The Miocene flora of Parschlug (Styria, Austria)revision and synthesis

Johanna  Eder

The macroflora of Rauenberg, Baden-Württemberg, Germany, is treated monographically. The plant-bearing sediments are marine, mainly well-bedded clay-to siltstones, the so-called Fischschiefer, which are part of the Bodenheim Formation. Based on nannoplankton they are dated to nannoplankton zone NP 23 (Rupelian, Lower Oligocene). The plant remains, mainly leaves and some fructifications, are preserved as compressions. The taxonomic assignment is based on gross morphology and cuticle characteristics. The flora yields marine algae and remains of the very diverse terrestrial flora. A total of 68 taxa, including three types of algae, one cycad, 12 conifers, and 49 dicots, among them 5 palms, are described. The following fossil species are described for the first time: Laurophyllum rauenbergense, Myrica obliquifolia, Distylium metzleri, ? Berchemia altorhenana, ? Ternstroemites maritiae, Trachelospermum kelleri, Oleinites altorhenana, O. rauen-bergensis, Dicotylophyllum badense, D. oechsleri, D. vesiculaeferens, D. ziegleri, ? Viscophyllum hendriksiae, and Cladites vesiculaeferens. Dicotylophyllum vesiculaeferens and Cladites vesiculaeferens bear peculiar, complex cuticular structures presumably representing salt-secreting glands. Both taxa are interpreted to derive from one plant species of yet uncertain systematic affinity. The flora bears a high proportion of broad-leaved, presumably evergreen taxa, whereas the diversity of modern Arcto-Tertiary taxa (sensu Kvaček 1994) is rather low. Most abundant are Platanus neptuni, Daphnogene cinnamomifolia, and Tetraclinis salicornioides. On the family level, Lauraceae (10 species) and Pinaceae (8) are most diverse, followed by Arecaceae (4–5), Cupressaceae, and Myricaceae (4 species each). Surprisingly, Fagaceae are documented solely by a single leaf of Eotrigonobalanus furcinervis f. haselbachensis, and the record of Pentaphyllaceae remains ambiguous (? Ternstroemites maritiae). Sloanea olmediaefolia is recorded for the first time from western parts of Europe. Remarkable is the presence of the rare cycad Ceratozamia floersheimensis. The following possible vegetation units are suggested: zonal broad-leaved sclerophyllous evergreen forests and an intrazonal coastal pine-laurel-palm association on near-coastal sandy soils, as well as gallery forests along streams. No records of swampy environments were recovered. The climate may be characterised as follows: Cfa climate in transition to Cwa (and Am or Af) climate sensu Köppen, mean annual temperature 19–24°C, mean annual precipitation 1300–1700 mm, mean temperature of the warm-est month 28–29°C, mean temperature of the coldest month 8–14°C, mean precipitation of the wettest month >230 mm, mean precipitation of the driest month 18–38 mm, wettest month between May and October, driest month between November and March. The warm period was the wetter one. The flora from Rauenberg most closely resembles that of Flörsheim (Kvaček 2004a) and shows relations to the Paratethys realm, for example the Tard Clay Formation. Relations to the floras from Saxony, Saxony-Anhalt, and North Bohemia, similar in age, are rather restricted: broad-leaved deciduous taxa are much less diverse, and the numerous presumably evergreen taxa and palms present in Rauenberg have not been recorded in the other regions, indicating a more complex vegetation differentiation than a simple north-south gradient. The high number of taxa of uncertain affinity at Rauenberg points to the need for further taxonomic studies of the flora of this time interval. Comparisons with European assemblages of the early Oligocene reveal that the vegetation diversity in Europe during this time interval is far from being well understood.

