order, and Ephedra sp. 3 shows branches of first and second order. Pollen of this type is found in

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the Asian clade and the North American clade of Ephedra. It is possible that Ephedra sp. 2 and 3 were produced by the same species and represent intraspecific

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morphological variability. Ephedra has been reported from middle Miocene

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palynofloras of western Turkey (e.g. Akgün et al., 2007; Bouchal et al., 2016a).

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Cupressaceae Richard ex Bartling

Cupressaceae gen. indet. 1 “non papillate”/Inaperturopollenites dubius (R. Potonié et

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Venitz) P. W. Thomson et Pflug (Plate I, 11)

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Occurrence: Present through the entire Tınaz section (Fig. 3). Remarks: Non papillate Cupressaceae pollen has been reported from Miocene palynofloras of western Turkey (e.g. Akgün et al. 2007). It is noteworthy that Cupressaceae macrofossils are rarely found in the Yatağan Basin (Gemici et al., 1990; Güner and Denk, 2014; Güner, 2016).

Cupressaceae gen. indet. 2 “papillate”/Inaperturopollenites concedipites (Wodehouse) Krutzsch (Plate I, 12)

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ACCEPTED MANUSCRIPT Occurrence: Present in samples S153594 to S153604. Remarks: Papillate pollen is found in a few subfamilies of extant Cupressaceae,

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namely Athrotaxioideae (Athrotaxis D. Don), Sequoioideae (Metasequoia Hu et W. C. Cheng, Sequoia Endlicher, Sequoiadendron J. Buchholz), Taiwanioideae (Taiwania

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Hayata), and Taxodioideae (Cryptomeria D. Don, Glyptostrobus Endlicher, Taxodium Richard), the remaining subfamilies produce non-papillate pollen (Van Campo-

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Duplan, 1951; Kvavadze, 1988).

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Cupressaceae gen. indet. 3 “papillate”/Sequoiapollenites gracilis Krutzsch (Plate I, 13) Occurrence: Single occurrences in S153601 and S153604.

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Remarks: In the pollen diagram (Fig. 3) ruptured and unruptured papillate pollen was

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combined.

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Pinaceae Sprengler ex F. Rudolphi

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Genus Cathaya Chun et Kuang Cathaya sp. 1/Cathayapollis scheuringii (Sivak) Ziembińska-Tworzydło (Plate VI, 10–13)

Description: Pollen, bisaccate, shape oblate, corpus circular to weakly rhombic in polar view, sacci spherical, attachment area of sacci narrow, diameter including sacci 60–75 µm (LM, SEM), corpus diameter 25–30 µm (LM, SEM), sacci width 40–50 µm (LM, SEM), sacci height 25–35 µm (LM); sacci with alveolate structuring; leptoma area psilate (LM), nanoechinate (SEM), cappa rugulate to verrucate, nanoechinate, perforate to fossulate (SEM), sacci nanoechinate, perforate (SEM). 22

ACCEPTED MANUSCRIPT Ocurrence: Single occurrences in S153594, S153602, and S153604. Remarks: Extant and fossil Cathaya pollen has nanoechinate exine sculpturing (e.g.

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Grímsson and Zetter, 2011; Liu and Basinger, 2000). Cathaya sp. 1 differs from C. sp. 2 by a smaller, circular corpus and narrow sacci attachment areas. The SEM analysis

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revealed a nanoechinate exine sculpturing for all specimens. Aberrant Cathaya pollen shows strong morphological similarities (small circular to rhomboidal corpus, large

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Bouchal et al. (2016a, p. 25, figs 7A–H).

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sacci) to pollen of Podocarpus in LM (compare Xi, 1986); for further remarks see

Cathaya sp. 2/Cathayapollis vancampoae (Sivak) Ziembińska (Plate VI, 14–15)

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Ocurrence: Present through the entire Tınaz section (Fig. 3).

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Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 25, figs 7E–H, as

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Cathaya sp.).

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Genus Cedrus Trew

Cedrus sp. (Plate I, 14) Ocurrence: Present through the entire Tınaz section (Fig. 3). Remarks: Extant and fossil Cedrus pollen is thickened in the cappa and saccus attachment area (Fujiki, 2003; Beug, 2004; Grímsson and Zetter, 2011). For descriptions and remarks see Bouchal et al. (2016a, p. 25, figs 7I–L).

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ACCEPTED MANUSCRIPT Genus Pinus Linnaeus Subgenus Pinus Linnaeus. (Diploxylon-pollen-type)

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Pinus subgenus Pinus sp. (Plate I, 15) Ocurrence: Present through the entire Tınaz section (Fig. 3).

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Remarks: Pinus subgenus Pinus (Diploxylon) pollen type is characterised by near spherical sacci with narrow attachment to the corpus (Hesse et al., 2009). For

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descriptions and remarks see Bouchal et al. (2016a, p. 27, figs 7Q–T).

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Subgenus Strobus (Lemmon) A. E. Murray (Haploxylon-pollen-type) Pinus subgenus Strobus sp. (Plate I, 16)

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Ocurrence: Present through the entire Tınaz section (Fig. 3).

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Remarks: Pinus subgenus Strobus (Haploxylon) pollen type is characterised by halfspherical sacci with broad attachment to the corpus (Hesse et al., 2009). For

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descriptions and remarks see Bouchal et al. Bouchal et al. (2016a, p. 27, figs 8A–D).

Genus Tsuga Carrière Tsuga sp./Zonalapollenites verrucatus Krutzsch ex Ziembińska-Tworzydło (Plate V, 1–4) Description: Pollen, monosaccate, shape oblate, circular in polar view, equatorial diameter 40–55 µm (LM), 35–45 µm (SEM); sculpturing rugulate (LM), leptoma psilate to weak rugulate, monosaccus rugulate, fossulate, proximal face rugulate, fossulate (SEM).

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ACCEPTED MANUSCRIPT Occurrence: Single occurrences in samples S153602 (Fig. 3) and S153621. Remarks: Tsuga pollen without echini is found only in extant T. canadensis (Linnaeus)

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Carrière and T. caroliniana Engelmann (Sivak, 1973). Tsuga pollen has been reported previously from middle Miocene palynofloras of western Anatolia: Tsugaepollenites

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igniculus (R. Potonié) R. Potonié et Venitz (Benda, 1971), Zonalapollenites maximus (Raatz) Krutzsch (Takahashi and Jux, 1991), Z. verrucatus (Kayseri-Özer et al.,

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2014a), Tsuga (Yavuz-Işık et al., 2011).

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3.5. Angiosperms 3.5.1. Monocots

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Commelinids

Poaceae Barnhart

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Poales Small

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Remarks: The taxonomic significance of pollen morphology is limited within Poaceae

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(cf. Page, 1978). In the pollen diagram (Fig. 3) Poaceae gen. indet. 1, 2 and 3 were combined.

Poaceae gen. indet. 1 (Plate I, 17–18) Occurrence: Present through the entire Tınaz section. Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 29, figs 8H–J, as Poaceae gen. indet. 1).

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ACCEPTED MANUSCRIPT Poaceae gen. indet. 2 (Plate I, 19–20) Occurrence: Present through the entire Tınaz section.

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Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 29, figs 8K–M).

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Poaceae gen. indet. 3 (Plate V, 5–7)

Description: Pollen, shape spheroidal, outline circular in equatorial view, diameter 40–

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50 µm (LM, SEM); eutectate, exine 1–1.5 µm thick (LM); ulcerate, ulcus diameter 2.5–3 µm (LM, SEM), annulus present (LM, SEM); sculpturing scabrate (LM),

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nanoechinate, nanoechini widely spaced, weakly rugulate (SEM). Occurrence: Present through the entire Tınaz section.

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Remarks: Poaceae gen. indet. 3 differs from Poaceae gen. indet. 1 and 2 by nanoechini

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that are not clustered. Nonareolate pollen grains with spaced nanoechini are found in several of extant Poaceae genera, e.g. Stipa Linnaeus, Diandrolyra Stapf, Glyceria

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Nuttall, Secale Linnaeus (Köhler and Elsbeth, 1979; Page, 1978).

Typhaceae Jussieu

In the pollen diagram (Fig. 3) Sparganium sp. and Typha sp. were combined in the category Typhaceae.

Genus Sparganium Linnaeus Sparganium sp. (Plate VIII, 10–12) Description: Pollen, shape spheroidal, outline circular, diameter 30–40 µm (LM, SEM); semitectate, exine 1.5–2.5 µm thick (LM), nexine thinner than sexine; ulcerate, 26

ACCEPTED MANUSCRIPT porus diameter 3–3.5 µm (LM, SEM); sculpturing reticulate (LM, SEM), muri crested with blunt nanoechini, muri width 0.5–1.5 µm (SEM), lumina of irregular shape.

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Occurrence: Present through the entire Tınaz section, but less regularly represented than Typha.

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Remarks: Dispersed Sparganium and Typha pollen monads are difficult to unambigiously determine under LM investigation (cf. Beug, 2004; Punt, 1975), under

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SEM investigation fossil and extant Sparganium pollen has muri crested with blunt excrescences whereas Typha has smooth to sharp edged crested muri (Grímsson et al.,

Genus Typha Linnaeus

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Typha sp. (Plate IX, 1–6)

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2015; Punt, 1975).

Description: Pollen, monad or planar tetrad, shape spheroidal, single pollen outline

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circular, tetrad diameter 40–50 µm (LM), 35–45 (SEM), single pollen diameter 22–30

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µm (LM), 20–26 µm (SEM); semitectate, exine 1.5–2 µm thick (LM), nexine thinner than sexine; ulcerate, porus diameter 4–5 µm (LM, SEM); sculpturing reticulate (LM, SEM), muri crested with ridges or smooth, muri commonly incomplete, lumina of irregular shape. Occurrence: Present through the entire Tınaz section, a single tetrad was encountered in sample S153621. Remarks: For remarks see Bouchal et al. (2016a, p. 29, figs 9A–C).

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ACCEPTED MANUSCRIPT Family Zingiberaceae Martinov Genus Spirematospermum Chandler

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Spirematospermum sp. (Plate VI, 1–3) Description: Pollen, shape spheroidal, outline circular, diameter (including echini) 19–

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23 µm (LM), 18–20 µm (SEM); possibly atectate, exine ca 1 µm thick (LM), sexine thinner than nexine; inaperturate (LM, SEM); sculpturing echinate (LM, SEM), echini

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basal diameters 0.4–1.4 µm (SEM), echini length 0.9–1.9 µm (SEM), echini constricted at base.

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Occurrence: Encountered in sample S153621.

Remarks: Pollen of this type has been associated with the extinct Zingiberaceae

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Spirematospermum (Heer) Chandler (Fischer et al., 2009; Kmenta and Zetter, 2013).

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Reports of morphologically similar pollen from middle Miocene palynofloras of western Turkey include Smilacipites Wodehouse (Takahashi and Jux, 1991, only pl.6,

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figs 2a–b) and possibly Echinigraminidites moravicus Krutzsch (Akgün and Akyol,

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1999).

3.5.2. Eudicots

Ranunculales Jussieu ex Berchthold et J. Presl Ranunculaceae Jussieu Ranunculaceae gen. indet. (Plate VI, 4–6) Description: Pollen, shape prolate, outline elliptic in equatorial view, length of polar axis pollen 38–42 µm (LM), (SEM), equatorial diameter 27–30 µm (LM), 29–40 µm (SEM); tectate, exine 2–2.5 µm thick (LM), nexine thinner than sexine; tricolpate, 28

ACCEPTED MANUSCRIPT colpus length 2/3 to 3/4 of polar axis; sculpturing microechinate (LM), microechinate, nanoechinate, perforate (SEM), microechini basal diameters 0.5–0.8 µm (SEM),

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microechini length 0.5–0.8 µm (SEM). Occcurrence: Single occurrences in samples S153605 to S153610 (Fig. 3).

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Remarks: Owing to its morphological similarities to the Ranunculus arcis type of Clarke et al. (1991) this specimen was assigned to Ranunculaceae. Previous reports of

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Ranunculaceae from middle Miocene palynofloras of western Turkey: Ranunculacidites sp. (Takahashi and Jux, 1991, pl. 15, figs 11a–b), Ranunculaceae

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(Yavuz-Işık et al., 2011), “Thalictrum-Typ” of Benda (1971)

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Core Eudicots

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Saxifragales Berchthold et J. Presl Altingiaceae Horaninow

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Genus Liquidambar Linnaeus

VI, 7–9)

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Liquidambar sp./Peripollenites stigmosous (R. Potonié) Pflug et P. W. Thomson (Plate

Description: Pollen, shape spheroidal, outline circular, diameter 31–43 µm (LM), 29– 40 µm (SEM); tectate, exine 1.5–2 µm thick (LM), nexine thinner than sexine; panto(15–20)porate (LM, SEM), pores circular to elliptic, pore diameter 2–8 µm (LM, SEM), porus membrane verrucate, rugulate, nanoechinate; sculpturing scabrate (LM), perforate, nanoechinate (SEM).

29

ACCEPTED MANUSCRIPT Occurrence: Present in sample S153595 (Fig. 3). Remarks: All extant Altingiaceae (Altingia Noronha, Liquidambar, Semiliquidambar

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H. T. Chang) produce pantoporate, perforate nanoechinate pollen grains with verrucate, rugulate, nanoechinate aperture membranes; in Liquidambar, pore shape

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varies from circular to elliptic; in Altingia, pores are circular (compare Bogle and Philbrick, 1980; Jones et al., 1995; Li et al., 2010; Miyoshi et al., 2011; Ickert-Bond

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and Wen, 2013). Recent phylogenetic studies of Ickert-Bond and Wen (2006, 2013) showed that most of extant Altingiaceae (except Altingia chinensis Oliver ex Hance,

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A. poilanei Tardieu, A. yunnanensis Rehder et E. H. Wilson) are nested within Liquidambar.

