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  • 1.
    Bouchal, Johannes M.
    Swedish Museum of Natural History, Department of Paleobiology.
    Zetter, Reinhard (Contributor)
    Pollen and spores of the uppermost Eocene Florissant Formation, Colorado: A combined light and scanning electron microscopy study2016In: Grana, Vol. 55, no 3, p. 179-245Article in journal (Refereed)
    Abstract [en]

    The uppermost Eocene Florissant Formation, Rocky Mountains, Colorado, has yielded numerous insect, vertebrate, and plant fossils. Three previous comprehensive palynological studies investigated sections of lacustrine deposits of the Florissant Formation and documented the response of plant communities to volcanic eruptive phases but overall found little change in plant composition throughout the investigated sections. These studies reported up to 150 pollen and spore phenotypes. In the present paper we used a taxonomic approach to the investigation of dispersed pollen and spores of the Florissant Formation. Sediment samples from the shale units containing macrofossils were investigated using light microscopy (LM) and scanning electron microscopy (SEM). The general picture of the palynoflora is in agreement with previous studies. However, the combined LM and SEM investigation provides important complementary information to previous LM studies. While a fairly large amount of previous pollen determinations could be confirmed, the purported taxonomic affinities of several pollen phenotypes need to be revised. For example, pollen referred to as Podocarpus or Podocarpidites sp. belongs to the Pinaceae Cathaya, Malus/Pyrus actually belongs to Dryadoideae, pollen of the form genus Boehlensipollis referred to as Proteaceae/Sapindaceae/Elaeagnaceae or Cardiospermum belongs to Sapindaceae but not to Cardiospermum, and pollen of Persicarioipollis sp. B with previously assumed affinities to Polygonaceae actually belongs to Thymelaeaceae. Pandaniidites and one type of Malvacipollis cannot be linked with Pandanaceae and Malvaceae. A few taxa are new records for Florissant (Ebenaceae: Diospyros; Mernispermaceae; Trochodendraceae: Tetracentron). In general, SEM investigations complement the LM palynological studies and improve the identification of dispersed pollen and spores and enable integration of data from dispersed fossil pollen into a wide range of comparative morphological, taxonomic, evolutionary, biogeographic, and phylogenetic studies.

  • 2.
    Bouchal, Johannes M.
    Swedish Museum of Natural History, Department of Paleobiology.
    The middle Miocene palynofloras of the Salihpaşalar lignite mine (Yatağan Basin, southwest Anatolia): environmental characterisationand comparison with palynofloras from adjacent basins2019In: Palaeobiodiversity and Palaeoenvironments, ISSN 1867-1594, E-ISSN 1867-1608Article in journal (Refereed)
    Abstract [en]

    As the third part of an ongoing investigation of middle Miocene palynofloras in the Yatağan Basin (YB), southwestern Anatolia, thepalynofloras of the Salihpaşalar lignite mine in the main YB were studied. Seven types of algal spores, aplanospores/zygospores orcysts, six types of lycophyte and fern spores, 12 types of gymnosperm pollen and 90 types of angiosperm pollen were identified. Of atotal of ca. 140 plant taxa described from the YB, over 10% are confined to the Salihpaşalar assemblage. Differences between coevalpalynofloras of the Sekköy Member might reflect changing or prograding depositional environments. A number of rare accessorialtaxa reflect these local differences: Pilularia, Valeriana, Drosera and Persicaria aff. amphibia only occur at Salihpaşalar and aretypical of shallow water or temporary ponds associated with a lake shore. Apart from this, all the palynofloras, originating from thelignite seams and overlying limnic limestones (uppermost Turgut and Sekköy Member), of the YB are strongly indicative of extensivewoody vegetation with a dominance of diverse Fagaceae and Pinaceae. In addition, a list comparing the well-documented YBpalynomorphs to morphologically similar palynomorphs of published late early to middle Miocene plant assemblages of westernAnatolian was compiled. Such a comparison reveals that in many instances different taxon names have been used to denote the sametaxa. Hence, resolving these synonymies is a prerequisite of any meaningful comparison of palynofloras in the region.

  • 3.
    Bouchal, Johannes M.
    et al.
    Swedish Museum of Natural History, Department of Paleobiology.
    Denk, Thomas
    An overview of the palynoflora of the Miocene Yatağan basin, Turkey2014Conference paper (Other academic)
    Abstract [en]

    The palynoflora of the lignite strip mines of the Yatağan basin, located in the Muğla province of western Turkey, is the focus of this study. Samples were taken from the Eskihisar, Salihpasalar and Tinaz mines. In the Yatağan basin two Miocene formations, formed from river and lake deposits, Eskihisar Formation (middle Miocene) and Yatağan Formation (late Miocene) have been designated. Both show a general lithology consisting of conglomerate, sandstone, claystone, limestone and tuffite, the mined/excavated lignite bearing strata are restricted to the Eskihisar Formation.

    Until now, pollen from the Yatağan basin has mostly been described according to conventional morphological nomenclature, using light microscopy (LM) only. In this study, the same individual pollen grains are investigated by using both, LM and scanning electron microscopy (SEM). The  high resolution pictographs allow a higher taxonomic resolution.

    The rich palynoflora (Table 1) is comprised of diverse spores (at least nine morphotypes), gymnosperm pollen from Cupressaceae, Gnetales, Pinaceae, and angiosperm pollen from Poaceae, Typhaceae, Altingiaceae,  Amaranthaceae (Chenopodieae), Anacardiaceae, Apiaceae, Betulaceae, Buxaceae, Caprifoliaceae (Dipsacoideae, Lonicera) Caryophyllaceae, Compositae (Asteroideae, Cichoriodeae), Cornaceae, Eucommiaceae, Fabaceae, Fagaceae (Fagus, Quercus, Trigonobalanopsis) Geraniaceae, Juglandaceae, Malvaceae, Myricaceae, Nymphaeaceae, Oleaceae, Palmae, Plumbaginaceae (Armeria, Plumbago), Polygonaceae (Rumex), Salicaceae, Sapindaceae (Acer), Smilacaceae, and Ulmaceae (Cedrelospermum, Ulmus, Zelkova).

