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Central Anatolia, Turkey): a Window Into Early Neogene Vegetation and Environments in the Eastern Mediterranean

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Central Anatolia, Turkey): a Window Into Early Neogene Vegetation and Environments in the Eastern Mediterranean Acta Palaeobotanica 57(2): 237–338, 2017 e-ISSN 2082-0259 DOI: 10.1515/acpa-2017-0011 ISSN 0001-6594 The early Miocene flora of Güvem (Central Anatolia, Turkey): a window into early Neogene vegetation and environments in the Eastern Mediterranean THOMAS DENK 1, TUNCAY H. GÜNER 2, ZLATKO KVAČEK 3 and JOHANNES M. BOUCHAL1,4 1 Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden; e-mail: [email protected] 2 Department of Forest Botany, Faculty of Forestry, Istanbul University, Istanbul, Turkey; e-mail: [email protected] 3 Institute of Geology and Palaeontology, Faculty of Science, Charles University, Prague, Czech Republic; e-mail: [email protected] 4 Department of Palaeontology, University of Vienna, Vienna, Austria; e-mail: [email protected] Received 14 June 2017; accepted for publication 24 October 2017 ABSTRACT. The early Burdigalian (MN3) plant assemblage of the Güvem area (northwestern Central Anatolia) is preserved in lacustrine sediments of the Dereköy pyroclastics. Its age is well constrained by radiometric dates of basaltic rocks bracketing the pyroclastics, making the Güvem flora one of the extremely few precisely dated early Miocene floras in the Mediterranean region. The rich assemblage of impression fossils comprises ferns and fern allies (2 species), gymnosperms (12 spp.) and angiosperms (129 spp.). Ilex miodipyrena sp. nov. is described as a new fossil-species. The most diverse families in the assemblage are the Fagaceae with 12 taxa and the Fabaceae with 12 leaf morphotypes and one fruit taxon. Aquatic plants are represented by seven taxa, riparian (including palms) and swamp forest elements by >35 taxa, and lianas by three taxa (Smilax spp., Chaneya). The relatively large number of aquatic and riparian/swamp elements is congruent with the rich fish, amphibian and reptile record of the Güvem area. Another characteristic feature of the plant assemblage is the presence of various lobed leaves which show similarities with modern species of different families (e.g. Alangium, various Malvales). Trees and shrubs growing on well-drained soils and forming closed-canopy and open-canopy forests are the most diversified group (>70 taxa). In terms of number of specimens in the collection and based on field observations, by far the most abundant leaf fossils belong to evergreen oaks of Quercus drymeja and Q. mediter- ranea and to various types of foliage that cannot be assigned to a particular extant or extinct genus of Fagaceae. These sclerophyllous trees must have covered vast areas surrounding the wetlands that developed during the early Miocene in the Güvem Basin. Based on a recent reassessment of the ecology and taxonomic affinity of these trees, they are considered to reflect humid temperate climatic conditions but with a brief drier season during the winter months. These forests are more similar to the laurel forests of the southeastern United States and those stretching in a narrow belt south of the Himalayas to eastern central China. The large number of Fabaceae may indicate the presence of warm subtropical environments but this is difficult to assess, as they are known for having wide ecological ranges today and in the past. All in all, a larger part of the plant taxa point to forested vegetation. This is in agreement with previous palynological studies which detected only small amounts of her- baceous and grass pollen. Open patches of vegetation may have been restricted to river banks and to rocky areas in a volcanic landscape. The biogeographic patterns detected for the early Miocene of the Güvem assemblage are manifold; most taxa are widespread Northern Hemispheric elements. A substantial part of the species migrated from Asia into Europe during the (late) Paleogene and reached Anatolia during the early Miocene (Fagus, Paliu- rus, Chaneya, Ailanthus, Quercus kubinyii, Davallia haidingeri, Acer angustilobum, A. palaeosaccharinum). Fewer taxa may have been in Anatolia before they migrated to Europe (e.g. Nerium, Smilax miohavanensis, Quercus sosnowskyi). Finally, very few taxa are Anatolian endemics (e.g. Ilex miodipyrena). KEYWORDS: macrofossils, palaeobotany, palaeoenvironment, palaeobiogeography, Miocene, Turkey Brought to you by | Naturhistoriska riksmuseet Authenticated Download Date | 12/20/17 3:00 PM 238 T. Denk et al. / Acta Palaeobotanica 57(2): 237–338, 2017 INTRODUCTION The Miocene period was the last period in the Earth’s history when particularly warm conditions favoured rich forest vegetation in the Northern Hemisphere, including the sub- arctic region (Mai 1995, Zachos et al. 2001, Denk et al. 2011). In the early