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Palaeogeography, Palaeoclimatology, Palaeoecology 530 (2019) 236–248

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Palaeogeography, Palaeoclimatology, Palaeoecology

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Early and of : Evidence from fossils, T dispersed , and petrified wood ⁎ Thomas Denka, , H. Tuncay Günera,b, Johannes M. Bouchala a Swedish Museum of Natural History, Department of Palaeobiology, Box 50007, 10405 Stockholm, Sweden b University Cerrahpaşa, Faculty of Forestry, Department of Botany, 34473 Bahçeköy, Istanbul, Turkey

ARTICLE INFO ABSTRACT

Keywords: The was a period of major palaeogeographic reorganization in the eastern Mediterranean region, Miocene climate during which time the Anatolian Plateau became subaerial and several intracontinental basins intermittently became connected to the Paratethys and Mediterranean seas. In this paper, we analyse early Miocene CLAMP and climate using leaf records, palynological assemblages, and fossil wood at 36 localities from western and Köppen signatures central Turkey, most of which have precise age control based on radiometric dating and faunal ages. Using the leaf flora of Güvem (Beş Konak, Keseköy), Climate Leaf-Analysis Multivariate Program (CLAMP) Burdigalian analyses and Köppen signatures were employed to infer a palaeoclimate typical of modern laurel forest regions. Based on the palynological records, abundance of various pollen-taxa was used as a measure of openness of vegetation and regional presence of major taxa. Most pollen floras are dominated by tree pollen (ranging from 85 to 98%) and indicated widespread afforestation. In the pollen diagrams, shifts in dominance from swamp forest elements (Taxodioideae) to well-drained (Pinaceae) indicate changes in lake levels or phases of basin development. Such shifts may have been associated with the development of more xeric forest vegetation. Wood anatomical features such as false tree rings further may indicate seasonal climate. Pollen diagrams and macrofossils reflect zonal and azonal broadleaf and needleleaf forest and extrazonal openvege- tation. The latter occurred in areas with shallow soils on volcanic rocks or (e.g. cycads, Dracaena), or coastal areas (herb dominance). Taxonomic composition and biogeographic affinities suggest laurel forest asa major forest biome on well-drained soils and ecotones between laurel forest and broadleaf deciduous forest biomes. A comparison with younger floras shows that these are neither more diverse nor more warmth-loving despite an increase in global temperature (Mid-Miocene Climatic Optimum) suggesting bottlenecks during previous () cooler times for warmth-loving taxa.

1. Introduction This exchange included African immigrants such as the Proboscidae (Koufos et al., 2003). While large parts of Minor were submerged during the There is conflicting evidence for the type of vegetation andpa- (Rögl, 1998, 1999), western and parts of central Turkey became sub- laeoenvironments that existed in during the early Miocene. aerial during the Oligocene (Rögl, 1999; Popov et al., 2004). During the Strömberg et al. (2007) analysed phytoliths from three early Miocene early Miocene, western Anatolia remained subaerial (e.g. Alcicek, 2010; localities in the Galatian Volcanic Province (GVP) and inferred rela- Ersoy et al., 2014), while emergent areas in central and east Turkey tively open vegetation such as savanna or open woodland. In contrast, were repeatedly transgressed by shallow seas (Paicheler et al., 1978; palynological data from the GVP suggest a flora dominated by and Popov et al., 2004; Poisson et al., 2016). During the later Burdigalian a mixed mesophytic forest (Yavuz-Işık, 2008; Yavuz-Işık and Demirci, (19–18 Ma), eastern Turkey emerged and the Tethys Seaway closed, 2009), and silicified woods from the GVP suggest diverse forest vege- resulting in a terrestrial connection between Anatolia and -Arabia tation during the early Miocene with some aridity (Akkemik et al., and Eurasia. The formation of a temporary land bridge (Gomphotherium 2016, 2017, 2018a, 2018b). land bridge) enabled faunal exchange at the base of mammal zone MN4 The objectives of this present paper are (i) to assess palaeoenvir- between Africa and Eurasia (Rögl, 1999; Harzhauser and Piller, 2007). onmental signal in 36 pollen, leaf and wood floras, (ii) to assess the

⁎ Corresponding author. E-mail address: [email protected] (T. Denk). https://doi.org/10.1016/j.palaeo.2019.05.042 Received 28 February 2019; Received in revised form 28 May 2019; Accepted 28 May 2019 Available online 31 May 2019 0031-0182/ © 2019 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/BY-NC-ND/4.0/). .Dn,e al. et Denk, T. Table 1 Early Miocene fossil localities (1 to 36; see Fig. 1) from Turkey considered in the present study. Abbreviations: (P) pollen, (L) leaf flora, (W) petrified wood, (PH) phytoliths, M Zone = planktonic foraminiferal zone, MN Zone = European mammal zone, NN Zone = nannofossil zone.

No. Locality Abbrev. Province Element and reference Age/ M Zone/ MN Zone/ NN Age constraint and reference Zone