  • 4.
    Bouchal, Johannes M.
    et al.
    Swedish Museum of Natural History, Department of Paleobiology.
    Denk, Thomas
    Swedish Museum of Natural History, Department of Paleobiology.
    Grímsson, F.
    Palynostratigraphical correlation of the excavated Miocene lignite seams of the Yatağan basin (Muğla Province, south-western Turkey)2016Conference paper (Other academic)
    Abstract [en]

    The excavated main lignite seams and overlying lacustrine sediments of the opencast mines Eskihisar, Salihpaşalar, and Tınaz, Muğla Province, south-western Turkey were investigated using a high taxonomic resolution palynological approach.

    The Eskihisar section comprises 47m and 56 samples of which 30 were usable for palynological analysis. The Tınaz section comprises 75 m and 29 samples of which 15 were usable for palynological analysis. Finally, the Salihpaşalar section comprises 25 m and 26 samples of which 16 were usable for palynological analysis. The age of the palynological sections is middle to late Miocene based on radiometric dating and vertebrate fossils.

     

    In order to investigate dispersed pollen and spores and their botanical affinities a combined light microscopy and scanning electron microscopy approach was used. The rich palynoflora comprises seven types of algal cysts (Botryococcus, Zygnemataceae), seventeen spore types (Lycopsida, Marsileaceae, Osmundaceae, Pteridaceae, Polypodiaceae), 14 types of gymnosperm pollen (Ephedraceae, Cupressaceae, Pinaceae), five types of monocotyledone pollen (Poaceae, Typhaceae) and ca 90 dicotyledone pollen types (Altingiaceae, Amaranthaceae, Anacardiaceae, Apiaceae, Aquifoliaceae, Asteraceae, Betulaceae, Campanulaceae, Cannabaceae, Caprifoliaceae, Caryophyllaceae, Ericaceae, Eucommiaceae, Euphorbiaceae, Fabaceae, Fagaceae, Geraniaceae, Juglandaceae, Lamiaceae, Linaceae, Lythraceae, Malvaceae, Myricaceae, Oleaceae, Onagraceae, Plumbaginaceae, Polygonaceae, Ranunculaceae, Rosaceae, Salicaceae, Sapindaceae, Sapotaceae, Ulmaceae).

     

    The objectives of this investigation were (1) to evaluate whether the three palynological sections were deposited at the same time, and (2) to show regional vegetation differences within a single sedimentary basin.

     

    We found three general pollen zones corresponding to different sedimentary settings and palaeoenvironments. The first pollen zone was linked to lignite formation (swamp forest, fern spores, Alnus, Decodon). The second pollen zone reflects lacustrine conditions (Typhaceae) and surrounding hinterland vegetation dominated by Fagaceae. The third pollen zone is dominated by herbaceous taxa, whereas woody taxa are less diverse and less abundant.

     

    In general, the three palynological sections are congruent in reflecting distinct pollen zones. However main vegetation types may be represented by different dominating taxa (e. g. Alnus dominace in Eskihisar and Tınaz localities while absent in Salihpaşalar) and rare taxa may differ between localities.

     

    Our results demonstrate that in order to achieve a comprehensive understanding of environmental and vegetation conditions in a sedimentary basin, a single palynological section (locality) may not capture the entirety of environmental conditions and changes.

  • 5.
    Bouchal, Johannes M.
    et al.
    Swedish Museum of Natural History, Department of Paleobiology.
    Denk, Thomas
    Swedish Museum of Natural History, Department of Paleobiology.
    Grímsson, F.
    Zetter, Reinhard
    The middle Miocene palynoflora and palaeoenvironments of Eskihisar (Yatağan Basin, southwestern Anatolia):: a combined LM and SEM investigation2016In: Botanical journal of the Linnean Society, ISSN 0024-4074, E-ISSN 1095-8339, Vol. 182, no 1, p. 14-79Article in journal (Refereed)
    Abstract [en]

    Anatolia was a crossroads for mammal migration during the Miocene due to intermittent land connections between Africa and Anatolia and persisting warm conditions. Here, we investigated a palynological section from middle Miocene sediments of Eskihisar (southwestern Anatolia) in order to establish biogeographic links of the palynoflora and to infer the palaeoenvironment. Four algal palynomorphs, nine spore taxa, eight gymnosperms, three monocots, and 67 dicot pollen types were encountered and investigated using the “single grain method” that combines light microscopy and scanning electron microscopy. Two pollen zones reflect different phases of basin development. Zonal vegetation remained fairly stable across the section and reflects heterogeneous environments including broad-leaved deciduous forest, subtropical forest, and sclerophyllous and semi-evergreen oak forest. Conifers were accessory elements in the broad-leaved deciduous forest communities and replaced these at higher elevations. Some herbaceous taxa (Plumbaginaceae) indicate scattered occurrences of sandy and/or rocky soils. Biogeographic affinities are general Northern Hemispheric, North American, and East Asian as also suggested by the macro fossil record. Only two taxa provide potential biogeographic links with the African flora. This suggests that biome shifts of plant taxa between African subtropical /tropical biomes and Anatolian (western Eurasian) temperate forests and shrublands may have been rare in the middle Miocene.

  • 6.
    Bouchal, Johannes M.
    et al.
    Swedish Museum of Natural History, Department of Paleobiology.
    Grímsson, F.
    Zetter, Reinhard
    Denk, Thomas
    Swedish Museum of Natural History, Department of Paleobiology.
    Some new pollen taxa from the middle Miocene of south western Anatolia2016Conference paper (Other academic)
    Abstract [en]

    In an ongoing study, focussing on the plant fossils and palynofloras of the lignite strip mines of the Yatağan basin(Muğla province), a number of pollen taxa, previously not reported from middle Miocene terrestrial sediments of Anatolia were encountered.