New floristic records from the eastern Karavanke/Karawanken and Kamniske Alpe/Steiner Alpen (Slovenia and Austria) are reported. Allium kermesinum is new for Austria; Arabis soyeri subsp. subcoriacea, Carex rupestris and Draba dubia are new for the Kamniske Alpe/Steiner Alpen; for Androsace hausmannii, Arabis stellulata, Carex ornithopodoides, Pedicularis rosea, Salix serpillifolia and Veronica fruticulosa new localities are presented. Furthermore, taxonomic problems in Oxytropis sect. Oxytropis and Arabis pumila sensu lato are discussed. l

The flora from Oberdorf (Styria, Austria, Miocene, Ottnangian), which was recently investigated taxonomically and flonstically, is here evaluated with the Coexistence Approach (CA). Using 127 relevant taxa, the obtained coexistence intervals of the mean annual temperature and mean annual precipitation fall into the ranges proposed by MELLER (1998) and MELLER & al. (1999), based on sociological comparisons of the fossil Oberdorf and the modern plant record. The ranges based on the CA are distinctly narrower with 15.7-17.6°C mean annual temperature and 1187-1322 mm mean annual precipitation; this more precisely defines the humid warm-temperate Cfa-climate.

Ann. Naturhist. Mus. Wien 105 A 45–159 Wien, Februar 2004 GEOLOGIE UND PALÄONTOLOGIE The Miocene Flora of Parschlug (Styria, Austria) – Revision and Synthesis By Johanna KOVAR-EDER1, Zlatko KVACEK2 & Margit STRÖBITZER-HERMANN3 (With 5 figures, 11 tables and 15 plates) Manuscript submitted on 23 October 2002, the revised manuscript on 21 January 2003 Abstract The first monographic treatment of the famous fossil flora of Parschlug (Styria, Austria) is presented. It comprises more than 60 plant species including 4 ferns, 5 conifers, and over 50 angiosperms. Described for the first time are Ulmus parschlugiana and Antholithes stiriacus. Newly combined are Berberis teutonica, B. (?) ambigua, Mahonia (?) aspera, Ternstroemites pereger, Cedrelospermum ulmifolium, Leguminosites hesperidum, L. dionysi, L. palaeogaeus, L. parschlugianus, Prinsepia serra, Cotinus (?) aizoon, and Ailanthus pythii. Diversified mesophytic elements prevail over a few dominant or common azonal woody taxa. Among the former, humid temperate components are relatively scarce and humid subtropical ones are rare, while subhumid, physiognomically sclerophyllous woody taxa are well represented. The age is considered as Karpatian/Early Badenian (late Early/early Middle Miocene) based on the floristic composition. Climatically this association indicates a drier warm-temperate/subtropical regime than documented from earlier and later Miocene times. Keywords: Macroflora, palaeoecology, palaeoclimate, floristic comparison, Miocene, Norian depression, Austria. Zusammenfassung Erstmals wird die Flora von Parschlug (Steiermark, Österreich) monographisch erfasst. Sie enthält mehr als 60 Pflanzenarten, davon 4 Farne, 5 Koniferen und mehr als 50 Angiospermen. Ulmus parschlugiana and Antholithes stiriacus werden erstmals beschrieben. Neu kombiniert werden Berberis teutonica, B. (?) ambigua, Mahonia (?) aspera, Ternstroemites pereger, Cedrelospermum ulmifolium, Leguminosites hesperidum, L. dionysi, L. palaeogaeus, L. parschlugianus, Prinsepia serra, Cotinus (?) aizoon, und Ailanthus pythii. Außer einigen dominierenden oder häufigen azonalen Gehölzen herrschen mesophytische Elemente vor. Unter diesen sind humid temperate nicht häufig und humid subtropische sogar selten. Aber subhumide, physiognomisch sklerophylle Gehölze sind reichlich vertreten. Basierend auf der floristischen Zusammensetzung wird ein karpatisch/unter-badenisches Alter (oberes Unter-/unteres Mittel-Miozän) angenommen. Im Vergleich mit älteren und jüngeren miozänen Floren deutet die Vergesellschaftung von Parschlug auf relativ trockenere warm-temperat/subtropische klimatische Verhältnisse hin. Schlüsselwörter: Makroflora, Paläoökologie, Paläoklima, floristische Vergleiche, Miozän, Norische Senke, Österreich. Johanna KOVAR-EDER, Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, D-70191 Stuttgart, Germany; e-mail: eder.smns@naturkundemuseum-bw.de. 2 Zlatko KVACEK, Charles University, Faculty of Science, Albertov 6, CZ-128 43 Praha 2, Czech Republic; e-mail: kvacek@natur.cuni.cz. 3 Margit STRÖBITZER-HERMANN, Geologisch-Paläontologische Abteilung, Naturhistorisches Museum, Burgring 7, P.B. 417, A-1014 Wien, Austria; e-mail: margit.stroebitzer@nhm-wien.ac.at. 1 46 Annalen des Naturhistorischen Museums in Wien 105 A Table of contents Introduction ................................................................................................................................................. 48 Geography and geological frame ................................................................................................................ 49 Material and methods .................................................................................................................................. 50 Systematics .................................................................................................................................................. 52 Osmunda parschlugiana (UNGER) ANDREÁNSZKY ...................................................................................... 52 Pronephrium stiriacum (UNGER) KNOBLOCH & Z. KVACEK ....................................................................... 52 Adiantum renatum UNGER .......................................................................................................................... 52 Salvinia cf. mildeana GOEPPERT .................................................................................................................. 53 Pinus sp. div. ............................................................................................................................................... 53 ? Cathaya sp. ............................................................................................................................................... 