1 Haymana HAY Ankara (W) Akkemik et al., 2018b Late Chattian/ 24.7 ± 1.9 Ma Radiometric (Ar/Ar): Atıcı et al., 2014 early Aquitanian 2 Sivas Basin SIV1 Sivas (P) Akkiraz et al., 2018 Aquitanian Unspecified Aquitanian Molluscs: Akkiraz et al., 2018 3 Muş Basin MUS Muş (P) Sancay et al., 2006 Aquitanian M1 (EM1–EM2) : Sancay et al., 2006 4 Kalkim-Gönen (Bengiler KAL Çanakkale (P) Üçbaş Durak and Akkiraz, 2016 Aquitanian 22.26 ± 0.06 Ma Radiometric (Ar/Ar): Üçbaş Durak and Akkiraz, 2016 section) 5 Tunçbilek TUB Kütahya (L) Mädler and Steffens, 1979; Aquitanian 22.12 ± 0.02 Ma to 21.56 ± Radiometric (Ar/Ar): Helvacı et al., 2017 (P) Akkiraz et al., 2012 0.04 Ma 6 Seytömer SEY Kütahya (P) Nakoman, 1968; (P) Yavuz-Işık, 2007; Aquitanian 22.12 ± 0.02 Ma to 21.56 ± Radiometric (Ar/Ar): Helvacı et al., 2017 (P) Akkiraz et al., 2012 0.04 Ma 7 Harami HAR Konya (P) Karayiğit et al., 1999; Aquitanian 22.268 to 21.936 Ma; Magentostratigraphy: Krijgsman et al., 1996 (P) Biltekin, 2018 MN1(–2) Vertebrates: de Bruijn et al., 1992; van den Hoek Ostende, 2001; Joniak et al., 2018 8 Kargı 2 KRG Çorum (P) Yavuz-Işık et al., 2011 Aquitanian MN1 Vertebrates: Saraç, 2003; NOW Community, 2019 9 Kılçak KCK Çankırı (P) Yavuz and Demırer, 2018 Aquitanian MN1–2 (–3) Vertebrates: de Bruijn and Saraç, 1992; de Bruijn et al., 1993; de Bruijn and von Koenigswald, 1994; Ünay, 1994; van den Hoek Ostende, 1992, 1995a, 1995b, 2001 10 Balya lignite mine BAL Balıkesir (L) Engelhardt, 1903; Denk et al., 2017b Aquitanian to 22.97 ± 0.23 to 18.72 ± 0.17 Radiometric (U-Pb): Aslan et al., 2017 Burdigalian Ma 11 Gördes Basin GRD Manisa (P) Akgün and Akyol, 1987 Aquitanian to 21.7 ± 0.04 to 16.4 ± 0.01 Radiometric (Ar/Ar): Purvis et al., 2005 Burdigalian Ma; MN3 Vertebrates: Peláez-Campomanes et al., 2018 237 12 Çan CAN Çanakkale (L) Mädler and Steffens, 1979; (P) Ediger, 1990; (P) Bozcu Aquitanian to 21.5 to 16.2 Ma for Ezine Radiometric (K/Ar): Borsi et al., 1972; et al., 2015 Burdigalian region Ercan et al., 1985; Yılmaz, 1990 13 Soma SOM Manisa (L) Mädler and Steffens, 1979; (P) Akgün et al., 1986; (P) Aquitanian to 20.72 ± 0.10 to 18.76 ± 0.05 Radiometric (Ar/Ar): Ersoy et al., 2014 Takahashi and Jux, 1991; (L, P) Gemici et al., 1991; (P) Burdigalian Ma; Vertebrates: Kaya et al., 2007 Akgün, 1993; (L) Erdei et al., 2010; (L) Güner and Denk, MN3 2012; (L) Denk et al., 2014, 2015, 2017b 14 Bigadiç Basin BIG Balıkesir (P) Akyol and Akgün, 1990 Aquitanian to 20.6 ± 0.7 to 17.8 ± 0.4 Ma Radiometric (Ar/Ar, K/Ar): Erkül et al., 2005 Burdigalian Palaeogeography, Palaeoclimatology,Palaeoecology530(2019)236–248 15 Güvem (including the sites GUV Ankara (P) This study; (L) Kasaplıgil, 1977; (L) Mädler and Steffens, Burdigalian 19.7 ± 0.6 to 17.9 ± 0.5 Ma; Radiometric (K/Ar): Wilson et al., 1997 Beş Konak and Keseköy) 1979; (L) Paicheler and Blanc, 1981; (PH) Strömberg et al., MN3 Vertebrates: de Bruijn et al., 1992; 2007; (P) Yavuz-Işık, 2008; (P) Yavuz-Işık et al., 2011; (L) van den Hoek Ostende, 2001; Wessels, 2009 Güner and Denk, 2012; (L) Denk et al., 2014, 2015, 2017a, b 16 Kocacay KOC Izmir (P) Kayseri-Özer et al., 2014 Burdigalian MN3–4 Vertebrates: Kaya et al., 2007 17 Kıbrısçık-Kuzca KUZ (W) Acarca Bayam et al., 2018 Burdigalian 20 to 18 Ma Radiometric (K/Ar): Toprak et al., 1996 18 Beypazarı- Aşağıgüney AGU Bolu (W) Acarca Bayam et al., 2018 Burdigalian 20 to 18 Ma Radiometric (K/Ar): Toprak et al., 1996 19 Beypazarı-Kıraluç KIR Bolu (W) Acarca Bayam et al., 2018 Burdigalian 20 to 18 Ma Radiometric (K/Ar): Toprak et al., 1996 20 Beypazarı-Mençeler MEN Ankara (W) Acarca Bayam et al., 2018 Burdigalian 20 to 18 Ma Radiometric (K/Ar): Toprak et al., 1996 21 İnözü Deresi kuzey taraf INL Ankara (W) Acarca Bayam et al., 2018 Burdigalian 20 to 18 Ma Radiometric (K/Ar): Toprak et al., 1996 22 İnözü Deresi güney taraf INO Ankara (W) Acarca Bayam et al., 2018 Burdigalian 20 to 18 Ma Radiometric (K/Ar): Toprak et al., 1996 23 Beypazarı-Karaşar Köyü KAR Ankara (W) Acarca Bayam et al., 2018 Burdigalian 20 to 18 Ma Radiometric (K/Ar): Toprak et al., 1996 24 Çamlıdere-Elmalı Köyü ELM Ankara (W) Acarca Bayam, 2018 Burdigalian 19.7 ± 0.6 to 17.9 ± 0.5 Ma Radiometric (K/Ar): Tankut et al., 1995 25 Kızılcahamam- Soğuksu SOG Ankara (W) Acarca Bayam et al., 2018 Burdigalian 19.7 ± 0.6 to 17.9 ± 0.5 Ma Radiometric (K/Ar): Tankut et al., 1995 26 Çamlıdere-Buğralar Köyü BUG Ankara (W) Acarca Bayam et al., 2018 Burdigalian 19.7 ± 0.6 to 17.9 ± 0.5 Ma Radiometric (K/Ar): Tankut et al., 1995 27 Çamlıdere-Pelitçik PEL Ankara (P) Yavuz-Işık and Demirci, 2009; Burdigalian 18.2 ± 0.5 to 18 ± 0.5 Ma; Radiometric (K/Ar): Wilson et al., 1997 (W) Akkemik et al., 2009 MN3–6 Vertebrates: Saraç, 2003 28 Seben-Kozyaka KOZ Bolu (W) Akkemik et al., 2016 Burdigalian 18.2 ± 0.8 to 17.4 ± 0.8 Ma Radiometric (K/Ar): Akkemik et al., 2016 29 Seben-Hoçaş HOC Bolu (W) Akkemik et al., 2016 Burdigalian 18.2 ± 0.8 to 17.4 ± 0.8 Ma Radiometric (K/Ar): Akkemik et al., 2016 30 Güdül GUD Ankara (W) Akkemik et al., 2017 Burdigalian 18.8 ± 0.6 to 16.9 ± 0.5 Ma Radiometric (K/Ar): Wilson et al., 1997 31 Sivas Basin SIV2 Sivas (P) Ocakoğlu et al., 2018 Burdigalian NN3 (=18.92 to 17.97 Ma) Nannofossils: Ocakoğlu et al., 2018 (continued on next page) T. Denk, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 530 (2019) 236–248

palaeoclimate for the GVP using the Climate Leaf-Analysis Multivariate Program (CLAMP) and Köppen signatures, and (iii) to use all floras to reconstruct major biomes in Anatolia during the early Miocene. We infer major biomes during the early Miocene of Anatolia using biome definitions in Prentice et al. (1992), Schroeder (1998), Olson et al. (2001), Woodward et al. (2004), and Körner (2013).