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Previous reports of Altingiaceae from middle Miocene palynofloras of western

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Turkey: Periporopollenites stigmosus (Gemici et al., 1991, pl. 6, fig. 9; Akgün and Akyol, 1999, figs 10.99, 11.94, 12.56; Akgün et al., 2007, Appendix A), Liquidambar

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(Yavuz-Işık, 2007; Yavuz-Işık et al., 2011; Akkiraz et al., 2012, pl. 4, fig. 21; Kayseri-

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Özer et al., 2014b; Bozcu et al., 2015), possibly “Caryophyllaceen-Typ” of Benda (1971, pl. 4, fig. 34). Fossil leaves of Liquidambar have been reported from the Miocene of Çan, Çanakkale (Mädler and Steffens, 1979) and the Yatağan Basin (Güner and Denk, 2014; Bouchal et al., 2015).

30

ACCEPTED MANUSCRIPT Fabids Malpighiales Jussieu ex J. Presl

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Euphorbiaceae Jussieu Euphorbiaceae gen. indet. (Plate I, 21–23)

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Occurrence: Present in samples S153594 and S153598 to S153603 (Fig. 3). Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 31, figs 9K–M, as

Genus Linum Linnaeus

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Linum sp. (Plate I, 24)

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Linaceae de Candolle ex Perleb

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Euphorbiaceae gen. indet.).

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Occurrence: A single specimen in sample S153621.

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Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 31, figs 10A–C).

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Salicaceae Mirbel

Genus Salix Linnaeus Salix sp. (Plate IV, 10–12) Description: Pollen, shape prolate, outline elliptic in equatorial view, length of polar axis 20–25 µm (LM, SEM), equatorial diameter 9–18 µm (LM, SEM); semitectate, exine 1.5–2 µm thick (LM), nexine thinner than sexine; tricolporate (LM); sculpturing reticulate (LM, SEM), lumina with free standing columellae, lumina irregular shaped and decreasing in size in colpus area, muri wedge-shaped (SEM). Occurrence: Present in sample S153599 (Fig. 3). 31

ACCEPTED MANUSCRIPT Remarks: Sohma (1993) investigated pollen morphology in extant Salix and identified eight reticulation types. Wedge-shaped muri (Type 2 of Sohma, 1993) are the most

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common form and are produced by several extant subgenera. Previous reports of Salix from middle Miocene palynofloras of western Turkey: Salix

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(Yavuz-Işık, 2007; Kayseri and Akgün, 2010, pl. 5, figs 60–63; Akkiraz et al., 2012, pl. 3, figs 23–24; Kayseri-Özer et al., 2014a; 2014b;), possibly Tricolpopollenites

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retiformis (P. W. Thomson) P. W. Thomson et Pflug (Akgün and Akyol, 1999, only figs 9.54, 10.72–73, 11.53).

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Fossil leaves of Salix have been reported from the Miocene of western Anatolia: Balya, Balıkesir (Engelhardt, 1903), Soma Basin (Gregor, 1990; Gemici et al., 1991),

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Şahınalı, Aydın (Gemici et al., 1993), and the Yatağan Basin (Gemici et al., 1990;

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Fabales Bromhead

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Güner and Denk, 2014; Güner, 2016).

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Fabaceae Lindley

Genus Podocarpium (A. Braun) Herendeen Podocarpium podocarpum (A. Braun) Herendeen/Tricolporopollenites wackersdorfensis Thiele-Pfeiffer (Plate IV, 13–15) Description: Pollen, outline circular to irregular, length of polar axis 38–42 µm (LM, SEM), equatorial diameter 30–35 µm (LM, SEM); eutectate, exine 1.5–2 µm thick (LM); tricolporate, colpus length 5/6 of polar axis; scabrate (LM), rugulate, perforatefossulate (SEM), rugulae segmented, microrugulae width 0.3–0.5 µm (SEM). Ocurrence: Single occurrences in samples S153594, S153595, and S153610 (Fig. 3). 32

ACCEPTED MANUSCRIPT Remarks: This pollen type corresponds in aperture and exine sculpturing to Tricolporopollenites wackersdorfensis and nearly identical (notably larger) pollen has

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been reported previously from the Yatağan Basin (Bouchal et al., 2016a, p. 33, figs 10D–F, as Podocarpium sp./Tricolporopollenites wackersdorfensis). This pollen type

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has been found in situ in fossil flowers of the extinct Fabaceae Podocarpium podocarpum (Liu et al., 2001); the authors did not give a size range for the in situ

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pollen. In Thiele-Pfeiffer’s (1980) original description of this fossil-species she mentioned that this pollen type is: “…very large, tricolporate pollen of spherical

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shape. Diameter range is mostly around 40 µm. Colpi very slim, …”; Liu (1985) differentiated two fossil-species Tricolporopollenites wackersdorfensis

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(=Fupingopollenites wackersdorfensis in Shatilova, 2009), larger than 40 µm, colpus

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with broad segmented margo, and Fupingopollenites minutus Liu (Liu, 1985), smaller than 39 µm, colpus with slim segmented margo; both have been reported from Eocene

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to Pliocene sediments of the Aegean Sea, the Black Sea and its surrounding region

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(Shatilova, 2009).

Fagales Engler

Betulaceae Gray Genus Alnus Miller Alnus sp./Alnipollenites verus (R. Potonié) R. Potonié (Plate I, 25) Occurrence: Present in samples S153594 to S153608 (Fig. 3), with a mass occurrence in S153599 (19.25 %).

33

ACCEPTED MANUSCRIPT Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 33, figs 10J–L, as

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Alnus sp.).

Genus Betula Linnaeus

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Betula sp./Trivestibulopollenites betuloides Pflug (Plate I, 26) Ocurrence: Present through the entire Tınaz section (Fig. 3).

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Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 34, figs 11A–C,

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as Betula sp.).

Genus Carpinus Linnaeus

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Carpinus sp./Carpinipites carpinoides (Pflug) Nagy (Plate I, 27)

PT

Occurrence: Present in samples S153621, and S153594 to S153606 (Fig. 3). Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 34, figs 11D–F,

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as Carpinus sp.).

Genus Corylus Linnaeus Corylus sp. (Plate I, 28) Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 34, figs 11G–I, as Corylus sp.). When investigated with LM, Corylus, Ostrya and Myricaceae pollen have strong morphological similarities (cf. Edwards, 1981). Because of this, and the suboptimal preservation in most samples of the Tınaz section, these three taxa have been combined within the category Corylus/Ostrya/Myrica (Fig. 3). This combined pollen 34

ACCEPTED MANUSCRIPT type is common in most samples of the Tınaz section. Of the three taxa, only Myrica has been reported from the macrofossil record of the Yatağan Basin (Güner and Denk,

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2014; Bouchal et al., 2015; Güner, 2016).

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Genus Ostrya Scopoli Ostrya sp. (Plate I, 29)

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Occurrence: The Corylus/Ostrya/Myrica pollen type is common in most samples of the Tınaz section.

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Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 34, figs 11J–L).

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Ostrya has been combined with Corylus and Myricaceae (see Corylus sp.).

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Fagaceae Dumortier Genus Fagus Linnaeus

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Fagus sp. (Plate VIII, 1–2)

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Occurrence: Present through the entire Tınaz section (Fig. 3). Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 36, figs 12A–F).

Genus Quercus Linnaeus In the Tınaz section, three types of oak pollen can be distinguished using SEM. Using LM, reliable assignment of dispersed oak pollen to a specific infrageneric group is impossible because the defining infrageneric characters (exine sculpturing) are only detectable in SEM (cf. Denk and Grimm, 2009). Owing to this factor and poor preservation of the samples, only pollen size was used to designate Quercus pollen of 35

ACCEPTED MANUSCRIPT the Tınaz section in the pollen diagram. “Quercus Large” approximately matching Quercus Group Quercus and Group Cerris, and “Quercus Small” approximately

PT

matching Quercus Group Ilex (see Denk and Grimm, 2010 for infrageneric groups of Quercus). Both “Quercus Large” and “Quercus Small” are commonly found through

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the entire Tınaz section (Fig. 3).

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Quercus sp. 1 (Quercus Group Cerris) (Plate VII, 1–3)

MA

Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 36, figs 12G–I).

Quercus sp. 2 (Quercus Group Ilex) (Plate VII, 4–6)

PT

ED

Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 38, figs 12J–L).

Quercus sp. 3 (Quercus Group Quercus) (Plate VII, 7–9)

CE

Remarks For descriptions and remarks see Bouchal et al. (2016a, p. 38, figs 13A–C).

Cerris.

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Pollen of Quercus Group Quercus is of much lower abundance than Quercus Group

Genus Trigonobalanopsis Kvaček et Walther Trigonobalanopsis sp. (Plate VII, 10–12) Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 38, figs 13D–F).

36

ACCEPTED MANUSCRIPT Juglandaceae de Candolle ex Perleb Engelhardioideae Iljinskaja

PT

In the pollen diagram (Fig. 3), both Engelhardioideae pollen types were combined. Engelhardioideae pollen is only present in low abundance in the lower part (Pollen

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zone 1 and 2) of the Tınaz section (S153594 to S153604).

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Engelhardioideae gen. indet. 1/Momipites quietus (R. Potonié) Nichols (Plate VIII, 3) Description: Pollen, shape oblate, outline triangular in polar view, equatorial diameter

MA

17–21 µm (LM), 15–19 µm (SEM); eutectate, exine 1.0–1.5 µm thick (LM), nexine thinner than sexine; triporate, pore diameter 1–2 µm (LM), ectoporus circular to

ED

elliptic, aperture sunken; psilate (LM), nanoechinate (SEM).

PT

Remarks: Engelhardioideae gen. indet. 1 has strong morphological similarities in size, apertures, outline, and exine sculpturing to extant Engelhardia Leschenault ex Blume

CE

pollen (Stone and Broome, 1975).

AC

Previous reports of morphologically similar pollen from middle Miocene palynofloras of western Anatolia: Engelhardtioipollenites microcoryphaeus (R. Potonié) R. Potonié, P. W. Thomson et Thiergart ex R. Potonié (=Triatripollenites coryphaeus microcoryphaeus) (Benda, 1971, pl. 2, fig. 19–21; Gemici et al., 1991, pl. 7, fig. 12; Takahashi and Jux, 1991, pl. 20, figs 8–9), Triatripollenites coryphaeus (R. Potonié) P. W. Thomson et Pflug (Gemici et al., 1993, pl. 3, figs 14–15; Akgün and Akyol, 1999, only figs 9.32–33, 11.30–31), Momipites quietus (Akgün et al., 2007, appendix A; Kayseri-Özer et al., 2014a), Engelhardtia (Kayseri and Akgün, 2010, only figs 5.22–39; Kayseri-Özer et al., 2014b, fig. 6.15). 37

ACCEPTED MANUSCRIPT Engelhardioideae gen. indet. 2/Momipites punctatus (R. Potonié) Nagy (Plate VIII, 4) Remarks: For descriptions and remarks see Bouchal et al. (2016a p. 38, figs 13G–L, as

PT

Engelhardioideae gen. indet.).

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Juglandoideae Genus Carya Nuttall

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Carya sp./Caryapollenites simplex (R. Potonié) Raatz (Plate VIII, 5) Occurrence: Present through the entire Tınaz section (Fig. 3).

MA

Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 40, figs 14A–C,

ED

as Carya sp.).

PT

Myricaceae A. Richard ex Kunth

Genus Morella Loureiro vel Myrica Linnaeus

CE

Morella vel Myrica sp. (Plate VIII, 6–7)

AC

Occurrence: The Corylus/Ostrya/Myrica pollen type is common in most samples of the Tınaz section.

Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 42, figs 14J–L).

Rosales Berchthold et J. Presl Cannabaceae Marinov Genus Celtis Linnaeus vel Pteroceltis sp. Maximowicz Celtis vel Pteroceltis (Plate VII, 13–15)

38

ACCEPTED MANUSCRIPT Description: Pollen, outline circular in polar view, diameter 29–34 µm (LM), 26–31 µm (SEM); tectate, exine 1–1.5 µm thick (LM), nexine thinner than sexine;

PT

stephano(3–4)porate (LM, SEM), pores circular, annulus sporadically present, pore diameter 2–3 µm (LM, SEM); scabrate (LM), perforate, nanoechinate (SEM).

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Occurrence: Present in samples S153602, S153603 (Fig. 3), and S153621. Remarks: Extant Celtis and monotypic Pteroceltis produce pollen with similar size

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range, shape, outline, exine sculpturing, and pore number (compare Zavada, 1983; Takahashi, 1989; Stafford, 1995; Li et al., 2010); the recovered specimens correspond

MA

in their morphological characters to both. Fossil dispersed pollen with affinities to extant Celtis have commonly been assigned to the form genus Celtipollenites (Nagy)

ED

Kohlman-Adamska et Ziembińska-Tworzydło (e.g. Stuchlik et al., 2014).

PT

Previous reports of Celtis from middle Miocene palynofloras of western Turkey: Celtis

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(Yavuz-Işık et al., 2011; Akkiraz et al., 2012, pl. 3, fig. 2).

AC

Ulmaceae Mirbel

Genus Cedrelospermum (Saporta) Manchester Cedrelospermum sp. (Plate VIII, 8) Occurrence: Present in samples S153601 to S153604, S153595, and S153610 (Fig. 3). Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 42, figs 15D–I).

39

ACCEPTED MANUSCRIPT Genus Ulmus Linnaeus Ulmus sp. (Plate VIII, 9)

PT

Occurrence: Present through the entire Tınaz section (Fig. 3).

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Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 42, figs 15J–L).

Genus Zelkova Spach

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Zelkova sp. (Plate VIII, 10)

Occurrence: Present through nearly the entire Tınaz section (Fig. 3).

MA

Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 44, figs 16A–C).

ED

Malvids

Geraniaceae Jussieu

PT

Geraniales Jussieu ex. Berchthold et Presl

CE

Genus Erodium L’Héritier ex Aiton

AC

Erodium sp. (Plate VIII, 11) Occurrence: Present in samples S153594, S153603, S153604, S153610, and S153617 (Fig. 3).

Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 44, figs 16D–F).

Myrtales Jussieu ex. Berchthold et Presl Lythraceae J. Saint-Hilaire Genus Decodon J. F. Gmelin Decodon sp. (Plate IX, 1–6) 40

ACCEPTED MANUSCRIPT Occurrence: Present through the entire Tınaz section (Fig. 3), pollen agglomerations were encountered in sample S153595.