54 Glyptostrobus europaeus (BRONGNIART) UNGER ........................................................................................ 54 ? Cupressus sp. ........................................................................................................................................... 55 Daphnogene polymorpha (A. BRAUN) ETTINGSHAUSEN ............................................................................. 55 Berberis teutonica (UNGER) KOVAR-EDER & Z. KVACEK comb. nov. ........................................................ 56 Berberis (?) ambigua (UNGER) KOVAR-EDER & Z. KVACEK comb. nov. ................................................... 56 Mahonia (?) aspera (UNGER) KOVAR-EDER & Z. KVACEK comb. nov. ..................................................... 57 Cercidiphyllum crenatum (UNGER) R. BROWN ........................................................................................... 57 Liquidambar europaea A. BRAUN .............................................................................................................. 58 Liquidambar sp. – fructus ........................................................................................................................... 58 Platanus leucophylla (UNGER) KNOBLOCH ................................................................................................. 58 Betula cf. dryadum BRONGNIART ................................................................................................................ 59 Betula vel Alnus sp. .................................................................................................................................... 59 Alnus julianiformis (STERNB.) Z. KVACEK & HOLY .................................................................................... 59 Alnus gaudinii (HEER) KNOBLOCH & Z. KVACEK ....................................................................................... 60 Fagus sp. – leaf ........................................................................................................................................... 60 Fagus sp. – cupule ...................................................................................................................................... 60 Fagus vel Alnus sp. ..................................................................................................................................... 61 Quercus drymeja UNGER ............................................................................................................................. 61 Quercus mediterranea UNGER .................................................................................................................... 62 Quercus zoroastri UNGER ........................................................................................................................... 62 cf. ? Gordonia oberdorfensis KOVAR-EDER ................................................................................................ 63 Ternstroemites pereger (UNGER) KOVAR-EDER & Z. KVACEK comb. nov. ................................................ 63 Myrica lignitum (UNGER) SAPORTA ............................................................................................................. 64 Myrica oehningensis (A. BRAUN) HEER ...................................................................................................... 65 Myrica sp. – fructus .................................................................................................................................... 65 Engelhardia orsbergensis (WESSEL & WEBER) JÄHNICHEN, MAI & WALTHER .......................................... 65 Engelhardia macroptera (BRONGNIART) UNGER ......................................................................................... 65 Tilia longebracteata ANDRAE ..................................................................................................................... 66 Craigia bronnii (UNGER) Z. KVACEK, BŮZEK & MANCHESTER .................................................................. 66 Ulmus plurinervia UNGER ........................................................................................................................... 66 Ulmus parschlugiana KOVAR-EDER & Z. KVACEK sp. nov. ....................................................................... 67 Cedrelospermum ulmifolium (UNGER) KOVAR-EDER & Z. KVACEK comb. nov. – foliage ........................ 68 Cedrelospermum stiriacum (ETTINGSHAUSEN) KOVAR-EDER & Z. KVACEK comb. nov. – fructus ............ 68 Zelkova zelkovifolia (UNGER) BŮZEK & KOTLABA ...................................................................................... 69 Celtis japeti UNGER ..................................................................................................................................... 70 Populus populina (BRONGNIART) KNOBLOCH .............................................................................................. 70 Populus sp. – fructus ................................................................................................................................... 70 Buxus cf. egeriana Z. KVACEK, BŮZEK & HOLY ........................................................................................ 71 cf. Rosa sp. .................................................................................................................................................. 71 Prinsepia serra (UNGER) KOVAR-EDER & Z. KVACEK comb. nov. ............................................................ 72 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 47 ? Prinsepia sp. ............................................................................................................................................ 73 Leguminosites hesperidum (UNGER) KOVAR-EDER & Z. KVACEK comb. nov. ........................................... 