Sarı et al., 2015 2. Material and methods

2.1. Stratigraphic framework Baş, 1986 NOW Community, 2019 In this study, we considered 36 plant fossil localities from lower Göktaş, 2013 ;

Koral et al., 2009 ; Miocene deposits of Anatolia to infer palaeoenvironments and palaeo- climate. In Table 1 we list the plant fossil localities and locality ab- Saraç, 2003 ;

Uchman et al., 2007 breviations along with information on the type of fossil assemblage, age, age constraints, and the appertaining references. The robust stra- tigraphic framework recently developed for many early Miocene plant

Age constraint and reference Radiometric (K/Ar): Karacık et al., 2013 Akdeniz and Konak, 1979 ; Ercan et al., 1995 ; assemblages of Turkey forms the basis for our assessment of environ- mental and climatic signals in these assemblages. Comparative studies of early Miocene floras in Turkey have been problematical because age estimates solely relying on palynological or plant macrofossil data often resulted in erroneous ages. For example, the flora of Güvem/Ankara was initially dated as basedon its leaf fossils (Kasaplıgil, 1977). Subsequent studies on vertebrate fossils and volcanic rocks from this area revealed an unequivocal early Miocene (Neogene European mammal zone 3, MN3) age (e.g. de Bruijn

Age/ M Zone/ MN Zone/Zone NN Cumaovasi Formation 17.9 ± 0.6 to 13 ± 0.4 Ma et al., 1992; Wilson et al., 1997). The leaf flora from Balya/Balıkesir described by Engelhardt (1903) was considered to be of late Miocene age but was later shown to be of early Miocene (Aquitanian) age (Aslan et al., 2017). The pollen floras of Tunçbilek/Kütahya and Soma/Manisa all were considered to be of middle age (ca. 13 Ma) in the

Stage Lower Miocene Age of the overlying Lower MioceneLower Miocene Unspecified lower Miocene Unspecified lower MioceneLower Miocene Ostracods: Lower Miocene Unspecified lower Vertebrates: Miocene Unspecified lower Miocene palynological subdivision of the Turkish Neogene by Benda (1971). Subsequently, vertebrate data (Kaya et al., 2007; Soma) and radio- metric data (Ersoy et al., 2014, Soma; Helvacı et al., 2017, Tunçbilek) indicated Aquitanian to Burdigalian ages for these localities. Similarly, for the leaf and pollen floras of Şahnalı (Şahinali)/Aydın, Gördes/ Denk et al., Manisa, and Tunçbilek, early and middle Serravallian ages were sug- gested by Akgün and Akyol (1987, 1999), Gemici et al. (1993), Akgün et al. (2007) and Kayseri-Özer (2017). In contrast, vertebrate data suggest early Miocene ages for these localities (Saraç, 2003)(Table 1). Akkiraz, 2011 ;

2.2. Plant fossil assemblages

Plant macrofossils () from the Güvem area (localities Beş Konak, Keseköy; Kasaplıgil, 1977; Paicheler and Blanc, 1981) have recently been revised (Denk et al., 2017a). We used angiosperm leaf Gemici et al., 1992 ; Gemici et al., 1992 ; (P) morphotypes described in this study as palaeoclimate proxies; palaeo- Akkemik et al., 2019 Güngör et al., 2019 Kayseri-Özer et al., 2014 Biltekin, 2017 climate was inferred with Climate Leaf Analysis Multivariate Program 2017b (P) (CLAMP; Yang et al., 2011; http://clamp.ibcas.ac.cn, last access: Oc- tober 10, 2018). Further leaf macrofossils from early Miocene strata were reported from Çan/Çanakkale (Mädler and Steffens, 1979), Soma/Manisa (Mädler and Steffens, 1979; Gemici et al., 1991), Şahnalı/Aydın (Mädler and Steffens, 1979; Gemici et al., 1993), and Tavşanlı/Kütahya SAH Aydın (L, P) DURGOK Balıkesir Çanakkale (W) (W) ERM Içel (P) CMV Izmir (L, P)

Abbrev. Province Element and reference (Mädler and Steffens, 1979). For these leaf floras, major revisions are lacking and preliminary, revised taxon lists are presented (Supple- mentary Table S3). Updates are based on newly collected material since 2010 (T. Denk, H.T. Güner, unpublished material; stored at Department of Forest Botany, Istanbul University Cerrahpaşa [ISTO]) and material stored at Bundesanstalt für Geowissenschaften und Rohstoffe, Hannover (BRG). In addition, single fossil-taxa have been described from Soma (cycads, Erdei et al., 2010; Mahonia, Güner and Denk, 2012; Dracaena, , Quercus, Denk et al., 2014, 2015, 2017a) and are considered as ( continued ) well. Pollen and spores from a section at the Keseköy mammal locality No. Locality 32 Cumaovasi 3334 Ermenek Şahinali 3536 Dursunbey Gökçeada

Table 1 (village Yukarı Kise; see van den Hoek Ostende, 2001; Wessels, 2009)

238 .Dn,e al. et Denk, T.

Table 2 Abundances of selected pollen taxa from early Miocene strata of Turkey and arboreal pollen/non-arboreal pollen ratios (see Supplementary Tables S1, S2, and S2a for full data set). For full locality information for the palynological sites see Table 1. PZ = Pollen zone; predominantly subtropical and tropical taxa are in bold face; grey shading indicates and Fagales. Pollen abundances: • = 0–1%, x = 1–5%, xx = 5–10%, xxx = > 10%, + = present no abundance data available, x(x) indicates single higher abundances in the pollen diagram. 239 Palaeogeography, Palaeoclimatology,Palaeoecology530(2019)236–248

aOne sample with Asteraceae peak in Kilcak 1 section. b85/15 Typhaceae peak. cZelkova and Ulmus not differentiated. T. Denk, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 530 (2019) 236–248

Table 3 Fossil wood records in early Miocene strata of Turkey (after Acarca Bayam, 2018; Acarca Bayam et al., 2018; Akkemik et al., 2009, 2016, 2017, 2018a, 2018b, 2019; Güngör et al., 2019). First number refers to in situ samples, second number to ex situ samples. See Table 1 for full locality information. Grey shading indicates conifers and . Locality HAY KOZ HOC KUZ AGU KIR MEN INL INO KAR ELM SOG BUG PEL GUD DUR GOK Glyptostroboxylon –/6 Pinus /Pinoxylon 1/3 1/4 1/3 –/1 1/1 –/1 –/1 Cedrus 1/7 –/1 –/1 –/1 –/2 Picea –/2 –/1 1/– Juniperus 6/3 2/6 Taxodium –/1 –/1 Cupressinoxylon –/1 Sequoia /Sequoioxylon –/1 –/1 –/1 –/1 –/1 –/1 "" –/3 –/1 Palmae 17/– –/2 Laurinoxylon –/3 Liquidambar 4/– Acer –/1 –/1 –/4 –/1 –/2 Alnoxylon –/1 Carpinoxylon –/1 Ostryoxylon –/1 Fagoxylon –/1 Quercoxylon sect. Ilex –/1 1/4 1/2 1/– 3/– 1/– 1/– –/1 –/3 1/– Prunus –/1 Pistacioxylon ufuki Ü.Akkemik & I.Poole –/1 Salix –/1 1/– 2/– Salix/Populus 3/4 23/4 Ulmus –/1 –/2 Zelkova 2/– 1/– Zelkovoxylon yesimae Ü.Akkemik & I.Poole –/3 Undetermined –/18 Total no. of specimens 5 26 64 6 17 14 10 4 4 1 18 1 2 2 6 1 1