PT

Remarks: Decodon sp. resembles fossil pollen morphotype 6 of Grímsson et al. (2012); for further descriptions and remarks see Bouchal et al. (2016a, p. 44, figs

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16G–I). The pollen aggregation (Pl. IX, 4–6) might represent the content of an

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immature anthere.

Malvales.

MA

Malvaceae Jussieu Malvoideae Burnott

ED

Malvoideae gen. indet. (Plate IX, 7–9)

PT

Description: Pollen, shape spheroidal, outline circular in equatorial view, equatorial diameter excluding echini 85–103 µm (LM), 66–77 µm (SEM); eutectate, exine 2.5–4

CE

µm thick (LM), sexine thinner than nexine (SEM); pantoporate, porus diameter ca 1

AC

µm (SEM); echinate (LM), echinate, perforate (SEM), echini basal diameters 1.6–4.8 µm (SEM), echini short (3–4.5 µm) and long (9–12 µm) (SEM), with perforations at their base.

Occurrence: Single occurrence in sample S153617. Remarks: Malvoideae gen. indet. is heavily corroded but has characteristic Malvoideae (pantoporate, echinate) pollen morphology (Christensen, 1986; Culhane and Blackmore, 1988). Malvoideae pollen has been reported previously from the Eskihisar lignite mine (Bouchal et al., 2016a, p. 46, figs 17A–C), but the Tınaz section specimen differs by its larger size, longer echini and different aperture shape. 41

ACCEPTED MANUSCRIPT Tilioideae Arnott Genus Tilia Linnaeus

PT

Tilia sp./Intratriporopollenites instructus (R. Potonié) P. W. Thomson et Pflug (Plate VIII, 12)

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Occurrence: Single occurrences in samples S153594, S153595, S153621, and S153601 to S153606.

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Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 46, figs 17D–F,

MA

as Tilia sp.).

Sapindales Jussieu ex. Berchthold et Presl

ED

Sapindales fam. et gen. indet.

PT

Sapindales fam. et gen. indet. (Plate IX, 10–12) Description: Pollen, shape prolate, elliptic in equatorial view, length of polar axis 28–

CE

34 µm (LM), 26–30 µm (SEM), equatorial diameter 20–25 µm (LM), 18–23 µm

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(SEM); semitectate, exine 2.5–3 µm thick, sexine thicker than nexine, costae present, costae reaching polar colpus endings (LM); tricolporate, ectocolpus 2/3 to 3/4 of polar axis (LM, SEM), endoporus framed by quadrangular costa (LM); striate to striatoreticulate (LM, SEM). Occurrence: Present in samples S153594 to S153610 (Fig. 3). Remarks: Prolate pollen with a distinct costa is commonly found in Anacardiaceae, Simaroubaceae and Rutaceae (e.g. Erdtman, 1952; Barento et al., 1987; Beug, 2004). Sapindales pollen of uncertain affinity has been reported from the Eskihisar lignite

42

ACCEPTED MANUSCRIPT mine by Bouchal et al. (2016a, ARS-type 1 and 2, p. 48, figs 17G–L, affinity of ARS-

PT

type 3 is to Nitraria, see below).

Nitrariaceae Lindley

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Genus Nitraria Linnaeus Nitraria sp. (Plate IX, 13–15)

NU

Description: Pollen, shape prolate, elliptic in equatorial view, length of polar axis 32– 38 µm (LM), 30–32 µm (SEM), equatorial diameter 28–33 µm (LM), 24–32 µm

MA

(SEM); tectate, exine 2.5–3 µm thick, sexine thicker than nexine, costae present, costae not reaching polar colpus endings (LM); tricolporate, ectocolpus length 3/4 to

ED

5/6 of polar axis (LM, SEM), endoporus circular, framed by circular costa (LM);

PT

striate (LM), perforate, striate, rugulate, fossulate (SEM), in colpus area striae fused to rugulae and fossulae, peforate in mesocolpium (SEM).

CE

Occurrence: Present in samples S153608 and S153610.

AC

Remarks: Long ectocolpi, circular to rhombic costae, and striatoreticulate to rugulate perforate exine sculpturing occur commonly in extant Nitrariaceae (Abdusalih and Xiaoling, 2003; Li et al., 2010; Hoorn et al., 2012). Pollen with affinity to Nitraria has commonly been assigned to Nitrariadites Zhuet Xi Ping in Zhu Zunghao et al. and Nitrariapollis Xi Yizhen et Sun Mengrong (Hoorn et al., 2012). Previous reports of pollen with morphological similarities to Nitrariaceae from Miocene palynofloras of southwestern Anatolia: Cupuliferoipollenites villensis P. W. Thomson (Benda 1971, pl. 3, figure 40), ASR-Type-pollen gen. indet. 3 (Bouchal et al. 2016a, figs 18A–C),

43

ACCEPTED MANUSCRIPT Tricolporopollenites turcianuns Takahashi et Jux (Takahashi and Jux, 1991, pl. 16,

PT

figs 26–29).

Sapindaceae Jussieu

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Genus Acer Linnaeus

Acer sp. 1 and 2 were combined in the pollen diagram (Fig. 3) under the category Acer

MA

Acer morphotype 1 (PlateX, 1–6)

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and are of common abundance in the Tınaz section.

Description: Pollen, shape prolate, elliptic in equatorial view, length of polar axis 34–

ED

41 µm (LM), 31–39 µm (SEM), equatorial diameter 26–31 µm (LM), 21–26 µm

PT

(SEM); semitectate, exine 2.5–3 µm thick, sexine thicker than nexine; tricolpate, ectocolpus length 3/4 to 5/6 of polar axis (LM, SEM); scabrate (LM), rugulate,

CE

fossulate, perforate (SEM).

AC

Occurrence: Present in most samples of the Tınaz section (Fig. 3), but less common than Acer morphotype 3 (striate). Remarks: Biesboer (1975) identified four major pollen types in extant Acer; striate (Type I), microreticulate (Type III), rugulate (Type II) and with granular sculpturing (Type IV). Acer morphotype 1 corresponds to Biesboer’s (1975) Type II. Pollen of this type is produced by extant Acer negundo Linnaeus, A. saccharinum Linnaeus and A. saccharum Marshall (Biesboer, 1975; Tian et al., 2001). Morphologically similar pollen has been reported from several Miocene European palynofloras (e.g. Fürstl,

44

ACCEPTED MANUSCRIPT 2002, Hausruck, Austria, pl. 13, figs 4–12; Kmenta, 2011, Altmittweida, Germany, pl.

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3, figs 1–6).

Acer morphotype 2 (PlateX, 7–9)

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Description: Pollen, shape prolate, elliptic in equatorial view, length of polar axis 42– 46 µm (LM), 35–38 µm (SEM), equatorial diameter 31–35 µm (LM), 22–24 µm

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(SEM); tectate, exine 3–3.5 µm thick, sexine thicker than nexine; tricolpate, ectocolpus length 3/4 to 5/6 of polar axis (LM, SEM); scabrate (LM), rugulate to

MA

striatoretiulate, fossulate, perforate (SEM). Occurrence: Present in sample S153617.

ED

Remarks: Acer morphotype 2 is intermediate between Acer morphotype 1 and 3,

PT

showing rugulate to striatoreticulate exine sculpturing.

CE

Acer morphotype 3 (Plate X, 10–15)

AC

Description: Pollen, shape prolate to spheroidal, circular to elliptic in equatorial view, lobate in polar view, length of polar axis 34–46 µm (LM), 31–38 µm (SEM), equatorial diameter 24–35 µm (LM), 21–28 µm (SEM); semitectate, exine 2–3 µm thick, sexine thicker than nexine; tricolpate, ectocolpus length 3/4 to 5/6 of polar axis (LM, SEM); striate (LM, SEM). Occurrence: Present through the entire Tınaz section (Fig. 3). Remarks: Acer morphotype 3 differs in its striate exine sculpturing from Acer morphotype 1. Striate exine sculpturing is the most common ornamentation in extant and fossil Acer pollen (Biesboer, 1975, Type I; Clarke and Jones, 1978; Tian et al., 45

ACCEPTED MANUSCRIPT 2001; Fürstl, 2002; Li et al., 2010; Stuchlik et al., 2014). Striate Acer pollen has been reported previously from middle Miocene palynofloras of western Turkey (e.g. Akgün

PT

et al., 2007; Bouchal et al., 2016a, Acer sp., p. 48, figs 18D–F). Previous studies demonstrated that pollen of Acer is only of limited taxonomic value for assignment to

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infrageneric clades, because pollen morphology is highly variable within these groups (Biesboer, 1975; Tian et al., 2001; Fürstl, 2002). It is noteworthy that seven Acer leaf

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species and three types of samaras have been reported from the Yatağan Basin (Güner,

MA

2016).

Genus Picrasma Blume

ED

Simaroubaceae de Candolle

PT

Picrasma sp. (Plate XI, 1–4)

Description: Pollen, shape prolate, outline elliptic in equatorial view, length of polar

CE

axis 38–42 µm (LM, SEM), equatorial diameter 27–31 µm (LM, SEM); semitectate,

AC

exine 3–3.5 µm thick (LM), nexine thinner than sexine, exine weakly thickened in endoporus area; tricolporate (LM); striate, reticulate, heterobrochate (LM, SEM), lumina decreasing in size towards colpus, sculpturing striate adjacent to colpus, lumina psilate weakly perforate. Occurrence: A single specimen in sample S153595. Remarks: The available specimen corresponds in size, aperture, and exine sculpturing (reticulate, striate in colpus area) to pollen of extant Picrasma (Institute of Botany and South China Institute of Botany, 1982; Li et al., 2010; Miyoshi et al., 2011). The botanical affinities of the enigmatic flower Chaneya (reported from Burdigalian 46

ACCEPTED MANUSCRIPT deposits of central Turkey, Beşkonak/Güvem in Paicheler and Blanc, 1981) have

PT

recently been shown to be with Picrasma (Wang and Manchester, 2000).

Amaranthaceae Jussieu

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Caryophyllales Jussieu ex Berchthold et Presl

Two types of Amaranthaceae pollen were combined in the pollen diagram (Fig. 3).

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Amaranthaceae gen. indet. 1 (Plate VIII, 13)

Occurrence: Amaranthaceae pollen is present through the entire Tınaz section; highest

MA

abundance (51 %) was encountered in sample S153608. Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 50, figs 18G–I, as

PT

ED

Amaranthaceae vel Chenopodiaceae gen indet. 1).

Amaranthaceae gen. indet. 2 (Plate VIII, 14)

CE

Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 50, figs 18G–I, as

AC

Amaranthaceae vel Chenopodiaceae gen indet. 2).

Caryophyllaceae Jussieu Caryophyllaceae gen. indet. (Plate VIII, 15–16) Description: Pollen, shape spheroidal, outline polygonal to weakly circular, diameter 27–32 µm (LM), 25–29 µm (SEM); eutectate, exine 3–4 µm thick (LM), nexine thinner than sexine, pantoporate, pori diameter 2.5–4.5 µm (SEM), number of pori 10– 16, pori sunken, operculate, operculum ornamented with 10–16 microechini (SEM),

47

ACCEPTED MANUSCRIPT narrow annulus (LM); scabrate (LM), microechinate, perforate (SEM), microechini evenly spaced.

PT

Occurrence: Caryophyllaceae pollen is rare in most samples from the Tınaz section (Fig. 3); only in sample S153617 is it rather abundant (13 %).

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Remarks: The closest morphological similarity (exine architecture, narrow annulus, exine sculpturing) of Caryophyllaceae gen. indet. is with the Moehringia trinervia type

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of Punt and Hoen (1995). Caryophyllaceae have previously been reported from middle Miocene palynofloras of western Turkey (see Bouchal et al., 2016a).

Plumbaginaceae Jussieu

ED

MA

This variable pollen type may include more than one biological species.

PT

Subfamily Staticoideae Kusnezow

Staticoideae gen. indet. (fine reticulate)(PlateXI, 5–7)

CE

Description: Pollen fragment, fragment diameter 40–46 µm (LM); semitectate, exine

AC

4–5 µm thick (LM), nexine thinner than sexine; microreticulate (LM, SEM), muri 0.5–1 µm wide (SEM), muri form a hexa- to pentagonal reticulum, muri crested with nanoechini, muri with vertical microrugulae. Occurrence: Single occurrence in sample S153621. Remarks: Nowicke and Skvarla (1977) identified two pollen types in extant Plumbaginaceae; the Plumbago type (clavate sculpturing, present in all Plumbaginoideae and Staticoideae pro parte) and Armeria type (reticulate to microreticulate sculpturing, Staticoideae pro parte). A distinct pollen dimorphism, including coarse- and fine-reticulate pollen grains, has been documented in genera 48

ACCEPTED MANUSCRIPT with Armeria type pollen (Erdtman and Dunbar, 1966; Nowicke and Skvarla, 1977; Turner and Blackmore, 1984; El-Ghazaly, 1991;). The fractured single specimen

PT

available can be assigned securely to the finely-reticulate Armeria type. Staticoideae gen. indet. 2 of Bouchal et al. (2016a, p. 52, figs 19J–L) is morphologically similar in

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reticulum sculpturing, but differs in its wider lumina.

NU

Polygonaceae Jussieu

Polygonum sp. (Pl.VIII. 17-18)

MA

Genus Polygonum Linnaeus

Occurrence: A single specimen in sample S153598 (Fig. 3).

PT

ED

Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 52, figs 20D–F)

Genus Rumex Linnaeus

CE

Rumex sp. (Pl.VIII, Fig. 19-20)

AC

Occurrence: Present in nearly the entire Tınaz section (Fig. 3). Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 52, figs 20A–C).