  • 7.
    Bouchal, Johannes M.
    et al.
    Swedish Museum of Natural History, Department of Paleobiology.
    Güner, Tuncay H.
    Swedish Museum of Natural History, Department of Paleobiology. Faculty of Forestry, Department of Forest Botany, Istanbul University Cerrahpa¸sa, 34473 Bahçeköy, Istanbul, Turkey.
    Denk, Thomas
    Swedish Museum of Natural History, Department of Paleobiology.
    Middle Miocene climate of southwestern Anatolia from multiple botanical proxies2018In: Climate of the Past Discussions, ISSN 1814-9340, E-ISSN 1814-9359, Vol. 14, p. 1427-1440Article in journal (Other academic)
    Abstract [en]

    The middle Miocene climate transition (MMCT) was a phase of global cooling possibly linked to decreasing levels of atmospheric CO2. The MMCT coincided with the European  Mammal Faunal Zone MN6. From this time, important biogeographic links between Anatolia  and eastern Africa include the hominid Kenyapithecus. Vertebrate fossils suggested mixed  open and forested landscapes under (sub)tropical seasonal climates for Anatolia. Here, we  infer the palaeoclimate during the MMCT and the succeeding cooling phase for a middle Miocene (14.8–13.2 Ma) of an intramontane basin in southwestern Anatolia using three2palaeobotanical proxies: (i) Köppen signatures based on the nearest-living-relative principle. (ii) Leaf physiognomy analysed with the Climate Leaf Analysis Multivariate Program (CLAMP). (iii) Genus-level biogeographic affinities of fossil floras with modern regions. The three proxies reject tropical climates for the MMCT of southwestern Anatolia and instead infer warm temperate C climates. Köppen signatures reject summer-dry Cs climates but cannot discriminate between fully humid Cf and winter-dry Cw; CLAMP reconstructs Cf climate based on the low X3.wet/X3.dry ratio. Additionally, we assess whether the palaeobotanical record does resolve transitions from the warm Miocene Climatic Optimum (MCO, 16.8–14.7 Ma) into the MMCT (14.7–13.9 Ma), and a more pronounced cooling at 13.9–13.8 Ma, as reconstructed from benthic stable isotope data. For southwestern Anatolia, we find that arboreal taxa predominate in MCO floras (MN5), whereas in MMCT floras (MN6) abundances of arboreal and non-arboreal elements strongly fluctuate indicating higher structural complexity of the vegetation. Our data show a distinct pollen zone between MN6 and MN7+8 dominated by herbaceous taxa. The boundary MN6 and MN7+8, roughly corresponding to a first abrupt cooling at 13.9–13.8 Ma, possibly might be associated with this herb-rich pollen zone.

  • 8.
    Bouchal, Johannes M.
    et al.
    Swedish Museum of Natural History, Department of Paleobiology.
    Güner, Tuncay H.
    Denk, Thomas
    Swedish Museum of Natural History, Department of Paleobiology.
    Palynological and palaeobotanical investigations in the Miocene Yatağan basin, Turkey: High-resoluton taxonomy and biostratigraphy2015Conference paper (Other academic)
    Abstract [en]

    The subject of this study is the palynology (biostratigraphic and taxonomic) and the plant remains of the lignite strip mines of Eskihisar, Salihpasalar, and Tinaz (Muğla province, western Turkey). In the Yatağan basin two Miocene to Pliocene formations are present, the Eskihisar Formation (early to middle Miocene) and the Yatağan Formation (late Miocene to early Pliocene). Both formations represent river and lake deposits consisting mainly of conglomerate, sandstone, claystone, limestone, tuffite, and intercalated lignite; the thickest, actively mined lignite seams occur in the Sekköy member of the Eskihisar Formation.

    Previous palynological studies of the palynoflora of the Yatağan basin mainly focussed on its biostratigraphic and palaeoclimatic significance, using conventional morphological nomenclature and light microscopy (LM).

    In this study the „single grain method“ is applied. Using this method, the same individual pollen grains are investigated by using both LM and scanning electron microscopy (SEM). The resulting high-resolution pictographs enable a much higher taxonomic resolution.

     

    The studied palynoflora is very rich and taxonomically diverse. Cryptogams are represented by more than ten spore morphotypes of at least three families (Osmundaceae, Pteridaceae, Polypodiaceae). Gymnosperm pollen is dominated by Cupressaceae, Gnetales (Ephedra), and Pinaceae (Cathaya, Keteleeria, Pinus). Angiosperm pollen can be assigned to 57 different genera belonging to Poaceae, Typhaceae, Altingiaceae, Amaranthaceae (Chenopodieae), Anacardiaceae, Apiaceae (three types), Asteraceae (Asteroideae, Cichoriodeae), Betulaceae (Alnus, Betula, Carpinus, Ostrya) Buxaceae, Campanulaceae, Caprifoliaceae (Lonicera), Caryophyllaceae, Dipsacaceae, Eucommiaceae, Euphorbiaceae, Fabaceae, Fagaceae (Fagus, Quercus, Trigonobalanopsis) Geraniaceae, Juglandaceae, Linnaceae (Linnum), Malvaceae, Myricaceae, Nymphaeaceae, Oleaceae (four different types), Plumbaginaceae (Armeria,), Polygonaceae (Rumex), Rosaceae, Sapindaceae (Acer), Ulmaceae (Cedrelospermum, Ulmus, Zelkova), and Zingiberales (Spirematospermum).