were reported by Yavuz-Işık (2008; 50 taxa) and Yavuz-Işık et al. 2.4. Arboreal/non-arboreal pollen ratios (AP/NAP) (2011). From this locality, phytoliths were investigated by Strömberg et al. (2007). For the present study, in addition, three-dimensionally We used the conservative threshold value of AP/NAP ≥3.85 as an preserved dispersed spores and pollen originating from concretions indicator of “tree-prevalent landscapes” (Favre et al., 2008), equivalent were investigated. The concretions derive from lacustrine diatomites to > 79% Arboreal pollen per sample. Favre et al. (2008) used aerial (pseudo-nodules according to Paicheler et al., 1978) occurring above photographs and directly estimated tree surface/herb surface ratios of the mammal layer of the Keseköy locality and roughly corresponding to 52 vegetation plots. They then established AP/NAP values from moss the level of sample EM3 of Strömberg et al. (2007). A single concretion polsters for each vegetation plot. Lowest AP/NAP values (< 1) corre- (hereafter concretion sample) was processed using standard methods sponded to open vegetation, while highest values (> 10) corresponded (40% HF to dissolve silica, 20% HCl to dissolve fluorspar; Halbritter to forest vegetation. A cut-off value of AP/NAP = 3.85 was calculated et al., 2018). Using the single grain technique and a combined light to safely distinguish herb and tree prevalent landscapes. Thus, AP microscopy (LM)/scanning electron microscopy (SEM) approach percentages > 79% are good predictors of forested landscapes. In ad- (Halbritter et al., 2018) about 100 taxa were recorded (Supplementary dition, we used threshold values predicting local presence (within a Figs. 1–5). SEM stubs with the pollen are stored at the Swedish Museum radius of 35 km) of European tree taxa (Lisitsyna et al., 2011). These of Natural History, Department of Palaeobiology. Besides, for several values are based on > 2000 sample sites across Europe (including a few published palynofloras (Table 2; Supplementary Tables S1, S2) bota- sites from Georgia and Armenia), and range between 10% (Pinus) and nical affinities were revised following Bouchal (2018) and 0.5% (e.g. Abies, Juniperus, Acer, Juglans). Other important taxa en- petrified wood floras were included in our comparative study aswell countered in the early Miocene of Anatolia such as Quercus, Fagus, (Table 3). Castaneoideae have values between 1.5 and 1%.

2.5. Climate-leaf analysis multivariate program (CLAMP) for inferring 2.3. Köppen signatures palaeoclimate

Köppen signatures use the main climate types in which genera CLAMP reconstructs several climate parameters using climatic thrive that are represented in a fossil plant assemblage. To determine signal encoded in leaf physiognomy (http://clamp.ibcas.ac.cn). Thirty- Köppen signatures from fossil plant assemblages, 26 Köppen–Geiger six leaf character states are recorded for all angiosperm leaves in a fossil climate types (Kottek et al., 2006; Peel et al., 2007; Rubel et al., 2017) assemblage. Using Canonical Correspondence Analysis (CANOCO, ter were mapped on distribution ranges of modern analogue taxa (1547 Braak, 1986) leaf physiognomic data from known (modern) calibration in 280 genera; Supplementary Material 1). To account for sites are compared with the leaf physiognomic data from the fossil site. ecological and climate niche evolution, infrageneric groups (sections) Because environmental data are known for the modern sites, the posi- or genera were used as modern analogues of fossil-taxa instead of tion of the fossil assemblage in leaf physiognomic space can be used to species (for details see Denk et al., 2013; Bouchal et al., 2018). reconstruct environmental data for the fossil site. Because the early Miocene plant assemblages have a strong Northern Hemispheric

240 T. Denk, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 530 (2019) 236–248 biogeographic signal with a particularly strong East Asian signal and from the depositional area, the latter possibly reflecting relatively drier because subtropical-tropical elements do occur in the assemblages, we conditions adjacent to the depositional area. used the calibration set PhysgAsia2 (with sites from and added to Northern Hemisphere sites). For comparison, we also ran CLAMP on PhysgAsia1 that has no tropical modern sites. 3.2. Leaf fossils