Asterids Ericales Berchthold et Presl Sapotaceae Jussieu Sapotaceae gen. indet. (PlateXI, 8–10) Description: Pollen, shape prolate, outline elliptic in equatorial view, length of polar axis 30–40 µm (LM), 24–32 µm (SEM), equatorial diameter 25–34 µm (LM), 20–28 49

ACCEPTED MANUSCRIPT µm (SEM); eutectate, exine 2–3.5 µm thick (LM), nexine thinner than sexine, nexine thickened in aperture area (costa), costa not reaching apical ends of colpus;

PT

tetracolporate, ectocolpus length 3/4 to 5/6 of length of polar axis, endoporus elliptic,

nanoverrucae and perforations evenly spaced. Occurrence: Present in sample S153604.

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framed by costa; scabrate (LM), nanoverrucate, perforate, weakly rugulate (SEM),

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Remarks: Harley (1991) comprehensively studied the pollen morphology of extant Sapotaceae and identified 12 major pollen types with 49 subtypes in total; the

MA

encountered specimen corresponds in size, aperture and exine sculpturing to Harley’s pollen type IA. This pollen type occurs in genera of subfamily Sapotoideae tribus

ED

Sapoteae (Swenson and Anderberg, 2005), e.g. Mimusops Linnaeus, Payena A. de

PT

Candolle, Madhuca Hamilton ex J. F. Gmelin. Genera of this tribus have a palaeotropical distribution (Pennington, 2004). Dispersed fossil Sapotaceae pollen has

CE

been assigned to the form genus Sapotaceoideaepollenites R. Potonié, P. W. Thomson

AC

et Thiergart ex R. Potonié (e.g. Stuchlik et al., 2014). Previous reports of pollen assigned to Sapotaceae from middle Miocene palynofloras of western Turkey: Tetracolporopollenites spp. (Ediger, 1990, fig. 6.17; Akgün and Akyol, 1999, figs 9.66–67, 11.90–92; Akgün et al., 2007, appendix A;), Sapotaceae (Yavuz-Işık, 2007; Yavuz-Işık et al., 2011; Akkiraz, 2011, fig. 9.PP;; Kayseri-Özer et al., 2014a; 2014b, fig. 6.37; Bozcu et al., 2015), possibly Tetracolpites sp. (Takahashi and Jux, 1991, pl. 15, fig. 6), possibly Araliaceae (Akkiraz et al., 2012, only pl. 4., fig. 5).

50

ACCEPTED MANUSCRIPT Lamiids Garryales Lindley

PT

Eucommiaceae Engler Genus Eucommia Oliver

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Eucommia sp./Tricolpopollenites parmularis (R. Potonié) P. W. Thomson et Pflug (Plate VIII, Fig. 21)

NU

Occurrence: Present through the entire Tınaz section (Fig. 3).

MA

Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 54, figs 20D–F).

Lamiaceae Martinov

PT

Ajugoideae Kosteletzky

ED

Lamiales Bromhead

Ajugoideae gen. indet. (Plate XI, 11–13)

CE

Description: Pollen, shape prolate, outline lobate to circular in polar view, equatorial

AC

diameter 35–41 µm (LM, SEM); eutectate, exine 2.5–3.5 µm thick (LM), nexine thinner than sexine; triolpate, colpus length length 2/3 to 3/4 of polar axis, colpus membrane sculptured; s microechinate (LM, SEM), perforate (SEM), microechinus diameter 0.3–1 µm (SEM), microechinus length 0.3–0.1 µm (SEM). Occurrence: Present in sample S153621. Remarks: Pollen of similar size, exine architecture, aperture, and exine sculpturing occurs in the subfamily Ajugoideae, e.g. Trichostema Gronovius, Schnabelia HandelMazzetti, Clerodendrum Linnaeus (Abu-Asab and Cantino, 1989; Trudel and Morton, 1992; Abu-Asab and Cantino, 1993; Suhua et al., 2003; Bendiksby et al., 2011). Under 51

ACCEPTED MANUSCRIPT LM investigation, this pollen type appears to have strong morphological similarity to Convolvulus sp. (Pl. XIII. 1–3), however scanning electron micrographs reveal

PT

distinctly smaller pores and microechini with a wider size range in Ajugoideae gen. indet. Morphologically similar pollen has been reported from several middle Miocene

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palynofloras of western Turkey: Tricolpopollenites sp. (Akgün and Akyol, 1999, figs 12.44–45), Tricolpopollenites robusteus (Song, Li et Zhong) Takahashi et Jux

MA

Oleaceae Hoffmannsegg et Link

NU

(Takahashi and Jux, 1991, pl. 14, fig. 14).

Oleaceae types 1 to 6 correspond in size, exine architecture, aperture, and

ED

microreticulate to reticulate exine sculpturing to extant and fossil Oleaceae pollen

PT

(Renault-Miskovsky et al., 1976; Nilsson, 1988; Punt et al., 1991; Sachse, 2001; Beug, 2004; Stuchlik et al., 2014). Pollen of Oleaceae has previously been reported from

CE

middle Miocene palynofloras of western Turkey (e.g. Akgün et al., 2007; Bouchal et

AC

al., 2016a) and is present in all samples of the Tınaz section. Precise determination to genus level is difficult, owing to the high degree of morphological variability and parallel evolution in pollen of Oleaceae [e.g. Olea Linnaeus (Nilsson, 1988); Fraxinus Linnaeus (compare Renault-Miskovsky et al., 1976; Punt et al., 1991; Guo et al., 1994; Jones et al., 1995; Li et al., 2010; Miyoshi et al., 2011; ). Therefore, oleaceous pollen has not been differentiated in the pollen diagram (Fig. 3).

52

ACCEPTED MANUSCRIPT Oleaceae gen. indet. 1 aff. Fraxinus (Plate XI, 14–16) Description: Pollen, shape spheroidal to prolate, outline elliptic to weakly circular in

PT

equatorial view, length of polar axis 28–32 µm (LM), 25–29 µm (SEM), equatorial diameter 24–28 µm (LM), 23–26 µm (SEM); semitectate, exine 2–3 µm thick (LM),

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nexine thinner than sexine; tricolporate, ectocolpus length 3/4 to 5/6 of polar axis; microreticulate (LM, SEM), muri width 0.4–0.7 µm (SEM), muri crested with weak

NU

perpendicular ridges, lumina narrow and of irregular shape, decreasing in size towards colpus, psilate (SEM).

MA

Occurrence: Single occurrences in samples S153601, S153605, and S153621. Remarks: This pollen type has closest morphological similarities (microreticulum,

ED

weak perpendicular ridges) to extant Fraxinus pollen (e.g. Guo et al., 1994; Punt et al.,

PT

1991). Bouchal et al. (2016a, p. 54 figs 21G–I) reported pollen with affinities to

CE

Fraxinus from the Eskihisar lignite mine.

AC

Oleaceae gen. indet. 2 (Plate XII, 1–3) Description: Pollen, shape spheroidal to slightly prolate, outline elliptic in equatorial view, length of polar axis 23–33 µm (LM), 20–28 µm (SEM), equatorial diameter 17– 23 µm (LM), 13–20 µm (SEM); semitectate, exine 2.5–3.5 µm thick (LM), nexine thinner than sexine; triolporate, ectocolpus length 1/2 to 2/3 of polar axis (LM); reticulate (LM, SEM), muri width 0.3–0.5 µm (SEM), muri crested with sharp edged ridges, perpendicular to muri, lumina wide and of irregular shape, lumina psilate (SEM).

53

ACCEPTED MANUSCRIPT Occurrence: Present in samples S153594 to S153610. Remarks: Oleaceae gen. indet. 2 differs from type 1 by having muri crested with

PT

sharp-edged perpendicular ridges and wider lumina. These characters are present in extant Linociera obtusifolia (Lamarck) H. Perrier, I. Degener et L. A. S. Johnson,

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Noronhia linocerioides H. Perrier, and Phillyrea angustifolia Linnaeus (CerceauLarrivall et al., 1984; Renault-Miskovsky et al., 1976; Sachse, 2001). A similar type of

NU

pollen has been reported previously from the Eskihisar lignite mine (Bouchal et al.,

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2016a, Oleaceae gen. indet. 2, p. 56, figs 21J–L).

Oleaceae gen. indet. 3 (Plate XII, 4–6)

ED

Description: Pollen, shape spheroidal to weakly oblate, outline circular in equatorial

PT

view, lobate to circular in polar view, length of polar axis 17–20 µm (LM), equatorial diameter 18–21 µm (LM), 20–24 µm (SEM); semitectate, exine 2–3 µm thick (LM),

CE

nexine thinner than sexine; tricolporate, ectocolpus length 1/2 to 2/3 of polar axis (LM,

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SEM); reticulate to microreticulate (LM, SEM), muri width 0.4–0.7 µm (SEM), muri crested with ridges and nanoechini, ridges perpendicular to muri, lumina of irregular shape and size, nexine with free-standing collumellae (SEM). Occurrence: Present through the entire Tınaz section. Remarks: Oleaceae gen. indet. 3 differs from Oleaceae gen. indet. 1 and 2 in smaller size, presence of weak nanoechini and lumina with free-standing collumellae. These characters are present in extant Olea Linnaeus, Fontanesia Labillardière and Osmanthus Loureiro (Guo et al., 1994; Li et al., 2010; Nilsson, 1988; RenaultMiskovsky et al., 1976; Sachse, 2001). A similar type of pollen has been reported from 54

ACCEPTED MANUSCRIPT the Eskihisar lignite mine, Yatağan Basin by Bouchal et al. (2016a, Oleaceae gen.

PT

indet. 3, figs 22A–C).

Oleaceae gen. indet. 4 (Plate XII, 7–9)

SC RI

Description: Pollen, shape spheroidal to oblate, outline circular in equatorial view, lobate to circular in polar view, length of polar axis 18–25 µm (LM), 17–22 µm

NU

(SEM), equatorial diameter 20–24 µm (LM), 19–21 µm (SEM); semitectate, exine 2–3 µm thick (LM), nexine thinner than sexine; tricolporate, ectocolpus length 1/2 to 2/3

MA

of polar axis (LM, SEM); reticulate to microreticulate (LM, SEM), muri width 0.4– 0.7 µm (SEM), muri crested with nanoechini and weak perpendicular ridges, lumina of

ED

irregular shape and size, lumina with free-standing collumellae (SEM).

PT

Occurrence: Present through the entire Tınaz section. Remarks: Oleaceae gen. indet. 4 is similar in pollen morphology (size, lumina with

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free-standing collumellae) to Oleaceae gen. indet. 3 but differs in having muri crested

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with distinct nanoechini. Such characters are found in extant Olea and Chionanthus Linnaeus (Guo et al., 1994; Nilsson, 1988; Punt et al., 1991; Renault-Miskovsky et al., 1976; Sachse, 2001). A similar type of pollen has been reported from the Eskihisar lignite mine, Yatağan Basin by Bouchal et al. (2016a, Oleaceae gen. indet. 4, figs 22D–F).

Oleaceae gen. indet. 5 (PlateXII, 10–12) Description: Pollen, shape spheroidal, outline circular in equatorial view, lobate to circular in polar view, length of polar axis 21–26 µm (LM), 18–22 µm (SEM), 55

ACCEPTED MANUSCRIPT equatorial diameter 18–23 µm (LM), 16–20 µm (SEM); semitectate, exine 2–2 .5µm thick (LM), nexine thinner than sexine; tricolporate, ectocolpus length 1/2 to 2/3 of

PT

polar axis (LM, SEM); reticulate (LM, SEM), muri width 0.4–0.6 µm (SEM), muri smooth, some having a central ridge, lumina of irregular shape and size, lumina with

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freestanding columellae (SEM). Occurrence: Present in sample S153621.

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Remarks: Oleaceae type 5 differs from types 1 to 4 in having smooth muri with a central ridge. Similar pollen is is produced by extant Olea and Osmanthus (Guo et al.,

MA

1994, figs 13A–D; Nilsson, 1988, fig. 1D).

ED

Oleaceae gen. indet. 6 (Plate XII, 13–15)

PT

Description: Pollen, shape spheroidal, outline elliptic in equatorial view, circular to lobate in polar view, length of polar axis 29–34 µm (LM), 27–32 µm (SEM),

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equatorial diameter 28–32 µm (LM), 26–30 µm (SEM); atectate, exine 3.5–5 µm thick

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(LM), nexine thinner than sexine; tricolporate, ectocolpus length 1/2 to 2/3 of polar axis (LM, SEM); reticulate (LM, SEM), muri width 0.8–1.2 µm (SEM), muri smooth, lumina of irregular shape and size, lumina psilate (SEM) Occurrence: A single specimen in sample S153621. Remarks: Oleaceae type 6 differs from types 1 to 5 in its larger size, wider lumen and wider muri. Similar pollen is produced by extant Ligustrum Linnaeus and Syringa Linnaeus (Guo et al., 1994; Li et al., 2010; Punt et al., 1991; Zhang, 1982).

56

ACCEPTED MANUSCRIPT Solanales Jussieu ex Berchthold et Presl Convolvulaceae Jussieu

PT

Genus Convolvulus Linnaeus Convolvulus sp. (Plate XIII, 1–3)

SC RI

Description: Pollen, shape prolate, outline elliptic in equatorial view, length of polar axis 41–46 µm (LM), equatorial diameter 30–36 µm (LM), 27–32 µm (SEM);

NU

eutectate, exine 3–5 µm thick (LM), nexine thinner than sexine; tricolpate; scabrate (LM), nanoechinate, perforate (SEM), nanoechinus diameter 0.2–0.4 µm (SEM).

MA

Occurrence: Single occurences in samples S153598, S153601, S153605, and S153621. Remarks: This pollen corresponds in pollen wall architecture, colpate aperture and

ED

echinate perforate exine sculpturing to extant Convolvulus (Sengupta, 1972; C.

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arvensis type in Cronk and Clarke, 1981; El-Ghazaly, 1991). Owing to the poor preservation of this pollen, it is difficult to distinguish from pollen of Ajugoideae

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under LM. Therefore, we provide full documentation (LM and SEM) of these

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specimens. Convolvulus pollen has been reported from the Eskihisar lignite mine, Yatağan Basin, by Bouchal et al. (2016a, p. 54, figs 21A–C).