     

    In addition, more than two thousand plant macrofossils were collected in the course of repeated field trips, including remains of Pinaceae, Berberidiaceae (Mahonia), Betulaceae (Alnus, Carpinus), Buxaceae (Buxus), Fagaceae (Fagus, Quercus), Lauraceae, Malvaceae (Tilia), Myricaceae (Myrica), Rosaceae, Salicaceae (Populus, Salix), Sapindaceae (Acer), Smilacaceae (Smilax), Typhaceae (Typha), Ulmaceae (Zelkova).

     

    A combined analysis integrating these rich and diverse plant macro- and microfossil records will lead to a better understanding and refined reconstruction of the vegetation in the Yatağan basin during the middle to late Miocene.

  • 9.
    Bouchal, Johannes M.
    et al.
    Swedish Museum of Natural History, Department of Paleobiology.
    Mayda, Serdar
    Natural History Museum, Ege University, 35100 Izmir, Turkey.
    Zetter, Reinhard
    University of Vienna, Department of Palaeontology, Vienna, Austria.
    Grímsson, Fridgeir
    University of Vienna, Department of Palaeontology, Vienna, Austria.
    Akgün, Funda
    Dokuz Eylül University, Department of General Geology, 35210 Izmir, Turkey.
    Denk, Thomas
    Swedish Museum of Natural History, Department of Paleobiology.
    Miocene palynofloras of the Tınaz lignite mine, Muğla, southwest Anatolia: taxonomy, palaeoecology and local vegetation change2017In: Review of Palaeobotany and Palynology, ISSN 0034-6667, E-ISSN 1879-0615, Vol. 243, p. 1-36Article in journal (Refereed)
    Abstract [en]

    Middle Miocene deposits exposed at the Tınaz lignite mine, Yatağan Basin, Muğla, southwestern Turkey, were palynologically investigated. The Tınaz lignite mine section belongs to the Eskihisar Formation. The lignite seam at the base of the section represents the uppermost part of the Turgut Member. Above, c. 65 m of clayey siltstone, limestone, and marls represent the Sekköy Member. Nine spores, 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 pollen zones were recognized, of which pollen zone 1 corresponds to the formation of the main lignite seam and reflects the change from a fluviatile to a lacustrine depositional setting. Pollen zones 2 and 3 and a transitional zone 2-3 reflect different stages of lake development and a shift in local vegetation from forested (pollen zones 1 and 2) to more open (transitional zone 2-3, zone 3). Interpreting changes in regional vegetation from pollen zones 1 to 3 is not straightforward as changes in the pollen spectra may be affected by changing contributions of airborne and water transported pollen and spores to the observed palynoassemblages. Age inference for the Tınaz lignite mine section has been complicated by the absence of datable ash layers, associated mammal faunas, or marine sediments. However, pollen zone 3 shares key features with the pollen spectrum recovered from the nearby mammal site Yenieskihisar (upper part of Sekköy Member) for which an age of 12.5-11.2 Ma has been suggested, and to the youngest pollen zone recovered from the mammal locality Çatakbağyaka, 10 km south of Tınaz, that probably represents mammal zone MN7/8 instead of MN5 or MN6 as previously suggested. In contrast, pollen zones 1 and 2 are fairly similar to the basal parts of the Çatakbağyaka pollen flora (uppermost parts of Turgut Member, basalmost parts of Sekköy Member). Furthermore, new mammal data from the Yatağan basin suggest that the layers below pollen zone 1 are MN4/5, and that carnivores cooccuring with pollen zone 1 in the main lignite seam of Eskihisar probably belong to MN6. Hence, a Langhian to Serravallian age can be inferred for pollen zones 1 and 2 of the Tınaz lignite mine section, and a late Serravallian age for pollen zone 3. Palaeobiogeographic relationships of the palynofloras are generally northern hemispheric, with many north temperate tree taxa showing modern disjunctions East Asia- NorthAmerica (Tsuga, Carya), East Asia- western Eurasia (Zelkova), East Asia- North America- western Eurasia (Liquidambar), or restricted to East Asia (Cathaya, Eucommia) or North America (Decodon). A few taxa belong to extinct lineages that have complex biogeographic patterns (Engelhardioideae, Cedrelospermum). The presence of Picrasma (Simaroubaceae) in the lower lignite layers of pollen zone 1 is remarkable, as the botanical affinities with the enigmatic flower Chaneya present in early to middle Miocene deposits of Turkey and Central Europe have recently been shown to be with Picrasma

  • 10.
    Bouchal, Johannes Martin
    Swedish Museum of Natural History, Department of Paleobiology. University of Vienna.
    EVOLUTIONARY TRENDS AND ECOLOGICAL DIFFERENTIATION IN EARLY CENOZOIC FAGACEAE OF WESTERN NORTH AMERICA2014In: American Journal of Botany, ISSN ISSN: 0002-9122., Vol. 101, no 8, p. 1332-1349Article in journal (Refereed)
    Abstract [en]

    • Premise of the study: The early Cenozoic was a key period of evolutionary radiation in Fagaceae. The common notion is that

    species thriving in the modern summer-dry climate of California originated in climates with ample summer rain during the

    Paleogene.

    • Methods: We investigated in situ and dispersed pollen of Fagaceae from the uppermost Eocene Florissant fossil beds, Colorado,

    United States, using a combined light and scanning electron microscopy approach.

    • Key results: Pollen types of Castaneoideae with affi nities to modern Castanea , Lithocarpus , and Castanopsis were recognized.

    Pollen of the extinct genus Fagopsis represents a derived type of Castaneoideae pollen. Infrageneric groups of Quercus were

    well represented, including pollen of Group Protobalanus. The taxonomic diversity of Fagaceae and of the total plant assemblage

    indicates a mosaic of microclimates, that range from pronounced to weakly seasonal climates and depend on slope aspect

    and elevation. Continental climatic conditions may have triggered the evolution of sclerophyllous leaves and adaptive radiation

    in Quercus and other taxa thriving today under distinctly summer-dry and winter-dry climates.