2.6. Definition of the Laurel Forest Biome As in the palynoflora, the macro flora of Güvem, Turkey, is richin conifers (11 fossil-taxa in at least eight genera). Fagales are represented The Laurel Forest Biome, as defined here, is characterized by with 11 genera (18 species), Rosaceae with four. Further, the broadleaf evergreen tree species with laurophyllous leaves (Schroeder, Acer is represented with at least three species (Supplementary Table 1998; Körner, 2013; temperate biome of Relyea and Ricklefs, S3). Among subtropical taxa, Smilax miohavanensis, Ilex miodipyrena, 2018). Laurophyllous leaf types are chiefly evergreen, with a coriaceous , diverse , and Simaroubaceae/Rutaceae occur. A texture, brochidodromous to eucamptodromous secondary venation, further characteristic of the leaf assemblage is the great number of large and entire or dentate margin. Although typically found in species of the lobed leaves that are difficult to assign to particular modern taxa but Lauraceae, laurophyllous leaves are also found in Magnoliaceae, show great similarities with extinct Malvaceae (Dombeyopsis, Laria) and Aquifoliaceae (Ilex), evergreen Fagaceae (e.g. , , extant Malvaceae (Sterculioideae) and the Alangiaceae (Alangium). Quercus), Rosaceae, Ericaceae, Rutaceae (Citrus) and others. In the Five other localities have yielded more or less rich leaf assemblages , , Eucryphia, and Knightia among (Fig. 1). Revised taxon lists for these localities are provided in Table 4 others, have laurophyllous leaves. A further characteristic feature of the and Supplementary Table S3. The macrofossil localities Balya, Tavşanlı, Laurel Forest Biome is the high diversity of conifers, among them many and Çan are moderately rich. Evergreen Fagaceae and other laur- relict conifers (e.g. Sequoia, Cryptomeria, Torreya and others; Schroeder, ophyllous leaf types (Ilex miodipyrena, Magnolia) dominate the assem- 1998; Körner, 2013; see Table 7). blage of Balya. The assemblages of Çan and Tavşanlı are dominated by azonal (riparian) taxa such as Alnus gaudinii and Liquidambar (Çan) and 3. Results Taxodium dubium (Tavşanlı). In contrast to these floras, the macro floras of Şahnalı and Soma are extremely diverse and the revised taxon lists 3.1. Palynological records presented here (Supplementary Table S3) do not include a large number of unidentified taxa and hence do not reflect the entire diversity of A general feature of early Miocene palynofloras in Turkey is the these floras (considering unpublished material collected by Denk and high diversity and relatively great abundance of pollen, in- Güner). The macroflora of Şahnalı is dominated by Fagales, similar to cluding and diverse Pinaceae. Cycads appear to be more the Güvem flora. Among Fagaceae, all fossil-species except Fagus are common in Aquitanian floras but potentially extend into the evergreen. Conifers are abundant but not as diverse as in Güvem Burdigalian at Soma (where they are known both from the pollen and (Torreya is recorded both in Güvem, Şahnalı, and Soma). Daphnogene macrofossil records). The same is true for palms. All floras are domi- (Lauraceae) is very common in some layers, where it co-occurs with nated by members of the Northern Hemispheric order Fagales re- Glyptostrobus. Other layers are dominated by the riparian element presented by at least fifteen genera (Table 2, Supplementary Table S1). Alnus. Dombeyopsis and Tilia are shared with the flora of Güvem. Fi- The palynological assemblage from the Keseköy concretion sample nally, the macro flora of Soma again is very diverse in Fagales taxa,but yielded > 65% pollen belonging to evergreen Fagaceae and 15% pollen evergreen Fagaceae are less dominant than in Balya, Güvem, and of conifers (Supplementary Table S2, S2a; Supplementary material Figs. Şahnalı. and Daphnogene are locally (specific layers) dominant, S1–S5). Similarly, a 40 m succession including the Keseköy mammal and so are Glyptostrobus and Pinus spp. This flora is distinct from other horizon in the lower part, shows highly uniform dominance of Fagales floras in the east Mediterranean by the common presence of Cycadaceae (Quercus, Engelhardioideae) and Pinaceae and low amounts of her- (Erdei et al., 2010) and Dracaena (Denk et al., 2014) and has a great baceous (Yavuz-Işık, 2008). number of unidentified leaf taxa that appear to be unique to thislo- A few taxa belong to families and genera extending into or pre- cality or shared with roughly coeval floras from southern France (Denk dominantly found within the (Cycas, Arecaceae, Olacaceae, T., Güner, T., unpublished data). , Simaroubaceae, Rutaceae tribus Citreae, Reevesia, The macro floras of Balya, Tavşanlı, Güvem, Soma, Çan, and Şahnalı Sapotaceae various pollen types, and Avicennia). Among these, only represent different vegetation types, among which riparian plants play Simaroubaceae and Sapotaceae and possibly Cycads are moderately a prominent role as they grew close to the depositional area (cf. Denk abundant in several floras. Herbaceous taxa generally occur inlow et al., 2015, 2017a). Azonal vegetation typically comprises members of abundances but Poaceae and Amaranthaceae reach up to 30% in early Taxodioideae and species. For example, Glyptostrobus and Aquitanian marine sediments of the Sivas Basin (Akkiraz et al., 2018) were important elements of azonal vegetation in Şahnalı, Güvem, and where they reflect coastal vegetation. Here occurs also the single record Soma. In addition, Myrica lignitum and palms might have been part of of Avicennia. this vegetation. In Tavşanlı, a Taxodium – Pinus – palm swamp forest This general picture is also seen in the arboreal/non-arboreal pollen was present, with minor contributions of Platycarya. Another taxodioid (AP/NAP) ratios and pollen threshold values of nineteen palynofloras Cupressaceae, Sequoia might have been part of azonal vegetation on from Aquitanian and Burdigalian strata (Table 2). Pollen from woody peaty soils (peat mires in Soma, Güvem; cf. Inci, 2002). A quite dif- and angiosperms dominate these assemblages with AP/ ferent azonal riparian forest is recorded from Çan, where Alnus and NAP ranging from 80/20 to 98/2 and thus indicating tree-prevalent Liquidambar were dominating elements. vegetation. Using the study of Lisitsyna et al. (2011) as a guideline for To reconstruct the regional (zonal) vegetation, plants that require local presence of particular tree taxa provides additional evidence for well-drained soils are informative. A common feature of the floras of forested vegetation during the early Miocene of Turkey. Pinus pollen is Güvem, Balya, and Şahnalı is the remarkable diversity of evergreen well above 10% in most of the investigated palynofloras suggesting Fagaceae and other laurophyllous plants (Tables 5, 6). It is noteworthy local presence of this tree genus. The same is true for a range of other that for Şahnalı the true diversity of Fagaceae most likely will be larger conifers and angiosperms including Quercus, Alnus, Engelhardioideae after revisions of both the pollen and the leaf floras. These kinds of and others (Supplementary Table S1). Values above and below the zonal forests are characteristic of the early Miocene of the eastern threshold value for Fagus suggests that this tree genus either formed Mediterranean region (Velitzelos et al., 2014; see below Discussion part of the local flora or was restricted to the hinterland further away section).

241 T. Denk, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 530 (2019) 236–248

Fig. 1. Early Miocene plant fossil localities used in this study (for full locality information for the 36 sites see Table 1). (A) Turkey showing the Galatian Volcanic Province (dark grey) and (B) Southern part of Galatian Volcanic Province.

Table 4 represented by a single wood type, assigned to Quercus sect. Ilex, and Early Miocene plant macrofossil (leaf) record of Turkey. See Supplementary thus are underrepresented in the wood record (Table 3). At some sites, Table S3 for full taxon lists. riparian elements such as Salicaceae and palms are abundantly pre- Characteristic taxon groups shared among early Miocene macro floras served as in situ stumps (e.g. Hoçaş site; Akkemik et al., 2016). Here, small-scale vegetation patterns are recorded and reflect riparian com- Typical elements of Laurel Forest Biome munities of Salix/Populus, palms, and Liquidambar versus communities Laurophyllous leaf fossil-taxa: of Quercus (evergreen type) and Juniperus reflecting well-drained stands Lauraceae Daphnogene Lauraceae gen. indet. of the hinterland and hence the regional vegetation. The same patterns Laurophyllum are known from the petrified forests of the Greek island Lesbos Magnoliaceae Magnolia (Velitzelos et al., 2014). In addition, from Lesbos, Mantzouka (2018) Myricaceae Comptonia reported lauraceous wood. Most recently, Güngör et al. (2019) reported Myrica Aquifoliaceae Ilex a diverse early Miocene wood flora from the Aegean island Gökçeada Fagaceae Quercus sect. Ilex including coastal, riparian and hinterland elements. Fagaceae aff. Eotrigonobalanus At Haymana, wood morphology (high vessel density, vessels with Fagaceae aff. Trigonobalanopsis narrow diameters) has been used as an indication of xeric growing Fagaceae gen. indet. conditions (Akkemik et al., 2018b). Further, at Beypazarı-Aşağıgüney Conifers (outside swamp forests): Pinus Sequoia (Bolu), false tree-rings were observed in Juniperus (Acarca Bayam et al., Torreya 2018). False tree-rings (intraannual density fluctuations) are common