Campanulids Aquifoliales Senft Aquifoliaceae Berchthold et Presl Genus Ilex Linnaeus Ilex sp. (Plate XIII, 4–7)

57

ACCEPTED MANUSCRIPT Description: Pollen, outline circular to weakly convex triangular in polar view, equatorial diameter 26–30 µm (LM), 24–28 µm (SEM); semitectate, exine 1–3 µm

PT

thick (LM), nexine thinner than sexine; tricolporate, ectocolpus indistinct (SEM); clavate (LM, SEM), two major types of clavae present: in the apocolpium and

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mesocolpium areas large clavae (length <2.5 µm, diameter <2.5 µm) with rugulate sculpturing (Pl. XXIII, 6) are developed; adjacent to the colpus and between large

evident.

MA

Occurrence: Present in sample S153604.

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clavae, small clavae or nanoverrucae (ca 0.2–0.4 µm diameter) (Pl. XXIII, 7) are

Remarks: Clavate exine ornamentation is charachteristic of extant Ilex pollen (e.g.

ED

Lobreau-Callen, 1975) and the fossil-genus Ilexpollenites Thiergart ex R. Potonié (e.g.

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Stuchlik et al., 2014).

Previous reports of Ilex from middle Miocene palynofloras of western Turkey:

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Ilexpollenites ilacus (R. Potonié) Thiergart ex R. Potonié and I. marginatus (R.

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Potonié) Thiergart (Takahashi and Jux, 1991, pl. 18, only figs 11–13; Akgün et al., 2007), “Ilex-Typen” of Benda (1971), Ilex (Yavuz-Işık, 2007; Akkiraz, 2011, fig. 9.OO; Akkiraz et al., 2012, pl. 4, figs 12–13; Bozcu et al., 2015). Fossil leaves of Ilex have been reported from the early/middle Miocene of western Anatolia of Balya, Balıkesir (Engelhardt, 1903) and the Soma Basin (Nebert, 1978; Gregor, 1990).

58

ACCEPTED MANUSCRIPT Asterales Link Asteraceae Berchthold et Presl

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Asteroideae types 1 to 4 (Pl. VIII, Figs. 22–25) can be assigned securely to subfamily Asteroideae (Punt and Hoen, 2009), but further attribution to a genus is not possible

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owing to their poor preservation. Asteroideae types have not been treated separately for the pollen diagram (Fig. 3). Asteraceae have been reported previously from middle

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Miocene palynofloras from western Turkey (e.g. Akgün et al., 2007; Bouchal et al., 2016a).

MA

Occurrence: Pollen of this subfamily is rare (0.25–1 %) in most samples of the Tınaz

ED

section, except in sample S153617 where it reaches 6.25 %.

PT

Asteroideae type 1 (Plate VIII, 22)

Description: Pollen, shape spheroidal, outline circular in equatorial view, length of

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polar axis 20–25 µm (LM), equatorial diameter 20–25 µm (LM) 18–22 µm (SEM);

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exine including echini 3–5 µm thick (LM), sexine thicker than nexine; tricolporate; echinate, perforate (LM, SEM), echini basal diameters 3–4 µm (SEM), echini blunt, echinus height 2–3 µm (SEM), perforations extending to upper third of echini.

Asteroideae type 2 (Plate VIII, 23) Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 45, figs 22J–L, as Asteroideae type 2).

59

ACCEPTED MANUSCRIPT Asteroideae type 3 (Plate VIII, 24) Description: Pollen, shape spheroidal, outline circular to lobate in polar view, polar

PT

equatorial diameter 40–70 µm (LM), 38–60 µm (SEM); exine including echini 6–15 µm thick (LM), sexine thicker than nexine; tricolporate; echinate, perforate (LM,

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SEM), echini basal diameters 4–5 µm (SEM), echini blunt, echini length 4–6 µm

NU

(SEM), perforations extending to upper third of echini.

Asteroideae type 4 (Plate VIII, 25)

MA

Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 45, figs 23A–C,

ED

as Asteroideae type 4).

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Cichorioideae

Cichorioideae types 1 and 2 were combined in the pollen diagram (Fig. 3). They have

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lophate/lacunate sculpturing and, therefore, can be assigned securely to

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Chichorioideae, with closest morphological similarity (presence of abporal and paraporal lacunae) to the Cichorium intybus type of Blackmore (1984). Occurrence: Cichorioideae pollen is present in samples S153610, S153617, and S153621.

Cichorioideae type 1 (Pl. VIII, Fig. 26)) Description: Pollen, shape spheroidal, outline polygonal, diameter 33–39 µm (LM), 26–31 µm (SEM); lophate, exine including lophae and echini 4–8 µm thick (LM), nexine thinner than sexine; tricolporate; lophate, echinate, perforate (LM, SEM), 60

ACCEPTED MANUSCRIPT echini basal diameters 2–3 µm (SEM, LM), echini length 1.5–2.5 µm (SEM), perforations reaching to lower third of echini, echini present only on lophae, lophae

PT

perforate, paraporal lacuna perforate, abporal lacuna psilate.Remarks: This pollen type is fairly similar to Cichorioideae gen. indet. of Bouchal et al. (2016a, fig. 23G). Owing

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to the poor preservation of the former, we cannot determine whether they belong to

Cichorioideae type 2 (Pl. VIII, Fig. 27)

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different taxa or not.

MA

Description: Pollen, shape spheroidal, outline polygonal, diameter 30–36 µm (LM), 27–32 µm (SEM); lophate, exine including lophae and echini 4–9 µm thick (LM),

ED

nexine thinner than sexine; tricolporate; lophate, echinate, perforate (LM, SEM),

PT

echini basal diameters 2–3 µm (SEM, LM), echini length 2–3 µm (SEM), perforations reach to lower third of echini, echini present only on lophae, lophae perforate,

CE

paraporal lacuna perforate, abporal lacuna psilate, perforations larger than in

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Cichorioideae type 1.

Campanulaceae Jussieu Campanuloideae Campanuloideae gen. indet. (Plate XIII, 8–10) Description: Pollen, spheroidal to oblate, outline circular in polar view, length of polar axis 20–30 µm (LM), equatorial diameter 26–35 µm (LM), 23–31 µm (SEM); exine including microechini 1.5–2.5 µm thick (LM), nexine and sexine about same thickness; triporate; microechinate (LM), microechinate, striato-microreticulate 61

ACCEPTED MANUSCRIPT (SEM), microechini basally divided, microechini basal diameters 0.4–0.7 µm (SEM), microechini length 0.5–1 µm (SEM).

PT

Occurrence: Present in samples S153601, S153602, S153610, and S153621. Remarks: Porate, microechinate, striato-microreticulate pollen is common in several

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Campanuloideae genera, e.g. Asyneuma Grisebach et Schenk, Campanula Linnaeus, Phyteuma Linnaeus, and Wahlenbergia Schrader ex Roth (e.g. Dunbar, 1975;

MA

Dipsacales Jussieu ex Berchthold et Presl

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Khansaria et al., 2012).

Caprifolioideae Eaton

PT

Genus Lonicera Linnaeus

ED

Caprifoliaceae Jussieu

Lonicera sp./Lonicerapollis grandis Grabowska (Plate XIII, 11–13)

CE

Description: Pollen, shape oblate, outline circular to convex triangular in polar view,

AC

equatorial diameter 53–61 µm (LM), 47–53 µm (SEM); eutectate, exine without echini 3–4 µm thick (LM), nexine thinner than sexine; brevitricolporate, apertures weakly protruding, ectocolpi short; echinate (LM), echinate, perforate (SEM), perforations irregular distributed, echini basal diameters 0.8–1.2 µm (SEM), echini length 1.5–2 µm (SEM), echini blunt. Occurrence: Present in samples S153595, S153601, and S153608. Remarks: Pollen of Lonicera has been reported previously from middle Miocene pollen floras of western Turkey (e.g. Akgün et al., 2007; Bouchal et al., 2016a).

62

ACCEPTED MANUSCRIPT Lonicera sp. 1 and 2 of Bouchal et al. (2016a, p. 58–60, figs 23J–L and 24A–C) differ

PT

by their more widely spaced and less numerous echini.

Linnaeoideae gen. indet. (Plate VIII, 28–29) Occurrence: Present in sample S153604.

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Linnaeoideae Rafinesque

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Remarks: For descriptions and remarks see Bouchal et al. (2016a, p. 60, figs 24D–F).

MA

Dipsacoideae Eaton

Pollen of this subfamily was combined in the pollen diagram (Fig. 3) and reaches its

ED

highest diversity in sample S153617. Dipsacoideae have been reported from middle

PT

Miocene palynofloras of western Turkey (e.g. Akgün et al., 2007; Bouchal et al.,

CE

2016a).

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Genus Dipsacus Linnaeus vel Cephalaria Schrader Dipsacus vel Cephalaria sp. 1 (Plate XIII, 14–16) Description: Pollen, shape spheroidal, outline circular to weakly lobate in polar view, length of polar axis 72–78 µm (LM), 64–69 µm (SEM), equatorial diameter 69–74 µm (LM), 61–67 µm (SEM); eutectate, exine 5–8 µm thick, nexine thinner than sexine (LM), nexine nearly absent in aperture area (LM); brevitricolpate to triporate, colpus length 1/4 to 1/3 of polar axis length, margo present (LM, SEM), margo formed by sexine (LM); microechinate (LM), microechinate, nanoechinate, perforate (SEM), margo perforate, nanoechinate. 63

ACCEPTED MANUSCRIPT Occurrence: Present in samples S153601, S153606, S153617, and S153621.

PT

Dipsacus vel Cephalaria sp. 2 (Plate XIV, 1–6) Description: Pollen, shape subspheroidal, outline convex triangular to hexangular in

SC RI

polar view, elliptic to weakly quadrangular in equatorial view, length of polar axis 81– 106 µm (LM), 84–91 µm (SEM), equatorial diameter 84–102 µm (LM), 61–67 µm

NU

(SEM); eutectate, exine including echini 8–16 µm thick, sexine twice the thickness of nexine, except in polar area sexine three times the thickeness and in aperture area

MA

nexine nearly absent; brevicolpate to porate (3), colpus length 1/4 to 1/3 of polar axis, margo present (LM, SEM), margo formed by sexine, aperture membrane ornamented

ED

with long echini (3–9 µm); echinate (LM), echinate, nanoechinate, perforate (SEM),

PT

margo psilate.

Occurrence: Present in samples S153605, S153610, and S153617.

CE

Remarks: Dipsacus vel Cephalaria sp. 2 pollen differs from sp. 1 mainly by greater

AC

dimensions and thicker exine. Both show close resemblance (exine architecture, aperture, margo presence, exine sculpturing) to pollen of extant species of Dipsacus and Cephalaria (e.g. Dipsacus fullonum type in Clarke and Jones, 1981; Barento et al., 1987; Beug, 2004; Miyoshi et al., 2011).

Genus Scabiosa Linnaeus Scabiosa sp. (Plate XIV, 7–9) Description: Pollen, shape prolate, outline hexagonal in polar view, elliptic to rhombic in equatorial view, length of polar axis 62–68 µm (LM), 43–56 µm (SEM), equatorial 64

ACCEPTED MANUSCRIPT diameter 49–57 µm (LM), 38–46 µm (SEM); eutectate, exine including echini 4–10 µm thick, nexine and sexine of same thickness except in polar area sexine thickened

PT

and in aperture area nexine nearly absent; brevitricolpate, colpus length 1/3 to 1/2 of polar axis (LM, SEM), aperture sunken; echinate (LM), echinate, microechinate,

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perforate (SEM), echini basal diameters 1.2–1.8 µm (SEM), echini length 2–3 µm (SEM).

NU

Occurrence: Present in samples S153608 and S153617.

Remarks: This pollen has closest similarities (outline, exine architecture, aperture,

MA

exine sculpturing) with pollen of extant Scabiosa (e.g. Barento et al., 1987; Beug,

ED

2004; Scabiosa columbaria type of Clarke and Jones, 1981; Miyoshi et al., 2011).

Apiaceae Lindley

PT

Apiales Nakai

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Various Apiaceae pollen types were combined for the pollen diagram (Fig. 3). For the

AC

description of outer and inner contours, the nomenclature of Punt (1984) was used. Apiaceae have previously been reported from middle Miocene palynofloras of western Turkey (e.g. Akgün et al., 2007; Bouchal et al., 2016a). Although an attempt was made to group the Apiaceae pollen into distinct types, we do not exclude the possibility that this family was more diverse than indicated here.

Apiaceae type 1 (PlateXIV, 10–13) Description: Pollen, shape prolate, length of polar axis 26–31 µm (LM), 20–24 µm (SEM), equatorial diameter including protruding apertures 15–19 µm (LM), 11–15 µm 65

ACCEPTED MANUSCRIPT (SEM); eutectate, exine 2–2.5 µm thick in polar area (LM), outer contour straight, apiculate, inner contour convex; tricolporate, slit-like ectocolpus with protruding

PT

sexine in porus area (LM, SEM), ectocolpus length 1/2 to 2/3 of polar axis, endoporus circular; scabrate (LM), microrugulate, fossulate, perforate (SEM), perforation density

SC RI

higher in apocolpium and mesocolpium (SEM), rugulae densely packed in equator and ectocolpus area (SEM).

NU

Remarks: Pollen of this type falls into the morphological range of the Aegopodium

Apiaceae type 2 (PlateXIV, 14–17)

MA

podagraria type of Punt (1984).

ED

Description: Pollen, shape prolate, length of polar axis 15–18 µm (LM), 12–15 µm

PT

(SEM), equatorial diameter including protruding apertures 8–11 µm (LM), 6–9 µm (SEM); eutectate, exine 1.5–2 µm thick in polar area (LM), outer contour straight,

CE

apiculate, inner contour convex; tricolporate, slit-like ectocolpus (LM, SEM),

AC

ectocolpus length 1/2 to 2/3 of polar axis, endopori circular; psilate (LM), microrugulate, fossulate, perforate (SEM), perforations more abundant in apocolpium and mesocolpium (SEM), rugulae densely spaced in equator and ectocolpus area (SEM). Remarks: Apiaceae type 2 is similar to type 1 (short colpus, outer and inner contour, exine sculpturing) but has a distinctly smaller size range.