    • Conclusions: Vegetation types similar to modern vegetation belts of the Coastal Ranges (chaparral, nemoral conifer forest)

    were established in the Front Range in the late Eocene. Coeval plant assemblages from the Coastal Ranges of California indicate

    distinctly subtropical, moist climates. Hence, characteristic elements found today in the summer-dry and winter-dry climates

    of Pacifi c North America ( Quercus Group Protobalanus, Notholithocarpus ) may opportunistically have dispersed into

    their modern ranges later in the Cenozoic. This scenario is in contrast to the evolution and migration patterns of their western

    Eurasian Mediterranean counterparts ( Quercus Group Ilex).

  • 11.
    Denk, Thomas
    et al.
    Swedish Museum of Natural History, Department of Paleobiology.
    Güner, Tuncay H.
    , Istanbul University, Istanbul, Turkey.
    Kvaček, Zlatko
    Charles University, Prague, Czech Republic.
    Bouchal, Johannes M.
    Swedish Museum of Natural History, Department of Paleobiology. johannes.bouchal@nrm.se.
    The early Miocene flora of Güvem (Central Anatolia, Turkey): a window into early Neogene vegetation and environments in the Eastern Mediterranean2017In: Acta Palaeobotanica, ISSN 0001-6594, E-ISSN 1427-6402, Vol. 57, no 2, p. 237-338Article in journal (Refereed)
    Abstract [en]

    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. mediterranea 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 herbaceous 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, Paliurus, 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).

  • 12.
    Denk, Thomas
    et al.
    Swedish Museum of Natural History, Department of Paleobiology.
    Velitzelos, Dimitrios
    National and Kapodistrian University of Athens, Faculty of Geology and Geoenvironment, Department of Historical Geology and Paleontology, Panepistimiopolis, Athens 15784, Greece.
    Güner, Tuncay H.
    Istanbul University, Faculty of Forestry, Department of Forest Botany, 34473 Bahceköy, Istanbul, Turkey.
    Bouchal, Johannes M.
    Swedish Museum of Natural History, Department of Paleobiology.
    Grímsson, F.
    University of Vienna, Department of Palaeontology, 1090 Vienna, Austria.
    Grimm, Guido
    Department für Paläontologie, Universität Wien, Wien, Austria.
    Taxonomy and palaeoecology of two widespread western Eurasian Neogene sclerophyllous oak species: Quercus drymeja Unger and Q. mediterranea Unger2017In: Review of Palaeobotany and Palynology, ISSN 0034-6667, E-ISSN 1879-0615, ISSN 0034-6667, Vol. 241, p. 98-128Article in journal (Refereed)
    Abstract [en]

    Sclerophyllous oaks (genus Quercus) play important roles in Neogene ecosystems of south-western Eurasia. Modern analogues (‘nearest living relatives’) for these oaks have been sought among five of six infrageneric lineages of Quercus, distributed across the entire Northern Hemisphere. A revision of leaf fossils from lower Miocene to Pliocene deposits suggests that morphotypes of the Quercus drymeja complex are very similar to a number of extant Himalayan, East Asian, and Southeast Asian species of Quercus Group Ilex and may indicate subtropical, relatively humid conditions. Quercus mediterranea comprises leaf morphotypes that are encountered in modern Mediterranean species of Quercus Group Ilex, but also in Himalayan and East Asian members of this group indicating fully humid or summer-wet conditions. The fossil taxa Quercus drymeja and Q. mediterranea should be treated as morphotype complexes, which possibly comprised different biological species at different times. Quercus mediterranea, although readily recognizable as a distinct morphotype in early to late Miocene plant assemblages, may in fact represent small leaves of the same plants that constitute the Quercus drymeja complex. Based on the available evidence, the taxa [GG1] forming the Q. drymeja complex and Q. mediterranea thrived in fully humid or summer-wet climates. The onset of the modern vegetational context of Mediterranean sclerophyllous oaks is difficult to trace, but may have been during the latest Pliocene/early Pleistocene.

  • 13. Grimm, Guido
    et al.
    Bouchal, Johannes M.
    Swedish Museum of Natural History, Department of Paleobiology.
    Denk, Thomas
    Swedish Museum of Natural History, Department of Paleobiology.
    Potts, Alastair
    Fables and foibles: A critical analysis of the Palaeoflora database and the Coexistence Approach for palaeoclimate reconstruction2016In: Review of Palaeobotany and Palynology, ISSN 0034-6667, E-ISSN 1879-0615, Vol. 233, p. 216-235Article in journal (Refereed)
    Abstract [en]

    The ‘Coexistence Approach’ is amutual climate range (MCR) technique combinedwith the nearest-living relative (NLR) concept. It has been widely used for palaeoclimate reconstructions based on Eurasian plant fossil assemblages; most of them palynofloras (studied using light microscopy). The results have been surprisingly uniform, typically converging to subtropical, per-humid or monsoonal conditions. Studies based on the Coexistence Approach have had a marked impact in literature, generating over 10,000 citations thus far. However, recent studies have pointed out inherent theoretical and practical problems entangled in the application of this widely used method. But so far little is known how results generated by the Coexistence Approach are affected by subjective errors, data errors, and violations of the basic assumptions. The majority of Coexistence Approach studies make use of the Palaeoflora database (the combination of which will be abbreviated to CA + PF). Testing results produced by CA + PF studies has been hindered by the general unavailability of the contents in the underlying Palaeoflora database; two exceptions are the mean-annual temperature tolerances and lists of assigned associations between fossils and nearest-living relatives. Using a recently published study on the Eocene of China,which provides the first and only insight into the data structure of the Palaeoflora database,we compare the theory and practice of Coexistence Approach using the Palaeoflora database (CA+PF).We show that CA+PF is riddled by association and climate data error.We reveal flaws in the application of the Coexistence Approach,which is often in stark contrast to the theory of the method. We show that CA + PF is highly vulnerable against numerous sources of errors, mainly because it lacks safeguards that could identify unreliable data. We demonstrate that the CA+PF produces coherent, pseudo-precise results even for artificially generated, randomplant assemblages. AlternativeMCR-NLR methods can surpass the most imminent deficits of the Coexistence Approach, and may be used as a stop-gap until more accurate bioclimatic and distribution data on potential Eurasian NLRs, and theoretically and statistically robust methods will become available. Finally, general guidelines are provided for the future application of methods using the mutual climatic range with nearest living relatives approach when reconstructing climate from plant fossil assemblages.