Typical elements of Humid Broadleaf Deciduous Forest in trees growing under seasonal (De Micco et al., 2016) but are Cercidiphyllaceae Cercidiphyllum as well found in flood-prone riparian settings (e.g. Young et al., 1993). Juglandaceae Carya Malvaceae Dombeyopsis Tilia 4. Palaeoclimate inference and biogeographical affinities Rosaceae Rosa Sorbus Acer 4.1. Köppen signatures for Güvem Formation Simaroubaceae Ailanthus Ulmaceae Cedrela About 1500 modern analogues representing 66 fossil-taxa identified Ulmus Zelkova to genus level were used to infer the Köppen signature of the Güvem flora. Using the pollen flora, fully humid (Cf) and winter dry(Cw) warm temperate climates contribute to 40% of all realized climate niches of 3.3. Petrified wood all taxa in the fossil plant assemblage. Tropical A climates are re- presented by 16%. In contrast, summer dry Mediterranean climates (Cs) Gymnosperms are well represented in wood floras across the and arid climates (Bs, BW) are not well-represented (11% and 7%, re- Galatian Volcanic Province, Turkey with the exception of the earliest spectively; Fig. 2). Tropical climates are not as prominent in Köppen Aquitanian site Haymana and the Burdigalian site Beypazarı-Karaşar signatures from the macrofloras. Köyü, where no wood was found. In contrast, Fagales are

Table 5 Climate parameters reconstructed by CLAMP for the Güvem flora.

Calibration dataset Climate parameter

MAT CMMT GROWSEAS GSP MMGSP X3.WET X3.DRY (°C) (°C) (months) (mm) (mm) (mm) (mm)

PhysgAsia2_HiResGridMetAsia2 (15.9-) 17.3 (-18.7) (7.1-) 8.9 (-11) 9-10 (1000-) 1400 (-1900) (100-) 130 (-170) (500-) 750 (-1000) (100-) 155 (-210) PhysgAsia2_HiResGridMetAsia1 (13-) 14.7 (-16) (3.8-) 6.5 (-9) 7-9 (800-) 1200 (-1500) (100-) 130 (-170) (500-) 650 (-820) (100-) 160 (-230)

MAT = mean annual temperature, CMMT = coldest month mean temperature, GROWSEAS = length of growing season, GSP = growing season precipitation, MMGSP = mean monthly growing season precipitation, X3.WET = precipitation of three consecutive wettest months, X3.DRY = precipitation of three consecutive driest months.

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Table 6 Classification of the Laurel Forest Biome.

Modern biomes Cenozoic Biomes

(Schroeder, 1998) (Olson et al., 2001) (Mai, 1995)

Laurel Forest Tropical and Subtropical Moist Broadleaf Forest Subtropical Rain and Laurel Forest; (northern range); Warm Temperate Evergreen Broadleaf Forest Temperate Broadleaf & Mixed Forest (Mixed Laurel Forest) (southern range) Laurel Forest with deciduous elements Temperate Broadleaf & Mixed Forest Warm Temperate Evergreen Broadleaf Forest (southern range); (Mixed Laurel Forest) Temperate Conifer Forest (p.p.) Nemoral Conifer Forest Temperate Conifer Forest (p.p.) Part of the above-mentioned biomes Ecotone Humid Broadleaf Tropical and Subtropical Moist Broadleaf Forest (northern range); Ecotone European Broadleaf Deciduous Forest and Laurel Forest Temperate Broadleaf & Mixed Forest Deciduous Forest and Warm Temperate Evergreen (southern range) Broadleaf Forest

4.2. CLAMP analysis for Güvem Formation western North America. Finally, there are only weak biogeographic affinities with the extant floras of and Africa. Nine oftentaxa Ninety-six operational taxonomic units (OTU) were scored for that are found in Australia and eight of the 17 taxa found in Africa are CLAMP (see Supplementary Material 2 for scoresheets and vector scores cosmopolitan. of all climate parameters). Due to the good preservation of leaf fossils most characters could be coded for each OTU resulting in a high 5. Discussion completeness statistic (almost 0.9), which gives a high level of con- fidence in the results. The reconstructed climate parameters suggest 5.1. Revised stratigraphic framework for early Miocene plant assemblages warm temperate conditions for mean annual temperature (mean of Turkey 17.3 °C) and coldest month mean temperature (mean 8.9 °C) and the duration of the growing season (ca. 9 months; Table 5). The relatively Several radiometric dates have become available during the past high growing season precipitation (1000–1900 mm) and the ratio of few (e.g. Atıcı et al., 2014; Ersoy et al., 2014; Akkemik et al., X3.WET to X3.DRY (ca. 5) further suggest precipitation seasonality but 2016; Aslan et al., 2017; Helvacı et al., 2017) considerably changing sufficient precipitation during the dry season (Cf climate). our understanding of plant fossil sites across Turkey. Specifically, sev- eral floras previously considered being of Serravallian (ca. 13 Ma)or 4.3. Biogeographic patterns for Güvem Formation late Miocene (< 11.6 Ma) age are now known to be of Aquitanian to Burdigalian age (ca. 23 to 18 Ma). Naturally, a robust stratigraphic The main biogeographic affinities of the early Miocene plant as- framework is the prerequisite for a meaningful evaluation of early semblages are Northern Hemispheric. This is illustrated using the Miocene vegetation and climate across large parts of western and combined leaf, wood and pollen record of the Güvem flora central Turkey. Hence, the approach of the present study was not to use (Supplementary Table S4). Most fossil taxa (60) are shared with the plant fossil data to date localities, but to use independent age con- flora of East (and Southeast) Asia and comprise relict conifers suchas straints for dating and to use plant macro- and microfossils to explore Calocedrus and Cathaya, and angiosperms such as Ailanthus and the palaeoenvironments and climate in the early Miocene of Turkey. Distylium. Likewise, a number of taxa present in the early Miocene of During recent years, it has become clear that traditional dating Turkey are today confined to the Americas and East and approaches such as palynostratigraphy and correlation of macro floras (Torreya, Engelhardioideae, Magnolia). A few taxa are American en- may be misleading. Apart from the cases mentioned in the present study demic elements [Smilax (havanensis group), Decodon, Comptonia]. (see introduction), prominent examples include the corrected ages of a Forty-seven and 43 taxa, respectively, are shared with the modern number of Himalayan-Tibet plant fossil localities (e.g. Su et al., 2018; floras of eastern North America and western North America. Arela- Markam Basin, south-eastern Tibet) and classical East Asian sites (e.g. tively high number of early Miocene taxa in Turkey are shared with Yu et al., 2017; Shanwang Formation, East ). In the Markam (22); of these, most are also found in eastern and Basin, the modern appearance of the flora suggested a Neogene age in

Fig. 2. Köppen signatures from the plant macrofossil and palynological record of Güvem (Beş Konak, Keseköy). The predominating climate types are warm temperate Cf (warm temperate fully humid) and Cw (warm temperate winter dry) climates. Moderate contribution of equatorial A climates underscores the ecotone situation of the inferred Laurel Forest Biome (for detailed explanation of the Köppen-Geiger climate classification see Kottek et al., 2006, Peel et al., 2007 and http://koeppen- geiger.vu-wien.ac.at/).