66

ACCEPTED MANUSCRIPT Apiaceae type 3 (PlateXV, 1–4) Description: Pollen, shape prolate, length of polar axis 26–30 µm (LM), 19–24 µm

PT

(SEM), equatorial diameter including protruding apertures 13–16 µm (LM), 9–12 µm (SEM); eutectate, exine 1.5–2 µm thick in polar area, costa 2.5–3 µm thick (LM),

SC RI

outer contour straight to weakly convex, inner contour convex; tricolporate, slit-like ectocolpus (LM, SEM), ectocolpi length 3/4 to 5/6 of polar axis, endopori circular to

NU

elliptic; scabrate (LM), microrugulate, fossulate (SEM), rugulae densely packed adjacent to ectocolpus (SEM).

MA

Remarks: Pollen of this type falls into the morphological range of Apium inundatum

ED

type of Punt (1984).

PT

Saniculoideae Thorne ex P. Royen

Saniculoideae gen. indet. (PlateXV, 5–8)

CE

Description: Pollen, shape prolate, outline weakly bone-shaped in equatorial view,

AC

length of polar axis 48–54 µm (LM), 44–50 µm (SEM), equatorial diameter 22–24 (LM), 18–21 µm (SEM); eutectate, exine 2–3 µm thick in polar area (LM), outer contour straight, weakly apiculate, inner contour straight to weakly convex; tricolporate, slit-like ectocolpus (LM, SEM), ectocolpi length 3/4 to 5/6 of polar axis, endopori elliptic, costae nearly extending around equator (endocingulum); scabrate (LM), microrugulate, fossulate, perforate (SEM), sculptural elements fused and perforations more abundant in apocolpium, central mesocolpium and colpus area (SEM), microrugulae loosely packed between central mesocolpium and aperture area (SEM). 67

ACCEPTED MANUSCRIPT Occurrence: Present in most samples of the Tınaz section (Fig. 3). Remarks: Saniculoideae gen. indet. corresponds (inner and outer contour, aperture,

PT

sculpturing) to the Astrantia (major) type pollen of various authors, e.g. Beug (2004), Cerceau-Larrivall et al. (1984), Punt (1984). Pollen of Saniculoideae has been reported

SC RI

previously from the Yatağan Basin (Bouchal et al., 2016a, figs 26A–C).

NU

3.5.3 Dicotyledonae incertae sedis Pollen type 1 (PlateXV, 9–11)

MA

Description: Pollen, shape oblate, outline circular, diameter 32–36 µm (LM), 30–34 µm (SEM); eutectate, exine 2–2.5 µm thick (LM), sexine and nexine about same

ED

thickness (LM); brevitricolpate, apertures weakly protruding in equatorial region,

perforate (SEM).

PT

colpus membrane microechinate; rugulate (LM), rugulate, nanoechinate, weakly

CE

Occurrence: Present in sample S153621.

AC

Remarks: We were unable to discern similarities with any particular modern family.

Pollen type 2 (PlateXV, 12–15) Description: Pollen, outline circular in polar view, equatorial diameter 32–37 µm (LM), 30–34 µm (SEM); semitectate, exine 2–2.5 µm thick, nexine thinner than sexine (LM); tricolporate, colpus membrane granulate, pori circular (LM, SEM); reticulate (LM, SEM), lumina decreasing in size in colpus adjacent and polar area (SEM). Occurrence: Present in samples 153594, 153595, and 153599 (Fig. 3).

68

ACCEPTED MANUSCRIPT Remarks: Morphological similarities are with Rubiaceae pollen; similar microreticulate, tricolporate pollen with granular colpus membrane is present for

PT

instance, in Coffea Linnaeus (Chinnappa and Warner, 1981) and Ophiorrhiza

SC RI

Linnaeus (Huang, 1972; Miyoshi et al., 2011).

Pollen type 3 (PlateXV, 16–18)

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Description: Pollen, shape spheroidal, outline circular in equatorial view, equatorial diameter 11–13 µm (LM), 10–12 (SEM); tectate, exine 0.8–1.2 µm thick (LM);

MA

tricolpate (LM, SEM); psilate (LM), microechinate, perforate (SEM). Occurrence: Present in sample S153621.

PT

ED

Remarks: We were unable to determine affinities with any particular modern family.

4. Discussion

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4.1. Pollen zones of the Tınaz lignite mine section

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Bisaccate gymnosperm pollen (2–40 %) and pollen of Fagaceae (5–30 %) are the dominant elements through the entire Tınaz lignite mine section. Three informal pollen zones (pollen zones 1, 2, and 3) can be distinguished. 4.1.1. Pollen Zones 1 and 2 Pollen zone 1 is associated with the lignite seam, its intercalated sediments, and the underlying siltstones (samples S153594–S153599). It is characterised by a high abundance of spores (3.75–59.25 %). Pollen of Salix and Liquidambar was observed only in this pollen zone; furthermore, an Alnus peak (<20 %) was encountered in sample S153599. This phase of basin development was characterised by lignite 69

ACCEPTED MANUSCRIPT accumulation and a change from fluvial (high-energy) to lacustrine (low-energy) environments. The lignite-producing swamp vegetation appears to have consisted

PT

mainly of ferns (Osmundaceae), Alnus, and Decodon; taxodiaceous Cupressaceae and Myricaceae were also part of this vegetation unit, but are of lower abundance in this

SC RI

pollen zone (papillate Cupressaceae, Corylus/Ostrya/Myricaceae in Fig. 2). Myrica leaf impressions occur locally and Taxodium sp. and Glyptostrobus europaeus

NU

(Brongniard) Unger are only rarely encountered in the macrofossil record of the lignite mines of the Yatağan Basin (Güner, 2016).

MA

Further characteristics of pollen zone 1 are relatively high amounts of Pinaceae bisaccate pollen, very low percentages of fairly diversefied herbaceous taxa, and

ED

moderate abundance of Fagus (up to >5 %) and Quercus (Quercus Group Ilex, Group

PT

Cerris, and Group Quercus; combined to ca 15 %). These represent vegetation units thriving on well-drained soils in the hinterland of the swamps.

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Lignite and fluviatile facies and their associated pollen taxa (Alnus, papillate

AC

Cupressaceae, fern spores) are difficult to correlate within a basin, because these facies wedged out laterally during depositon. Therefore, pollen zone 1 is only of limited use for intrabasinal and regional correlation. Pollen zone 2 is represented in the lacustrine marlstones and clayey limestones above the main lignite seam. Here, spores are much less common (<1.5 %), pollen of Typhaceae is more abundant in several samples (10–34 %), and a high diversity and abundance of woody angiosperm taxa (22–49 %) is evident. Quercus pollen amounts to ca 30 %.

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ACCEPTED MANUSCRIPT It is very likely that vegetation types of pollen zones 1 and 2 in the Tınaz section comprise forest vegetation and that various degrees of herbaceous pollen mostly

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represent moist subhabitats of the lake shore vegetation (Figs 2, 3). Arboreal pollen (AP) range from >70 % to >90 % in pollen zone 1, which is derived from lignite-

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dominated layers. AP pollen percentages are lower in pollen zone 2, representing lacustrine sedimentation. Although most of the samples have >60 % AP, the two

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topmost samples of this zone have less than 50 % AP. This is due to high levels of Typhaceae and Amaranthaceae pollen, but the abundance and diversity of woody

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angiosperm pollen remain high. Rather than reflecting a shift to generally open landscapes, this may reflect changes in the lake shore/water table accompanying the

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development of a Typha belt and pioneer vegetation. This has also been observed in

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Quaternary sediments of Turkey (van Zeist et al., 1975). 4.1.2. Transitional zone and pollen zone 3

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Samples S153608 and S153610 represent a transitional zone between pollen zones 2

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and 3. Here, a change in the ratio of herbaceous and woody taxa is apparent. In sample S153608, Amaranthaceae (51 %) dominate, and a decrease in diversity and abundance of woody angiosperm taxa (13.5 %) is evident. In sample S153610, the abundance (28.75 %) and the diversity of woody angiosperm taxa increases again but also the diversity in herbaceous taxa has increased. Woody angiosperms present in pollen zones 1 and 2, such as Ilex, Carpinus, Engelhardioideae, Tilia, Sapotaceae, and Celtis do not occur in this transitional zone and the following pollen zone 3. It is suggested, that during this phase, the lake margin was shallow enough for the development of a reed belt that fell dry episodically and was colonized by pioneer plants (e.g. 71

ACCEPTED MANUSCRIPT Amaranthaceae). The only occurrence of Nitraria in the Tınaz section is from this transitional zone. Today, Nitraria is a typical element of the semi-arid to arid belt

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stretching from northeast Asia to northern Africa, commonly occurring on saline soils (salt deserts, salt marshes) and on coastal sand dunes (Sheahan, 2011).

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The transitional zone between pollen zones 2 and 3 shows values of AP between 20 and 50 %. Amaranthaceae are the most abundant group and Quercus pollen decreases

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to ca 5–6 % as compared to >30 % in the level below. Along with the presence of Nitraria, this may indicate locally more open vegetation, with a diverse woody flora

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further from the lake. However, it may also reflect the shallowing of the lake and associated decreased influx of water-transported pollen. Keeping in mind that 95 % of

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wind-transported pollen originates from within ca 1 km radius of the area of

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sedimentation (Klaus, 1987; Traverse, 2007), this would mean that a stronger filter acted on pollen transported from further away and that the pollen diagram is biased

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towards local plants growing around the lake.

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Pollen zone 3 is represented by sample S153617 and is characterised by a high abundance and diversity of (wind-pollinated) herbaceous pollen taxa (Amaranthaceae, Asteraceae, Caryophyllaceae, Dipsacoideae, Polygonaceae) and a decrease in the abundance and diversity of woody pollen taxa. Ephedra reaches more than 10 %. Although Amaranthaceae probably grew close to the lake, Caryophyllaceae and Asteraceae may have been part of contrasting vegetation types. Values of arboreal pollen (AP) are below 20 %. Small amounts of wind-pollinated Cathaya, Cedrus, Pinus and Cupressaceae s.str. indicate the presence of woody vegetation at various distances from the area of sedimentation (see below 4.2.). 72

ACCEPTED MANUSCRIPT Among woody angiosperms, low numbers of Eucommia and Fagus pollen indicate that humid temperate vegetation did not develop in close vicinity to the lake. Large

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Quercus pollen (mainly corresponding to Quercus Group Cerris) is slightly more abundant (ca 2 %) than small Quercus pollen (corresponding to Quercus Group Ilex)

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and points to the presence of oak woodlands closer to the lake, possibly co-occurring with Ephedra. Poaceae are rare and Artemisia is entirely absent. Overall, this would

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indicate locally open vegetation and/or reduced influx of water-transported pollen. The Eskihisar lignite mine section of Bouchal et al. (2016a) did not include the upper

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parts of the Sekköy Member, hence, reported pollen associations similar to pollen zone

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1 and 2.

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4.2. Comparison to the Eskihisar and Yenieskihisar lignite mine sections 4.2.1 Eskhihisar lignite mine section

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The palynofloras of the Eskihisar (Bouchal et al., 2016a) and the Tınaz lignite mine

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sections (pollen zones 1 and 2) are similar, differing only in the presence/absence of rare taxa. For instance, pollen of Celtis, Ilex, Liquidambar, Ranunculaceae, Salix, Sapotaceae, Sparganium, Campanuloideae, and Tsuga were not reported by Bouchal et al. (2016b), but occur regularly in middle Miocene sediments of Tınaz and Salihpaşalar (pers. observ. J. Bouchal, 2016) in the Yatağan Basin, and in several other localities of western Turkey (e.g. Akgün et al., 2007; Benda, 1971). Ericaceae, Onagraceae, Rosaceae, and Buxaceae have not been encountered in the Tınaz lignite mine section, although pollen of these families is present in the Eskihisar lignite mine section (Bouchal et al., 2016a) and several localities of western Turkey 73

ACCEPTED MANUSCRIPT (e.g. Akgün et al., 2007; Benda, 1971). Buxus, although absent from the pollen record of Tınaz, is among the 10 most abundant macrofossils in strata above the main lignite

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seam in the Tınaz lignite mine (Güner, 2016). This shows that finding or not finding insect-pollinated woody plants in the pollen record is to some extent stochastic.

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Pollen zone 1, recognized in the Tınaz and the Eskihisar lignite mine sections (Bouchal et al., 2016a), is restricted to the fluviatile and lignite facies and contains

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abundant spores of ferns and fern allies at both localities. Although Osmundaceae, Alnus and Decodon dominate the entire pollen zone 1 of the Eskihisar lignite mine

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section (Bouchal et al., 2016a, fig. 2), Alnus and Osmundaceae show abundance peaks only in single samples of pollen zone 1 in the Tınaz lignite mine section. Decodon

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4.2.2 Yenieskhihisar section

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shows no marked abundance peaks in the entire Tınaz lignite mine section (<4.25 %).

Yavuz-Işık et al. (2011, fig. 3) and Benda (1971) investigated the pollen flora of the

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Yenieskihisar mammal locality (Fig. 1.2, southern part of Eskihisar mine, upper most

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Sekköy Member, MN7/8, e.g. Sickenberg et al. 1975; Atalay, 1980; Fortelius, 2016), and reported a pollen assemblage with a moderately high abundance of herbaceous taxa (ca 30 %) including, for example, Apiaceae, Asteraceae, Amaranthaceae, Campanulaceae, Caryophyllaceae, Rumex, and Pinaceae (ca 20 %). Other common woody angiosperm taxa are deciduous oaks, Ulmus, Zelkova, Alnus, and Carya. Based on lithostratigraphic correlation (Fig. 1), this pollen assemblage clearly is younger than pollen zone 3 of the Tınaz lignite mine section. Yavuz-Işık et al. (2011) reported 20 % undifferentiated Pinaceae pollen and 10 % Zelkova pollen in that section. These values are much higher than those reported here for pollen zone 3. 74

ACCEPTED MANUSCRIPT Overall, the ratio of arboreal pollen to non-arboreal pollen is ca 20/80 in pollen zone 3 of the Tınaz lignite mine section but is 70/30 in the sample from Yenieskihisar,

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possibly indicating more closed local vegetation for the younger Yenieskihisar pollen assemblage. This is interesting because the occurrence of the rhinoceros Beliajevina

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grimmi Heissig, 1974 in the Yenieskihisar faunal assemblage (Fortelius, 2016) has been taken as an indication of open grass-dominated landscapes (Geraads and Saraç,

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2003).