73 Leguminosites dionysi (UNGER) KOVAR-EDER & Z. KVACEK comb. nov. .................................................. 74 Leguminosites palaeogaea (UNGER) KOVAR-EDER & Z. KVACEK comb. nov. ........................................... 74 Leguminosites parschlugianus (UNGER) KOVAR-EDER & Z. KVACEK comb. nov. ..................................... 74 Podocarpium podocarpum (A. BRAUN) HERENDEEN .................................................................................. 74 Phaseolites securidacus UNGER .................................................................................................................. 75 "Acacia" parschlugiana UNGER .................................................................................................................. 75 "Juglans" parschlugiana UNGER ................................................................................................................. 75 Paliurus tiliifolius (UNGER) BŮZEK ............................................................................................................. 76 Paliurus favonii UNGER .............................................................................................................................. 76 Berchemia multinervis (A. BRAUN) HEER ................................................................................................... 77 Acer tricuspidatum BRONN ......................................................................................................................... 77 Acer pseudomonspessulanum UNGER emend. STRÖBITZER-HERMANN ....................................................... 77 Acer integrilobum WEBER sensu WALTHER ................................................................................................ 78 Acer sp. div. – fructus ................................................................................................................................. 79 Toxicodendron herthae (UNGER) Z. KVACEK & WALTHER ........................................................................ 80 Cotinus (?) aizoon (UNGER) KOVAR-EDER & Z. KVACEK comb. nov. ........................................................ 80 Ailanthus pythii (UNGER) KOVAR-EDER & Z. KVACEK comb. nov. ............................................................ 81 Ailanthus confucii UNGER ........................................................................................................................... 82 Fraxinus primigenia UNGER ....................................................................................................................... 82 Nerium sp. ................................................................................................................................................... 82 Smilax sagittifera HEER emend. HANTKE .................................................................................................... 83 Monocotyledoneae gen. et sp. indet. .......................................................................................................... 84 "Celastrus" europaea UNGER ...................................................................................................................... 84 "Cornus" ferox UNGER ................................................................................................................................ 84 "Evonymus" latoniae UNGER ....................................................................................................................... 85 "Quercus" daphnes UNGER ......................................................................................................................... 85 Antholithes stiriacus KOVAR-EDER & Z. KVACEK sp. nov. ........................................................................ 86 Cypselites sp. ............................................................................................................................................... 86 Saportaspermum sp. .................................................................................................................................... 87 ? Chaneya sp. .............................................................................................................................................. 87 Dicotylophyllum sp. 1 ................................................................................................................................. 88 Dicotylophyllum sp. 2 ................................................................................................................................. 88 Dicotylophyllum sp. 3 ................................................................................................................................. 88 Dicotylophyllum sp. 4 ................................................................................................................................. 89 Dicotylophyllum sp. 5 ................................................................................................................................. 89 Dicotylophyllum sp. 6 ................................................................................................................................. 89 Dicotylophyllum sp. div. ............................................................................................................................. 89 Taphonomy ................................................................................................................................................. 92 Palaeoecology, sociology, and climate ....................................................................................................... 92 Parschlug in the context of other floras along the Norian depression ....................................................... 95 The flora of Parschlug in Central European context .................................................................................. 96 Southern Germany (Randeck Maar and Upper Freshwater Molasse) ........................................................ 