243 T. Denk, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 530 (2019) 236–248

Table 7 Haymana based on the common presence of small vessel diameters and Characteristic families of the modern Laurel Forest Biome. high vessel density in three taxa (Quercus, Pistacia, and Zelkova; Family Occurrence in early Miocene of Anatolia Akkemik et al., 2018b). An alternative explanation could be that these woods represent branches and not stems, which is also indicated by Cupressaceae Abundant in leaf, wood, and pollen record their dimensions (Akkemik, pers. comm. 2018-11-08). It is difficult, Pinaceae Abundant in leaf, wood, and pollen record based on the presence of only three taxa (Quercus, Pistacia, and Zelkova) Taxaceae Present in leaf record Tree ferns – to infer xeric or mesic conditions as this combination of genera may Aquifoliaceae Present in leaf and pollen record occur both in distinctly humid but also in highly seasonal communities. Present in leaf record In another paper, Akkemik et al. (2018a) compared petrified wood of Celastraceae – Acer from three Miocene localities of the Galatian Volcanic Province Clethraceae Present in pollen record (GVP) to wood of extant Acer from Turkey using xeromorphy ratios. All Fagaceae Abundant in leaf, wood, and pollen record Flacourtiaceae – of their wood samples produced signal corresponding to modern trees Hamamelidaceae incl. Altingiaceae Present in leaf, wood, and pollen record thriving under xeric conditions. However, since the position of the Illiciaceae – wood within the tree is not known it is difficult to assess the ecological Lauraceae Abundant in leaf fossil record significance of these findings. Vessel size within roots, trunkand Magnoliaceae Present and moderately diverse in leaf and pollen record branches vary significantly (see examples given in Akkemik et al., Myricaceae Present in leaf and pollen record 2018a). Furthermore, the common occurrence of false tree-rings in Myrsinaceae – Juniperus (Acarca Bayam et al., 2018) might suggest increased season- Present and moderately diverse in pollen ality. False tree-rings or intraannual density fluctuations (IADFs) are record common among trees and growing under Mediterranean Palms Present in leaf, wood, and pollen record Pittosporaceae – summer dry climates. Thus, many Mediterranean conifers and both Rosaceae Present in leaf, wood, and pollen record deciduous and evergreen species can rapidly adjust their cambial Rutaceae Present in pollen record activity to changing soil water availability (De Micco et al., 2016). The – frequency of IADFs varies considerably between species and among sites (De Micco et al., 2016) and IADFs are triggered during arid periods Woody plant families typical of the Laurel Forest biome (North Hemisphere: China, ; SE North America, , ; from Schroeder, 1998). but also during wet periods as earlywood-like cells in latewood. In addition, IADFs also occur in flood-prone riparian settings under humid traditional palaeobotanical papers, whereas new radiometric dating temperate climates (Mitsch, 1984; Young et al., 1993; Falcon-Lang suggests an Eocene to Oligocene age. Similarly, the classical mid-Mio- et al., 2001). Acarca Bayam et al. (2018) assigned fossil wood with false cene site Shanwang has been re-dated to early Miocene (similar to the tree-rings from the early Miocene GVP to the extant genus Juniperus. Burdigalian Güvem flora). This has major implications when assessing Juniperus usually occurs on well-drained soils and therefore false tree- evolutionary pathways in particular taxa and comparing vegetation rings in this wood may indicate precipitation seasonality rather than development at a continental scale. flooding. There has been some confusion about the palaeoenvironments of Turkey during the early Miocene. Changing lake δ18O values across 5.2. Early Miocene environments and biome reconstruction for Turkey central Turkey during the middle Aquitanian indicates changes in re- gional climates including more arid conditions (Lüdecke et al., 2013). The early Miocene environmental signal detected in this study is This is consistent with the structural and palaeogeographic evolution of unambiguous. AP/NAP ratios of 98/2–80/20, pollen percentages of this region during the early Miocene with regional marine transgres- individual tree taxa indicating the local and regional presence of trees, sions, lacustrine mini-basins and uplifted highs (Popov et al., 2004; leaf assemblages with high to very high proportions of evergreen Poisson et al., 2016; Akkiraz et al., 2018). In addition, pronounced broadleaved and needleleaf plants, and in situ fossilized trees suggest volcanism in the GVP and in western Turkey added to the topographic that forest vegetation was widespread during the early Miocene across variability. Tectonics and shifts in lake levels hence suggest fluctuations western and central Turkey. Furthermore, both the palynological and of wetter and drier periods. Biltekin (2018) documented basin devel- wood assemblages suggest close similarity with modern-day Schroeder's opment in the lacustrine Ilgın Basin, Konya, central Turkey, during the concept of the Laurel Forest Biome (Tables 6, 7). Laurophyllous taxa Aquitanian using a 40 m thick sequence. Three pollen zones were represented in the pollen records of Turkey are Sapotaceae, Ilex, Re- characterized by dominance of (1) Taxodioideae, (2) Cedrus, and (3) evesia, Citreae, Myrtaceae, Engelhardioideae, and evergreen Fagaceae. Pinus. Herbaceous taxa were nearly absent from all three pollen zones Of these, Sapotaceae, Reevesia, Citreae, and Myrtaceae are not recorded and the respective dominating tree taxa were also abundant in pollen in the leaf fossil record (possibly, because they are difficult to recognize zones in which they did not dominate. Biltekin (2018) interpreted this without additional information from leaf epidermal features). In con- as reflecting a trend from a warm and humid climate to cooler anddrier trast, Lauraceae are only found in the macrofossil record, as their pollen conditions in the Pinaceae dominated pollen zones. Joniak et al. (2018) generally has a very low fossilization potential (very little sporopollenin speculated that the change from swamp forest to well-drained forest deposited in the exine, e.g. Traverse, 2007). A further characteristic may have been accompanied by an opening up of the vegetation. This is feature of the Laurel Forest Biome (as opposed to tropical forest biomes) difficult to test, as both Cedrus dominated and Pinus dominated forests is the prominent role of conifer taxa (19 genera in modern forests; may perhaps be open forests. However, in general, a change Schroeder, 1998; Körner, 2013), which are absent from the tropics. This from humid to relatively drier environments does not necessarily cor- underscores the ecotone situation of laurel forests between tropical and respond to a change from closed to more open vegetation. temperate biomes (Fig. 3). Strömberg et al. (2007) investigated three sites located in the GVP Although the Laurel Forest Biome is characterized by perhumid (including Keseköy) using plant silica (phytoliths) and inferred open conditions (Prentice et al., 1992; Schroeder, 1998; Körner, 2013), some grass-dominated landscapes for Turkey and adjacent areas during the regions have less rainfall during the warm summer months (e.g. wes- early Miocene. Our data from 33 sites unambiguously show the con- tern North America, ) and hence xeromorphic plants are tinuous presence of forest vegetation across western and central Turkey. to be expected at different places such as on limestone or volcanic rocks It is noteworthy that sample EM3 (“laminated siltstone with fossil leaf or in coastal settings. impressions”) of Strömberg et al. (2007) has a distinct signal of open- Accordingly, open and/or xeric conditions have been inferred for habitat grasses. From approximately the same level, our concretion