  • 14. Grímsson, F.
    Meller, Barbara (Contributor)
    Bouchal, Johannes M. (Contributor)
    Swedish Museum of Natural History, Department of Paleobiology.
    Grimm, Guido (Contributor)
    Combined LM and SEM study of the middle Miocene (Sarmatian) palynoflora from the Lavanttal Basin, Austria: Part III. Magnoliophyta 1 – Magnoliales to Fabales2015In: Grana, ISSN 0017-3134, E-ISSN 1651-2049, Vol. 54, p. 85-128Article in journal (Refereed)
    Abstract [en]

    Previous studies on the palynoflora from the Lavanttal Basin show that it contains a rich

    assemblage of spores and gymnosperm pollen. Present and ongoing investigations of

    dispersed angiosperm pollen suggest a high diversity within this group, and due to the

    excellent preservation of the material some rare pollen types are recognized. The Magnoliales

    to Fabales pollen record documented here contains 30 different taxa. Only a few pollen types

    are assigned to Magnoliids (four taxa); these are rare in the pollen record. Similarly, the

    Commelinids comprise five taxa and are also rare. Most of the angiosperm pollen originate

    from Eudicots, 21 taxa. Of the angiosperm taxa documented here, Magnolia , Carex ,

    Ranunculaceae, Platanus , Trochodendron , Buxus , Cercidiphyllum , Daphniphyllum ,

    Distylium , Fortunearia , Parrotia , Parthenocissus , Vitis , Euphorbia , Salix , and

    Papilionoideae are recorded for the first time from the Lavanttal Basin. This also includes the

    first fossil pollen record of Trochodendron  worldwide and the first reliable pollen record of

    Daphniphyllum . Several of the taxa described here had a wide Northern Hemispheric

    distribution from Eocene until the end of the Miocene. Also, key relatives of the fossil taxa

    are presently confined to humid warm-temperate environments, suggesting a very mild

    climate during the middle Miocene (Sarmatian) of the Lavanttal area. Some of the taxa

    encountered also support previous observations that the sediments of the Lavanttal Basin

    accumulated in a lowland wetland environment. This is based on pollen from aquatic taxa

    thriving in lakes, streams and swamps, and pollen of terrestrial plant taxa occupying margins

    of lakes and streams, backswamps, floodplains, river plains, and hummocks. Other 

     angiosperm pollen clearly originate from plants thriving on drier substrates, reflecting various

    vegetation units of the mixed evergreen/deciduous broad-leaved/conifer forests surrounding

    the wetland basin.

  • 15. Grímsson, F.
    Bouchal, Johannes M. (Contributor)
    Swedish Museum of Natural History, Department of Paleobiology.
    Meller, Barbara (Contributor)
    Zetter, Reinhard (Contributor)
    Combined LM and SEM study of the middle Miocene (Sarmatian) palynoflora from the Lavanttal Basin, Austria: part IV. Magnoliophyta 2 – Fagales to RosalesIn: Grana, ISSN 0017-3134, E-ISSN 1651-2049Article in journal (Refereed)
    Abstract [en]

    An ongoing investigation of the middle Miocene (Sarmatian) palynoflora from the Lavanttal Basin continues to show that it contains an extremely rich assemblage of angiosperm taxa. The Fagales to Rosales pollen record documented here contains 34 different taxa belonging to the Betulaceae (Alnus, Betula, Carpinus, Corylus, Ostrya), Fagaceae (Castanea, Fagus, Quercus Groups Cerris, Ilex, Cyclobalanopsis, Quercus/Lobatae), Juglandaceae (Engelhardioideae, Carya, Juglans, Pterocarya), Myricaceae (Morrella vel Myrica), Cannabaceae (Celtis), Elaeagnaceae (Elaeagnus), Rhamnaceae, Rosaceae (Prunus) and Ulmaceae (Cedrelospermum, Ulmus, Zelkova). Two of the pollen types represent extinct genera, Trigonobalanopsis and Cedrelospermum, and are also reported for the first time from the Lavanttal Basin along with pollen of Rhamnaceae and Prunus. The different types of Quercus pollen are now affiliated with Groups Cerris, Cyclobalanopsis, Ilex and Quercus/Lobatae based on sculpturing elements observed using scanning electron microscopy (SEM). Köppen signatures of potential modern analogues of the fossil Fagales and Rosales suggest a subtropical (Cfa, Cwa) climate at lower elevation and subsequent subtropical to temperate climate with altitudinal succession (Cfa→Cfb/Dfa→Dfb; Cwa→Cwb→Dwb) in the Lavanttal area during accumulation of the palynoflora. Most of the fossil taxa have potential modern analogues that can be grouped as nemoral and/or merido-nemoral vegetation elements, and the diversity of Fagales indicates a varying landscape with a high variety of niches.