It should be noted, however, that modern pollen spectra obtained from surface samples

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from a complex landscape in the Middle Atlas of Morocco (Bell and Fletcher, 2016) contained (30–) 45–80 (–90) % arboreal pollen depending on the individual sample.

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Thus, marked local differences in abundance of particular pollen types should be

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expected even for coeval sites in the Yatağan Basin given the distinct geographical setting of the Eskihisar lignite mine and the Tınaz lignite mine. Both the Eskihisar and

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the Tınaz lignite mines are isolated from the remaining sub-basins of the Yatağan

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Basin by small mountains (Becker-Platen, 1970).

4.3. Other early to middle Miocene palynofloras from southwestern Anatolia 4.3.1 Palynoflora of the Çatakbağyaka mammal site The Çatakbağyaka mammal site is located to the south of the Yatağan Basin (Fig. 1.2, Sekköy Member, village Bağyaka near Meke, Muğla province; Atalay, 1980; Sickenberg et al., 1975). This locality has been investigated palynologically by YavuzIşık et al. (2011, fig. 3) and Jiménez-Moreno (2005, fig. 4.77). Although the pollen diagrams shown in these studies do not include spores or algal palynomorphs, the two 75

ACCEPTED MANUSCRIPT samples of Çatakbağyaka of Yavuz-Işık et al. (2011) and samples 1 and 2 of JiménezMoreno (2005) correspond in abundance and composition mainly to pollen zones 1

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and 2 of the Eskihisar and Tınaz mine sections described here. Abundant or common taxa (e.g. Cedrus, Pinus, nonpapillate Cupressaceae, deciduous and evergreen

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Quercus, Fagus, Carya, Acer, Amaranthaceae, Apiaceae) and rare pollen taxa [e.g. papillate Cupressaceae (as “Taxodium type”), Ephedra, Euphorbiaceae, Ericaceae,

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Engelhardioideae (as “Engelhardia”), Buxaceae, Geraniaceae, Plumbaginaceae] correspond to the taxa comprising pollen zones 1 and 2 of the Eskihisar and Tınaz

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mine sections.

Sample 3 in fig. 4.77 of Jiménez-Moreno (2005) records an increase in herbaceous

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taxa (e.g. Apiaceae, Asteraceae, Amaranthaceae) and a decrease in abundance and

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diversity of arboreal pollen (ratio AP/NAP ca 50/50). This trend is also evident in the transition zone between pollen zones 2 and 3 and in pollen zone 3 of the Tınaz lignite

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mine section. In addition, both studies reported several rare mega-mesothermic

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elements (mostly single occurrences in Jiménez-Moreno, 2005), such as Alchornea, Mussaenda-type, Passifloraceae, Mimosa or Acacia. Unfortunately, these taxa were not figured from this locality. Considering the revised age of the Çatakbağyaka mammal locality (MN7/8), sample 3 of Jiménez-Moreno (2005) may correspond to the time period when the mammal fauna was deposited. Overall, this illustrates that using pollen assemblages alone, it is difficult to safely correlate sediments from Yenieskihisar (MN7/8), Tınaz pollen zone 3 (younger than MN7/8), and Çatakbağaka sample 3 (MN7/8).

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ACCEPTED MANUSCRIPT 4.3.2 Palynoflora of the Ören basin lignite mines Kayseri (2010) investigated three stratigraphic sections (Ikizler Section, Sekköy

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Section I and II) of the Sekköy Member (=Sekköy Formation in Kayseri, 2010) of the Ören Basin lignite mines (Sekköy lignite mine in Fig. 1.2). All three sections host

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abundant spores (e.g. Laevigatisporites haardti), papillate and nonpapillate Cupressaceae (Inaperturopollenites spp., Cupressacites spp.), Alnus (Alnipollenites

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verus), and Fagaceae pollen. The composition of the palynofloras of the Sekköy Member (Formation) of the Ören basin mainly concur in their compositon with the

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pollen floras of the Yatağan Basin (Akgün et al., 2007; Bouchal et al., 2016a; this study). The main difference is the intermittently high abundance of Cupressaceae in

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the Ören sections.

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Kayseri-Özer et al. (2014a) and Kayseri and Akgün (2010) studied the palynoflora of the Kultak section, Ören region, south of the Yatağan Basin. The lignites containing

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the palynoflora are stratigraphically below a mammal fauna assigned to MN5–6 (17–

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12.5 Ma, Fortelius, 2016). The Kultak palynoflora was deposited in a deltaic environment, which explains the rare occurrence of the mangrove Avicennia. Nevertheless, the Kultak flora resembles both the Tınaz and the Eskihisar lignite mine floras (pollen zone 2) by the moderate to abundant presence of Cedrus and Pinus, abundant Quercus, moderately abundant Oleaceae, Zelkova/Ulmus and others. The Kultak pollen flora differs by abundant Engelhardioideae; pollen of this subfamily is more commonly found in older (Paleogene to early Miocene) localities (Akgün et al., 2007). According to Beaudouin et al. (2005), such differences may be attributable to

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ACCEPTED MANUSCRIPT changing fluvial input of pollen to the depositional site. Alternatively, it may reflect slightly warmer lowland conditions for the Kultak palynoflora.

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From the same area (Ören region), Kayseri-Özer et al. (2014a) reported two further palynofloras, derived from lacustrine depositional settings, which have some

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similarities with the floras from the Yatağan Basin. The pollen floras from the Hüsamlar Member originate from alternating layers of marls and lignite; from the

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latter, two mammal fossils were recovered and a tentative age of MN4b–5 has been proposed (Kayseri-Özer et al., 2014a). The first mammal recovered, was

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Gomphotherium angustidens Cuvier, 1817, with primitive tooth morphology, possibly suggesting a late Burdigalian age; the other, Prosanthorinus sp., a European

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rhinoceros with similarities to congeneric specimens from France. The Karacaağaç

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coal mine palynoflora is very similar to the Yatağan Eskihisar and Tınaz lignite mine floras (abundant Pinus, Cedrus, Quercus spp.). Abundant palm pollen, a riparian

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indicator, is not recorded from the Eskihisar and Tınaz lignite mine floras. Finally, the

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Hüsamlar flora reported by Kayseri-Özer et al. (2014a) is also similar to the Yatağan Eskihisar and Tınaz lignite mine floras, with the exeption of the high proportion of Taxodioideae pollen in the Hüsamlar flora.

4.4. Biogeographic relationships of the palynofloras of the Tınaz lignite mine section In general, pollen and spores are less useful for inferring biogeographic relationships than macrofossils because pollen and spores usually are not informative at the species level. Exceptions are, for instance, the extinct genus Tricolpopollenites wackersdorfensis (Fabaceae) that can be linked to the fossil plant Podocarpium 78

ACCEPTED MANUSCRIPT podocarpus. However, in many cases, generic assignments of pollen and spores can be made, in particular when assisted by SEM observation. In this case, biogeographic

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affinities can be established at a coarser level. Various biogeographic patterns were identified for the Tınaz section palynoflora (Table S 3). Most common are taxa with a

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modern Cosmopolitan distribution. Included here are both woody and herbaceous angiosperms. Taxa with a northern hemispheric distribution are also common; this

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group comprises chiefly temperate tree genera. Classical modern relictual and disjunct distributions of formely widespread taxa are evident for example in plants today

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confined to East Asia (Cathaya, Eucommia), North America (Decodon), North America-East Asia (Carya, Tsuga), and western Eurasia-North America-East Asia

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(Fagus, Liquidambar, and Ostrya).

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Few taxa display biogeographic relationships with the Palaeotropics/Africa. These are Euphorbiaceae aff. Epiprinae, Sapotaceae aff. Sapoteae, and Oleaceae, gen. indet. 2

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with morphological affinities with Linociera and Noronhia. The presence of Picrasma

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pollen (Simaroubaceae) in the lower lignite layers of pollen zone 1 is remarkable, as the botanical affinities of the enigmatic flower Chaneya, present in early to middle Miocene deposits of Turkey and Central Europe, have recently been shown to be with Picrasma (Wang and Manchester, 2000).

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ACCEPTED MANUSCRIPT 5. Summary

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In this study, we document pollen and spores from a 75 m thick section of the Tınaz lignite mine, Tınaz Subbasin, Yatağan Basin, Muğla. We identified three informal

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pollen zones, of which the lower zones 1 and 2 are dominated by arboreal pollen. A transitional zone just above pollen zone 2 records increased herbaceous pollen (30–50

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%) with substantial amounts of Amaranthaceae and the presence of Nitraria pollen. The upper zone 3 is separated from the transitional zone by 35 m of sediment devoid

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of palynomorphs, and is characterised by high amounts of herbaceous pollen (ca 80 %) along with still diverse but less abundant arboreal pollen. The pollen spectra of the

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transitional zone and pollen zone 3 reflect either more open local vegetation or a

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change in the water table of the lake resulting in temporarily exposed lake shores. The Yenieskihisar mammal site (MN7/8) from the adjacent Eskihisar sub-basin clearly is

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younger than pollen zone 3 of the Tınaz sub-basin. Although the published palynoflora

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of Yenieskihisar is dominated by arboreal taxa and is indicative of a complex landscape comprising forest and more open vegetation, open grass-dominated landscapes have been inferred from mammal fossils. Comparisons of the Yatağan Basin palynofloras with other published southwestern Turkish palynofloras of middle and late Miocene age illustrate that based on palynological evidence alone it is difficult to safely infer ages of plant-bearing sediments. Biogeographic links of the Tınaz lignite mine palynoflora are mainly with temperate regions of the Northern Hemisphere. A few taxa show African biogeographic links,

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ACCEPTED MANUSCRIPT and a single taxon, Nitraria, belongs to a lineage that has a modern diversity centre in

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Central and northeastern Asia.

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Acknowledgements

Thanks are due to the staff of the Tınaz lignite mine for facilitating field work. Tuncay

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Güner, Istanbul, kindly organized the field work in the Yatağan Basin. We thank two anonymous reviewers for their valuable comments and Steve McLoughlin for proof

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reading the final version of this manuscript. This study was funded by the Swedish Research Council, grant to TD (project number 2012-4378). FG received funding from

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the Austrian Science Foundation (FWF; project number P24427-B25). A Riksmusei

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CE

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Vänner travel grant is acknowledged.

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ACCEPTED MANUSCRIPT Figure Captions

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Figure 1. Geographical and regional geological setting of early to middle Miocene vertebrate localities (VL) and lignite mines (LIM) in the Yatağan Basin and Ören

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basin area. 1. Map showing the geographical position of the Yatağan Basin and Ören basin (2). 2. Regional geological map of the Yatağan Basin and Ören basin (simplified

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map based on Atalay, 1980; Becker-Platen, 1970; Kayseri, 2010; Inaner et al., 2008). Late early to middle Miocene LIM and VL: (A) Karacaağaç LIM, (B) Hüsamlar LIM

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(MN4e–5), (C) Kultak VL (MN5–6) (Kayseri, 2010, Kayseri et al., 2014a). Middle Miocene LIM and VL with lithostratigraphic and pollen correlation: (D) Eskihisar

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LIM section, (E) Yenieskihisar VL, note that fluviatile mammal-bearing sediments are

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younger than sediments exposed at the Tınaz lignite mine. Lacustrine sediments below the mammal horizon at Yenieskihisar correspond to the sediments comprising pollen

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zone 3 at Tınaz. (F) Bağyaka LIM (Inaner et al., 2008), (G) Tınaz LIM (this study),

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(H) Çatakbağyaka VL (MN7/8; this study), (I) Sekköy LIM (Kayseri, 2010). References for age inferences: (a) this study (S. Mayda, unpublished data), (b) NOW (Fortelius, 2016), (c) Yavuz-Isik et al. (2011), (d) Jimenez-Moreno (2005), (e) Biltekin et al., (2014), (F) Kayseri (2010) and Kayseri et al. (2014a), (g) Wessels, 2009.

Figure 2. Lithostratigraphic units of the Eskihisar Formation from which sediment samples were taken. The sampled section includes the uppermost part of the Turgut Member, which is here considered to be early middle Miocene (Langhian to Serravallian) in age. 103

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Figure 3. LM pollen count diagram of the Tınaz mine section showing percentages of

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taxa. Abundance in %. + = rare (<2.5 %); x = Botryococcus colonies present, xx = B. colonies frequently encountered; * taxon comprises several genera or infrageneric

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groups discernible in SEM but not in LM. Hence these genera are not considered in the pollen diagram. L. = Laevigatosporites; T. = Tricolporopollenites; LO = Lonicera

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sp.; LIN = Linnaeoideae ; S = small; L = large. N = 400 pollen per sample. Sample

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numbers are indicated next to the simplified stratigraphic section.

Plate I.

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LM overview (1–3, 5–17, 19, 21, 24–29), SEM overview (22) and SEM detail (4, 18,

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20, 23) micrographs of Botryococcaceae, Zygnemataceae, algal cysts, Osmundaceae, Pteridaceae, spores of uncertain affinities, Cupressaceae, Pinaceae, Poaceae,

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lignite mine.