244 T. Denk, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 530 (2019) 236–248

Fig. 3. Thermic vegetation zones and hygric variants with modern natural vegetation types (biomes; after Schroeder, 1998). Vegetation types discussed in the text are highlighted in grey. See Table 6 for corresponding modern and fossil biomes of Olson et al. (2001) and Mai (1995). sample from Keseköy and the palynological section of Yavuz-Işık (2008) to the ecotone between Temperate Broadleaf and Mixed Forest and have a distinct forest signal using threshold values of individual tree Tropical and Subtropical Moist Broadleaf Forest of Olson et al. (2001; taxa and the overall AP/NAP. In addition, the layers above and below Table 6). According to Prentice et al. (1992) the distribution of warm- this level are very rich in plant macrofossils indicating the presence of temperate evergreen trees usually is limited by mean temperatures of laurel forest dominated by evergreen Fagaceae and diverse conifers the coldest month (CMMT) below 5 °C with absolute minima > c. (Denk et al., 2017a). The Laurel Forest Biome that we inferred as the −10 °C (Fig. 3), whereas Temperate Broadleaf and Mixed Forests are zonal vegetation for large parts of Turkey may have provided numerous constrained by a CMMT of −2 to 5 °C. Both biomes have in common ecological niches (as seen today in the different forest regions re- rainfall values > 65% of annual evaporative demand. Thus, rainfall is presenting this biome) that could have sustained a diversity of open distributed seasonally but does not limit plant growth during the habitat grasses and hence is not in conflict with the high diversity of growing season. This ecotone situation is also illustrated by the CLAMP grasses reported by Strömberg et al. (2007). Poaceae are under- results, where CMMT is reconstructed above 5 °C with the PhysgA- represented in the plant macrofossil record, and their diversity is not sia2_HiResGridMet Asia2 calibration dataset whilst Asia1 reconstructs reflected in the pollen record as grass pollen commonly lacks diagnostic values slightly below 5 °C (Table 5). Further X3.wet to X3.dry ratios features even when investigated with SEM. Nevertheless, our data are indicate some seasonality but no water stress during the growing in conflict with Strömberg et al.'s (2007) interpretation of relatively season. Finally, it is important to stress that the reconstructed climate open vegetation such as savanna or open woodland in the early Mio- does not necessarily allow a differentiation of xeric or mesic forest (see cene GVP. Perhaps the phytolith data were biased towards lake margin Section 4.2.). However, xeric conditions possibly present in Cedrus open habitats, where the openness was maintained by fluctuating lake dominated forest communities are not equivalent to open environments levels. Under the right conditions it would not need a very large area of (Mayer and Aksoy, 1986). grass to source a lot of phytoliths (cf. Bremond et al., 2004). Climate inferences from Köppen signatures also suggest the ecotone In most early Miocene palynofloras from Turkey, arboreal pollen situation between tropical and warm temperate (subtropical) forest dominated distinctly (mainly > 90%, see Table 2). The only palyno- biomes. This is expressed in a strong C signature (ca. 40% Cf and Cw flora strongly deviating from this general pattern is from theKar- Köppen climate types) along with a moderate A signature (16% equa- acaören Formation, Sivas Basin (SIV1). Here, Amaranthaceae and torial A climates). The tropical signal is more pronounced in the pollen Poaceae are abundant (combined values range from > 35–10%, record (16%) than in the macro fossil record (12%). A possible ex- Akkiraz et al., 2018). In contrast to the other terrestrial sections con- planation could be the relatively high amount of unassigned entire sidered in our study, the Karacaören Formation comprises marine se- margined leaf fossils. Also, there is a higher amount of pantropical fa- diments and the more pronounced herb signal reflects the presence of milies identified from the pollen record than is recorded from theleaf coastal salt marshes adjacent to the depositional area. record. It is noteworthy that the tropical signature recorded in the leaf In a wider East Mediterranean context, the inferred Laurel Forest flora from Güvem (12%) is considerably higher than the onefrom Biome for the early Miocene of western and central Turkey appears to floras of southwestern Turkey (9%, Bouchal et al., have extended to (Lesbos, Kimi, Velitzelos et al., 2014; 2018; Supplementary Table S5). This illustrates that climatic trends as Mantzouka et al., 2016; Mantzouka, 2018) and to Bosnia and Croatia seen in globally averaged climate curves (cooler Burdigalian, warmer (Kvaček et al., 1993; Jiménez-Moreno et al., 2009). ) not necessarily are seen in local floras. Thus, tropical plant groups may have gone through bottlenecks prior to mid-Miocene 5.3. Early Miocene climate of Turkey warming and therefore show reduced diversity in middle Miocene floras (e.g. Sapotaceae). A qualitative assessment of 35 macro floras and pollen floras from Turkey strongly suggested the Laurel Forest Biome was the dominant 6. Conclusions biome in western and central Turkey during the Aquitanian and Burdigalian. The Laurel Forest Biome of Schroeder (1998) corresponds In this study, we used a fossil dataset of pollen and spores, leaves,

245 T. Denk, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 530 (2019) 236–248 and wood to investigate biomes and associated climate in the early Akkiraz, M.S., 2011. Vegetation and climate in the Miocene deposits of southern side of Miocene of Turkey. A novelty of this study is that a robust stratigraphic Büyük Menderes Graben, Şahinali-2 core, SW Turkey. Bull. Geosci. 86, 859–878. Akkiraz, M.S., Akgün, F., Utescher, T., Wilde, V., Bruch, A.A., Mosbrugger, V., Üçbaş, framework is used for early Miocene plant localities in Turkey. S.D., 2012. Palaeoflora and climate of lignite-bearing lower−middle Miocene sedi- Radiometric dates and dates derived from mammal faunas and marine ments in the Seyitömer and Tunçbilek sub-basins, Kütahya Province, Northwest fossils reveal that a number of localities traditionally considered being Turkey. Turk. J. Earth Sci. 21, 213–235. 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