  • 16. Grímsson, F.
    et al.
    Bouchal, Johannes M.
    Swedish Museum of Natural History, Department of Paleobiology.
    Grimm, Guido
    Zetter, Reinhard
    Evaluating the mid Miocene paleoclimate of Lower Carinthia (Austria) based on high resolution palynological studies from the Lavanttal Basin2016Conference paper (Other academic)
    Abstract [en]

    Köppen signatures (Denk et al. 2013, Grímsson et al. 2015) can be used to generalize the climatic niche occupied by potential modern analogues (PMA) of fossil plants (here: palynological) assemblages. The Köppen climate system distinguishes climate zones by certain abiotic parameters or combinations thereof and represents them in a three letter code referring to the general climate types (first letter), the seasonal distribution of precipitation (second letter) and the seasonal distribution or general level of warmth (third letter). Based on their Köppen signatures PMAs can be categorized as arctic-alpine, boreal, nemoral, meridio-nemoral, tropical-meridional, tropical, eurytropical, and/or semihumid meridional vegetation elements (see also Denk et al. 2013, Velitzelos et al. 2014, Grímsson et al. 2015). Based on the climatic preferences of their PMAs, the Fagales and Rosales lineages present at the Lavanttal site rule out tropical (A-)climates and climates with pronounced (summer) draught (B-, Cs-, Ds-climates). The same holds for boreal/subarctic climates with short but humid summers (Cfc, Dfc, Dfd, Dwc). The Fagales are represented by 23 lineages at the Lavanttal site including genera that are today composed (predominately or exclusively) of nemoral and meridio-nemoral elements. This points to climate conditions not unlike those found today in the lowlands and adjacent mountain regions of the (south-)eastern United States, the humid-meridional region of western Eurasia (e.g. northern Italy, Black Sea region, western Caucasus), central and southern China, or Honshu (Japan). These regions are characterised by subtropical conditions at lower elevations (Cfa-, Cwa-climates) and subsequent altitudinal successions: Cfa! Cfb/Dfa! Dfb in eastern United States, western Eurasia, central China and Japan, or Cwa! Cwb! Dwb in southern China. The climax vegetation in these areas are mixed mesophytic forests and various mixed evergreen/deciduous broad-leaved forests, characteristic for the humid and semi-humid, summer-rain areas of the meridional and nemoral zone. (Co-)Dominant genera in these forests are the various members of the northern hemispheric Fagales. Important indicator taxa include Fagus , one of the most common and widespread genera in temperate, mixed mesophytic forests of North America, China and Japan, and Quercus Group Ilex, a co-dominant group in the East Asian monsoon influenced, winter-dry or fully humid southern foothills of the Himalayas and montane regions of south-western and central China. Equally informative is Corylus, and the co-occurrence of Carya , Juglans , Pterocarya  and Engelhardioideae, pinpointing towards forests as today found in south-western China and the warm subtropical parts of the southeastern United States.

  • 17.
    Güner, Tuncay H.
    et al.
    Swedish Museum of Natural History, Department of Paleobiology. Faculty of Forestry, Department of Forest Botany, Istanbul University Cerrahpa¸sa, 34473 Bahçeköy, Istanbul, Turkey.
    Bouchal, Johannes M.
    Swedish Museum of Natural History, Department of Paleobiology.
    Köse, Nesibe
    University of Istanbul, Faculty of Forestry, Department of Forest Botany, Istanbul, Turkey.
    Göktaş, Fikret
    Mayda, Serdar
    Denk, Thomas
    Swedish Museum of Natural History, Department of Paleobiology.
    Landscape heterogeneity in the Yatağan Basin (southwestern Turkey) during the middle Miocene inferred from plant macro fossils2017In: Palaeontographica. Abteilung B, Palaophytologie, ISSN 0375-0299, Vol. 296, p. 113-171Article in journal (Refereed)
    Abstract [en]

    Plant macro fossils from the lignite mines of Eskihisar, Tınaz, and Salihpaşalar (Yatağan Basin, southwestern Anatolia) were investigated. The fossils were collected from marls overlying the exploited lignite seams and represent three subbasins within the main Yatağan Basin. The age of the Eskihisar lignite seam is well constrained by vertebrate fossils (MN 6, middle Miocene). Further, lithological and palynological correlation suggests that the lignite seams and overlying marls in the three lignite mines were formed at the same time. Three distinct zonal vegetation types are reflected in the local plant assemblages: (i) In Eskihisar, Fagus and evergreen Quercus mediterranea-Q. sosnowskyi communities formed important parts of the zonal vegetation along with the deciduous Quercus kubinyii; (ii) in Tınaz, Quercus sosnowskyi-Q. mediterranea-Q. drymeja communities occurred, while Fagus is rarely encountered in the macrofossil record. (iii) In Salihpaşalar, Quercus mediterranea and Q. drymeja are the most abundant elements, while Fagus and Q. sosnowskyi are absent or nearly so. This demonstrates that local environmental conditions within a geographically restricted region varied and probably were controlled by slope aspects, edaphic conditions, and river drainage. Overall, the zonal vegetation is characterized by a high diversity of evergreen and deciduous oaks belonging to Quercus subgen. Cerris sect. Ilex and sect, Cerris and the local dominance of Fagus. The riparian vegetation was dominated by Populus, Salix and Acer, whereas Alnus and taxodiaceous Cupressaceae and ferns were very rare or absent. The mass occurrence of Quercus sosnowskyi in the Yatağan Basin floras is biogeographically interesting, as this distinct sclerophyllous oak has previously been known to occur only in late Miocene sediments of northern Greece and Abkhasia (Georgia). 

  • 18. Meller, Barbara
    Grímsson, F.
    Zetter, Reinhard
    Hassler, Andreas
    Hofmann, Christa-Charlotte
    Middle Miocene macrofloral elements from the Lavanttal Basin, Austria, Part I. Ginkgo adiantoides (Unger) HeerArticle in journal (Refereed)
    Abstract [en]

    A new locality, at Schaßbach (Carinthia, Austria), within the Neogene Lavanttal Basin has yielded numerous well preserved early Badenian (Langhian) plant macrofossils. This paper, which is the first in a series of papers that describe the macro-fossil remains from Schaßbach, provides a geological and chronostratigraphic framework of the study area and summarizes previous research on plant macrofossils from the Lavanttal Basin. Here, Cainozoic leaf fossils of Ginkgo with preserved cuticles showing epidermal features are described for the first time from Austria, and from the pre-Pliocene of the Central Paratethys region. The Ginkgo foliage remains are currently one of the oldest Cenozoic fossils representing this genus in Central Europe. The fossils are considered to reflect trees growing outside lowland wetland areas and originated from the riparian vegetation. Based on the current habitat and fossil occurrence of Cainozoic ginkgos, the early Badenian flora in the Lavanttal area likely endured a warm temperate and humid climate.