Miocene, tropi- cal waters flowed between the Indian Ocean and Atlantic, causing mild and equable condi- tions in western Eurasia (Rögl 1998). In Tur- key, the Pontides to the north and the Taurids to the south were not as high as today, and the Anatolian Plateau comprised lowlands, fresh- water lakes, and marshes (Popov et al. 2004, Schildgen et al. 2012, 2014). Subsequently, intermittent collision of the Afro-Arabian plate with Eurasia created a physical connec- tion between Africa and western Eurasia (Har- zhauser et al. 2007) which enabled migration of a wide range of large mammals into western Eurasia (Gomphotherium landbridge). There are remarkably few early Miocene plant macrofossil localities in the Mediterra- nean region that have been thoroughly stud- ied in more recent times and that are species- rich. From Spain, the macrofloras of La Rinconada (Ribesalbes, Castellón) and Mora de Rubielos (Teruel) have been described by Barrón and Postigo-Mijarra (2011), Posti go- Mijarra and Barrón (2013), and Barrón and Diéguez (2001). The Chattian/Aquitanian flora of Armissan (Aude, southern France) described by Saporta (1865) has never been revised. The same is true for the Chattian flo- ras of Aix-en-Provence and Manosque/Bois d’Asson (see Nury 1988 for the age of the Chattian floras). From Bosnia, Kvaček et al. (1993) described a moderately rich leaf flora. From Greece, Velitzelos (2002) published a preliminary revision of the rich flora of Kimi (Unger 1867), and Velitzelos et al. (1981) described leaf fossils of Lesvos. In addition, a number of early Miocene localities with dis- placed and in situ wood remains have been described from the island of Lesbos (e.g. Süss & Velitzelos 2010) and from Turkey (Akkemik Fig. 1. A–C. Geographical and regional geological setting of et al. 2009, 2016). early Miocene fossil localities in the Güvem area. A. Map The fossil sites in the vicinity of Güvem showing the geographical position of the Galatia Volcanic (Kızılcahamam district, Ankara Province; Province (GVP) and the Güvem Basin (B). B. Regional geo- logical map (simplified map based on Paicheler 1978; Tankut northwestern Central Anatolia; Fig. 1) have et al. 1998). C. Map showing Kasaplıgil’s collection localities been studied by palaeontologists for almost (simplified map based on Paicheler 1978);1 – Aköz locality (= vertebrate locality of Paicheler et al. 1978); 2 – Çerkeş road 50 years. Vertebrates were studied by Pai- “lone pine near the cemetery” locality; 3 – Demirciler locality; cheler et al. (1978), Hoek Ostende (2001), 4 – Beşkonak locality; 5 – Karga creek locality Brought to you by | Naturhistoriska riksmuseet Authenticated Download Date | 12/20/17 3:00 PM T. Denk et al. / Acta Palaeobotanica 57(2): 237–338, 2017 239 Rückert-Ülkümen (2003), Lopez et al. (2004), et al. 1996, Wilson et al. 1997, Tankut et al. 1998). and Dubois et al. (2010), among others. Among The GVP evolved as a consequence of the collision the recovered animal fossils are fishes, tad- of the Afro-Arabian and Eurasian plates (Rögl 1998). Potassium-argon dating of volcanic rocks determined poles, frogs, turtles, insectivores, and rodents. that the major phase of volcanic activity was during Most are typical of temperate and equatorial the early Miocene (17–20 Ma; Wilson et al. 1997). The monsoon climates in Southeast Asia (Hoek Güvem Basin is one of several volcanic complexes Ostende 2001). One exception is the Cteno- within the GVP. It is characterized by thick fluvio- lacustrine sedimentary infills alternating with pyro- dactylidae, rodents that are today typical of clastics. The ca 1100 m thick Güvem Formation (Tan- arid landscapes in Africa but which, based on kut et al. 1998) unconformably overlies Çukurviran their fossil record, are thought to have possibly dacite that has been dated to 19.7 ± 0.6 Ma (Wilson undergone substantial niche evolution. et al. 1997). The Dereköy pyroclastics (300 m; Fig. 2) Plant fossils discovered and collected by lie at the base of the Güvem Fm, built up by effusive volcanic material (andesitic blocks, ash flows, lapilli Baki Kasaplıgil in the late 1960s are now tuffs) and interfingered by lacustrine sediments (clay- stored at the University of California Museum stones, diatomite). From this lacustrine sequence of Paleontology (UCMP). They form the basis (Beşkonak sequence in Paicheler 1978) originate of the present study. Kasaplıgil intended to vertebrate, insect and plant fossils. On top of the publish a monograph on the flora of Güvem but Dereköy pyroclastics follow the Balta rhyolite (70 m), Bakacak andesite (200 m) and Oğlakcı dacite (50 m; unfortunately he died before he could complete Fig. 2). The latter two formations yielded K-Ar ages his monograph (http://ucmp.berkeley.edu/blog/ archives/939). A preliminary report was pub- lished by Kasaplıgil in 1977. At around the same time, Paicheler and Blanc (1981) pub- lished the
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