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Euphorbiaceae, Linaceae, and Betulaceae palynomorphs encountered in the Tınaz

1. Botryococcus cf. B. brauni (S153621). 2. Spirogyra sp. 1/Ovoidites lanceolatus (S153605). 3–4. Algal cyst indet. 1 (S153610), (4) detail. 5. Algal cyst indet. 2/Sigmopollis pseudosetarius (S153598). 6. Osmunda sp./Baculatisporites major (Raatz) Krutzsch, EV (S153621). 7. Osmunda sp./Baculatisporites primarius (Wolff) P. W. Thomson et Pflug, PV (S153621). 8–9. Pteris sp., (8) PV (S153595), (9) PV (S153604). 10. Monolete spore fam. indet./Laevigatosporites haardti (S153621), EV. 11. Cupressaceae gen. indet. 1 “non papillate”/Inaperturopollenites dubius (S153606). 12. Cupressaceae gen. indet. 2 “papillate”/Inaperturopollenites concedipites 104

ACCEPTED MANUSCRIPT (S153621), PV. 13. Cupressaceae gen. indet. 3 “papillate”/Sequoiapollenites gracilis (S153604), EV. 14. Cedrus sp., (S153596), EV. 15. Pinus subgenus Pinus sp.

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(S153595), DV. 16. Pinus subgenus Strobus sp. (S153595), EV. 17–18. Poaceae gen. indet. 1 (S153606), (17) DV, (18) exine detail. 19–20. Poaceae gen. indet. 2

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(S153621), (19) DV, (20) exine detail. 21–23. Euphorbiaceae gen. indet. (S153621), (21) top EV, bottom PV, (22) EV, (23) exine detail. 24. Linum sp. (S153621), PV. 25.

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Alnus sp./Alnipollenites verus (S153621), PV. 26. Betula sp./Trivestibulopollenites betuloides (S153621), PV, atrium visible. 27. Carpinus sp./Carpinipites carpinoides

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(S153621), PV. 28. Corylus sp. (S153621), PV. 29. Ostrya sp. (S153605), PV. EV (equatorial view), PV (polar view), DV (distal view). Scale bar – 10 µm (1–3, 5–17,

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19, 21–22, 24–29), 1µm (4, 18, 20, 23).

Plate II.

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LM overview (1, 4, 7, 10), SEM overview (2, 5, 8, 11) and detail (3, 6, 9, 12)

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micrographs of zygospores or aplanospores of Zygnemataceae and fungal uredospore. 1–3. 10–12 Puccinia sp. (S153610). 4–6. Spirogyra sp. 2/Ovoidites microfoveolatus (S153605). 7–9. Spirogyra sp. 3/ Ovoidites pococki (S153601). 10–12. Spirogyra sp. 4/ Ovoidites raatzi (S153601). Scale bars – 10 µm (1, 2, 4, 5, 7, 8, 10, 11), 1 µm (3, 6, 9, 12).

Plate III. LM overview (1, 4, 7, 10), SEM overview (2, 5, 8, 11) and detail (3, 6, 9, 12) micrographs of dispersed Sphagnaceae, Pteridaceae and spores of uncertain 105

ACCEPTED MANUSCRIPT identification. 1–3. Sphagnum sp./Stereisporites stereoides (S153621), (1) PV, (2–3) PRV. 4–6. Pteridaceae gen. indet./ Cingulatisporites macrospeciosus (S153602), (4)

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PV, (5) top EV, bottom PRV, (6) proximal detail . 7–9. Trilete spore fam. indet. 1/cf. Convolutispora ampla (S153621), (7) PV, (8–9) DV. 10–12. Trilete spore fam. indet.

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2/cf. Stereisporites (Stereigranisporis) granulus (S153599), (10) PV, (11–12) DV. EV (equatorial view), PV (polar view), DV (distal view), PRV (proximal view). Scale bars

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– 10 µm (1, 2, 4, 5, 7, 8, 10, 11), 1 µm (3, 6, 9, 12).

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Plate IV.

LM overview (1, 4, 7, 10, 14), SEM overview (2, 5, 8, 11) and detail (3, 6, 9, 12, 13,

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15) micrographs of dispersed Ephedraceae and Cathaya pollen. 1–3. Ephedra sp.

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1/Ephedripites (Ephedripites) regularis (S153610), EV. 4–6. Ephedra sp. 2/Ephedripites (Distachyapites) tertiarius (S153621), EV. 7–9. Ephedra sp.

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3/Ephedripites (Distachyapites) lusaticus (S153608), EV. 10–13. Cathaya sp.

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1/Cathayapollis scheuringii (S153604), (10) PV, (11) DV, (12) saccus detail, (13) corpus detail. 14–15. Cathaya sp. 2/Cathayapollis vancampoae (S153595), (14) PV, (15) saccus detail. EV (equatorial view), PV (polar view), DV (distal view). Scale bars – 10 µm (1, 2, 4, 5, 7, 8, 10, 11, 13), 1 µm (3, 6, 9, 12, 13, 15).

Plate V. LM overview (1, 5, 8, 11, 14), SEM overview (2, 6, 9, 12, 15, 16) and detail (3, 4, 7, 10, 13) micrographs of dispersed Pinaceae, Poaceae and Typhaceae pollen. 1–4. Tsuga sp./Zonalapollenites verrucatus (S153621), (1) PV, (2) DV, (3) saccus detail, (4) 106

ACCEPTED MANUSCRIPT leptoma detail. 5–7. Poaceae gen. indet. 3 (S153621), EV. 8–10. Sparganium sp. (S153598), (9) EV. 11–16. Typha sp. 11–13. Monad (S153621). 14–16. Typha sp.

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tetrad/Tetradomonoporites typhoides Krutzsch (S153621). EV (equatorial view), PV (polar view), DV (distal view). Scale bars – 10 µm (1, 2, 4, 5, 7, 8, 10, 11), 1 µm (3, 6,

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9, 12).

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Plate VI.

LM overview (1, 4, 7, 10, 13), SEM overview (2, 5, 8, 11, 14) and detail (3, 6, 9, 12,

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15) micrographs of dispersed Zingiberaceae, Ranunculaceae, Altingiaceae, Salicaceae, and Fabaceae pollen. 1–3. Spirematospermum sp. (S153621). 4–6. Ranunculaceae gen.

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indet. (S153606), EV. 7–9. Liquidambar sp./Peripollenites stigmosous (S153595). 10–

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12. Salix sp. (S153599), EV. 13–15. Podocarpium podocarpum/ Tricolporopollenites wackersdorfensis (S153595), EV. EV (equatorial view), PV (polar view). Scale bars –

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10 µm (1, 2, 4, 5, 7, 8, 10, 11, 13, 14), 1 µm (3, 6, 9, 12, 15).

Plate VII.

LM overview (1, 4, 7, 10, 13), SEM overview (2, 5, 8, 11, 14) and detail (3, 6, 9, 12, 15) micrographs of dispersed Fagaceae and Cannabaceae pollen. 1–3. Quercus sp. 1 (Quercus Group Cerris) (S153621), EV. 4–6. Quercus sp. 2 (Quercus Group Ilex) (S153621), EV. 7–9. Quercus sp. 3 (Quercus Group Ilex) (S153621), EV. 10–12. Trigonobalanopsis sp. (S153621), EV. 13–15. Celtis vel Pteroceltis sp. (S153621),

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13, 14,), 1 µm (3, 6, 9, 12, 15).

Plate VIII.

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LM overview (1, 3–6, 8–15, 17, 19, 21–28) and SEM micrographs (2, 7, 16, 18, 20, 29) of dispersed Fagaceae, Juglandaceae, Myricaceae, Ulmaceae, Geraniaceae,

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PV, (2) exine detail. 3. Engelhardioideae gen. indet. 1/Momipites quietus (S153621),

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PV. 4. Engelhardioideae gen. indet. 2/Momipites punctatus (S153621), PV. 5. Carya sp./Caryapollenites simplex (S153621), PV. 6–7. Morella vel. Myrica sp. (S153621),

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(6) PV, (7) exine detail. 8. Cedrelospermum sp. (S153604), PV. 9. Ulmus sp.

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(S153621), PV. 10. Zelkova sp. (S153621), PV. 11. Erodium sp. (S153610), PV. 12. Tilia sp./Intratriporopollenites instructus (S153621), PV. 13. Amaranthaceae gen. indet. 1 (S153621). 14. Amaranthaceae gen. indet. 2 (S153621). 15–16. Caryophyllaceae gen. indet. (S153621), (16) exine detail. 17–18. Polygonum sp. (S153517), (17) EV, (18) exine detail. 19–20. Rumex sp. (S153617), (19) EV, (20) exine detail. 21. Eucommia sp./Tricolpopollenites parmularis (S153621), EV. 22. Asteroideae type 1 (S153621), EV. 23. Asteroideae type 2 (S153621), left PV, right EV. 24. Asteroideae type 3 (S153621), PV. 25. Asteroideae type 4 (S153617). 26. Cichoroideae type 1 (S153621), PV. 27. Cichoroideae type 2 (S153621), PV. 28–29. 108

ACCEPTED MANUSCRIPT Linnaeoideae gen. indet. (S153604), (28) PV, (29) exine detail. EV (equatorial view), PV (polar view). Scale bar – 10 µm (1, 3–6, 8–15, 17, 19, 21–28), 1µm (2, 7, 16, 18,

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Plate IX.

LM overview (1, 4, 7, 10, 13), SEM overview (2, 5, 8, 11, 14) and detail (3, 6, 9, 12,

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15) micrographs of dispersed Lythraceae, Malvaceae, and Sapindales pollen. 1–6. Decodon sp. 1–3. (S153595), EV. 4–6. Pollen aggregation (S153595). 7–9.

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Malvoideae gen. indet. (S153617). 10–12. Sapindales fam. et gen indet. 1 (S153610),

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EV. 13–15. Nitraria sp. (S153610), EV. EV (equatorial view). Scale bars – 10 µm (1,

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Plate X.

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2, 4, 5, 7, 8, 10, 11, 13, 14), 1 µm (3, 6, 9, 12, 15).

LM overview (1, 4, 7, 10, 13), SEM overview (2, 5, 8, 11, 14) and mesocolpium detail (3, 6, 9, 12, 15) micrographs of dispersed Sapindaceae pollen. 1–6. Acer morphotype1. 1–3. (S153602), EV. 4–6 (S153621), EV. 7–9. Acer morphotype 2 (S153617), EV. 10–15. Acer morphotype 3. 10–12. (S153621), EV. (S153621), EV. 13–15. (S153621), EV. EV (equatorial view). Scale bars – 10 µm (1, 2, 4, 5, 7, 8, 10, 11, 13, 14), 1 µm (3, 6, 9, 12, 15).

Plate XI. 109

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(polar view). Scale bars – 10 µm (1, 2, 4, 5, 7, 8, 10, 11, 13, 14), 1 µm (3, 6, 9, 12, 15). Plate XII.

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LM overview (1, 4, 7, 10, 13), SEM overview (2, 5, 8, 11, 14) and detail (3, 6, 9, 12, 15) micrographs of dispersed Oleaceae, and Convolvulaceae pollen. 1–3. Oleaceae

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type 2 (S153621), EV. 4–6. Oleaceae type 3 (S153621), (4 top) EV, (4 bottom, 5) PV.

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7–9. Oleaceae type 4 (S153621), (7 top) EV, (7 middle, bottom; 8) PV. 10–12. Oleaceae type 5 (S153621), EV. 13–15. Oleaceae type 6 aff. Ligustrum/Syringia

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(S153621), EV. EV (equatorial view), PV (polar view). Scale bars – 10 µm (1, 2, 4, 5,

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7, 8, 10, 11, 13, 14), 1 µm (3, 6, 9, 12, 15).

Plate XIII.

LM overview (1, 4, 8, 11, 14), SEM overview (2, 5, 9, 12, 15) and sculpture detail (3, 6, 7, 10, 13, 16) micrographs of dispersed Convolvulaceae, Aquifoliaceae, Campanulaceae, Caprifolioideae, and Dipsacoideae pollen. 1–3. Convolvulus sp. (S153621), EV. 4–7. Ilex sp. (S153604), (4–5) PV, (6) mesocolpium, (7) colpus membrane. 8–10. Campanuloideae gen. indet. (S153621), PV. 11–13. Lonicera sp./Lonicerapollis grandis (S153608). 14–16. Dipsacus vel Cephalaria sp. 1 110

ACCEPTED MANUSCRIPT (S153621), PV. EV (equatorial view), PV (polar view). Scale bars – 10 µm (1, 2, 4, 5,

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8, 9, 11, 12, 14, 15), 1 µm (3, 6, 7, 10, 13, 16).

Plate XIV.

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9. Scabiosa sp. (S153617), (1 right) PV, (1 bottom, 2) EV. 10–13. Apiaceae type 1 (S153598), EV. 14–17. Apiaceae type 2 (S153610), EV. EV (equatorial view), PV

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(polar view). Scale bars – 10 µm (1–5, 7, 8, 10, 11, 14, 15), 1µm (6, 9, 12, 13, 16, 17).

Plate XV.

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LM overview (1, 5, 9, 12, 16), SEM overview (2, 6, 10, 13, 17), detail (3, 7, 15, 18),

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and aperture detail (4, 7, 11, 14) micrographs of dispersed Apiaceae and Dicotyledone pollen of uncertain affinity pollen. 1–4. Apiaceae type 3 (S153604), (1–2) EV, (3) apex, (4) colpus and mesocolpium detail. Saniculoideae gen indet (S153621), (5–6) EV, (7) apex, (8) colpus and mesocolpium detail. 9–11. Pollen type 1 (S153621), EV. 12–15. Pollen type 2 (S153604), (12–13) PV. 16–18. Pollen type 3 (S153621), PV. EV (equatorial view), PV (polar view). Scale bars – 10 µm (1, 2, 5, 6, 9, 10, 12, 13, 16, 17), 1µm (3, 4, 7, 8, 11, 14, 15, 18).

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Fig. 1 112

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Plate I

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Plate II

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Plate III

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Plate IV

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Plate V

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Plate VI

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Plate VII

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Plate VIII

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Plate IX

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Plate X

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Plate XI

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Plate XII

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Plate XIII

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Plate XIV

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Plate XV

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ACCEPTED MANUSCRIPT Highlights

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- Middle Miocene palynofloras of the Tınaz lignite mine, Yatağan Basin, SW Anatolia, are investigated - High taxonomic resolution is achieved by a combined LM and SEM approach - Mammals co-occurring with palynofloras provide age constraints - Palaeogeographic relationships of the pollen floras are mainly northern hemispheric

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