  • 19.
    Pott, Christian
    et al.
    LWL-Museum of Natural History, Münster, Germany.
    Bouchal, Johannes M.
    Swedish Museum of Natural History, Department of Paleobiology.
    Choo, Thereis Y.S.
    Yousif, Rihab
    Bomfleur, Benjamin
    Westfälische Wilhelms-Universität, Münster, Germany.
    Ferns and fern allies from the Carnian (Upper Triassic) of Lunz am See, Lower Austria: A melting pot of Mesozoic fern vegetation2018In: Palaeontographica. Abteilung B, Palaophytologie, ISSN 0375-0299, Vol. 297, no 1–6, p. 1-101Article in journal (Refereed)
    Abstract [en]

    The floral assemblage from the Carnian (Upper Triassic) of Lunz-am-See in Austria contains a rich and diverse array of ferns and fern allies, most of which have been considered only marginally, some have been described but more than a century ago and the majority have never been illustrated. Therefore, we here provide a detailed analysis of these taxa accompanied by extensive descriptions, detailed illustrations and colour restorations of the plants. We identified 17 species of lycophytes, sphenophytes and pteridophytes assignable to 14 genera in seven families. One taxon could be identified to class level only. One new genus of dipteridacean ferns (viz. Digitopteris C.Pott et Bomfleur gen. nov.), so far monotypic, is here described from Lunz with its type species Digitopteris repanda C.Pott et Bomfleur sp. nov. The Lunz flora comprises an interesting mixture of eusporangiate Marattiales and leptosporangiate Osmundales, Gleicheniales and Schizaeales; the former representing a rather primitive group of ferns. The co-occurrence of eusporangiate and leptosporangiate ferns renders the Lunz flora important given the fact that eusporangiate ferns were predominating over more modern, leptosporangiate forms during the Paleozoic, whereas they typically constitute only subordinate components in fern vegetation since the mid-Mesozoic until today. Among the leptosporangiate forms, the species from Lunz represent, due to their Triassic age, more primitive forms, which probably were rather growth-restricted thriving as herbaceous understorey layer among the shade of the eusporangiate, tree-like marattialean ferns.

  • 20.
    Velitzelos, Demitrios
    et al.
    Athens University.
    Bouchal, Johannes Martin
    Swedish Museum of Natural History, Department of Paleobiology. University of Vienna.
    Denk, Thomas
    Swedish Museum of Natural History, Department of Paleobiology.
    Review of the Cenozoic floras and vegetation of Greece2014In: Review of Palaeobotany and Palynology, ISSN 0034-6667, Vol. 204, p. 56-117Article in journal (Refereed)
    Abstract [en]

    Oligocene to Pleistocene floras of Greece are reviewed based on published and unpublishedmaterial. Oldest plant-bearing sediments of Rupelian–Chattian age are exposed in eastern Thrace (Evros) and were deposited after the closure of the Turgai Seaway. They contain a blend of (i) taxa that migrated to western Eurasia from the East (Alnus, Fagus), (ii) characteristic Oligocene taxa (Nyssa altenburgensis, Ampelopsis hibschii), and (iii) extinct (Eotrigonobalanus, Quasisequoia) and modern genera (Calocedrus, Quercus Group Lobatae) from older epochs. Coastal palmswamps and laurel forests of the hinterland indicate a subtropical, fully humid to winter-dry climate (Cfa, Cwa according to Köppen). The Aquitanian–Burdigalian plant assemblage of Lesbos is intermediary between Evros and the Burdigalian floras of Euboea sharing taxa with Evros (palms), and with Euboea and early Miocene floras of Anatolia (Güvem, Tilia). In the early Miocene (Burdigalian) floras of Euboea, species of Quercus Group Ilex (Quercus drymeja, Quercus mediterranea) characteristic of fully humid or winter-dry (monsoon) climates (Cf, Cw) became dominant elements in well-drained forests. Floristic links are with late Oligocene to middle Miocene floras of Central Asia (Tilia), AsiaMinor (cycads, Quercus Group Ilex, Tilia), and South and Central Europe (cycads, Quercus Group Ilex, palms). Middle Miocene floras are restricted to the Aegean Islands (Chios). Biogeographic links arewith early to lateMiocene floras of Central Europe (Parrotia, Podocarpium) andwithmiddle Miocene floras of Anatolia (Parrotia). UpperMiocene plant-bearing sedimentary formations aremost abundant in Greece and exposed on the Ionian Islands, Greece mainland to East Macedonia, Peloponnese, Aegean Islands, and Crete. Overall, the fossil plant assemblages from Greece mainland are indicative of fully humid conditions during this time (Cfa), with Fagus and oaks of Quercus Group Ilex being dominant elements. Seasonality may have been more pronounced on the Peloponnese and the Aegaean Islands and Crete, expressed by the rare occurrence of Fagus in the fossil records of these areas. The palaeobotanical records from Samos unambiguously point to the presence of forest vegetation during early Tortonian to Messinian (Cwa) when the famous vertebrate faunas of Samos were deposited. The Pliocene is characterized by the regional occurrence of modern types of deciduous oaks mainly of Quercus Group Cerris and Quercus subsect. Galliferae. East Asian links persist in Fagus, Quercus, and Cupressaceae,North American ones in Sabal; several othermesophytic taxa fromprevious periods are recorded as well. The modern sclerophyllous Mediterranean vegetation, thriving in a warm summer dry climate (Csa), cannot be traced prior to the Pleistocene based on the palaeobotanical record.

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