96 Cypris Clay flora, western Bohemia ........................................................................................................... 98 Miocene of Hungary ................................................................................................................................. 101 Miocene of Greece .................................................................................................................................... 101 Age of the flora of Parschlug .................................................................................................................... 117 Acknowledgements ................................................................................................................................... 117 References ................................................................................................................................................. 118 Index of species ........................................................................................................................................ 125 48 Annalen des Naturhistorischen Museums in Wien 105 A Fig. 1: Geographical and geological position of Parschlug. Background geological map from OBERHAUSER (1980). Introduction The Miocene flora of Parschlug in southeastern Austria is famous thanks to the numerous species currently used in the palaeobotanical literature that are based on it. Most of them were described by F. UNGER and, to a lesser extent, by C. v. ETTINGSHAUSEN. The descriptions and illustrations of the material including the types are scattered over 13 publications edited in the 19th century. However, the flora of Parschlug was never treated monographically because ETTINGSHAUSEN (1878 b) was unable to finish his initiated study intended for such a monograph. Plant fossils from Parschlug are located in many European collections, of which the most extensive are certainly those housed in the Natural History Museum and the Geological Survey, both in Vienna, and the Botanical Institute of the Karl-FranzensUniversity and the Landesmuseum Joanneum, both in Graz. Rich fossil plant material was collected during the times of mining activity in this area, which started in the early 1800s and ended in 1959. We have studied all the mentioned collections as well as the collection at the Montan-University in Leoben and some others in Germany and Hungary. Because of immense number of samples collected at Parschlug, we were only able to examine the most important parts of the collections and have certainly overlooked some interesting, rare fossils. We took advantage of the database called "Palaeontological Types in Austrian Collections" http://www.oeaw.ac.at/oetyp/palhome.htm), which enabled us to locate the preserved type specimens and originals. As a result we present here a monograph including various revisions of elements occurring in the flora of Parschlug hoping that this will be of use for other students of Tertiary palaeobotany. Because of known difficulties with identifying foliage, which in the case of Parschlug is devoid of useful epidermal anatomy, not all entities have been assigned to the natural system and many problems remain to be resolved in future studies. We believe that our investigations are a good starting point for such an endeavour. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 49 Fig. 2: The Parschlug coal basin in the Norian depression (simplified from SACHSENHOFER et al. 2002: fig. 1). In our investigations the genus Acer was prepared by M. STRÖBITZER-HERMANN, all other taxa jointly by the other two authors. Geography and geological frame The Parschlug basin is situated about 5 km north of Kapfenberg in Styria – ca. 15°17´ E longitude / 47°28´ N latitude according to Austrian Map, map OEK 1:50 000, sheet 133 (fig.1). As a consequence of the Miocene lateral extrusion of the Eastern Alps, the Mur / Mürz fault system – known also as the Norian depression – developed between the eastern Alpine margin and the later Tauern window. This yielded several coal-bearing pull-apart basins and half-grabens between Hart / Gloggnitz and Tamsweg. The Parschlug basin is an eastern one situated in the lower Mürz valley; it is classified by NEUBAUER et al. (2000) as a pull-apart basin (fig. 2). Contrary to other basins along the Norian depression (e.g. the Fohnsdorf basin) the Parschlug basin was never in focus of geological investigations. The available information is largely based on data given by UNGER (1848), PETRASCHEK (1922-1929), WEBER & WEISS (1983), and most recently SACHSENHOFER (in SACHSENHOFER et al. 2002). In the Parschlug basin, sands and sandstones overlie the basement. A 4-8 m thick coal seam ("Parschlug seam") follows in the section. At the western margin of the basin the seam dips about 45° towards east, while the dipping decreases towards the centre to 12°. Step faults running NNW/SSE and dipping towards NW cut the seam in several fault 50 Annalen des Naturhistorischen Museums in Wien 105 A blocks (German: Schollen). Towards the east and northwest the seam thickness decreases, and the seam splits up and wedges out. Therefore, mining activity was focussed formerly on the southwestern part of the depression. Clays and marls with up to 10 cm thick marlstone-ironstone intercalations (German: Toneisensteinbänke) overlie the seam. At certain levels they bear plant remains known as the famous flora of Parschlug. UNGER (1848) figured and described a profile taken in 1843 at "dem über dem Fabriksgebäude befindlichen Stollen des Graf´schen Bergbaus", which reflects the local situation at that time (fig. 3). The ash content of the coal is relatively high (up to 40 %). The sulphur content of 4 to 7 % and the preservation of gastropod shells (Planorbis aplanatis) in an inter-seam indicate a relatively high pH-value (around 7) of the mire. Material and Methods ETTINGSHAUSEN, in his old catalogue (housed at the Institute of Botany, University of Graz), recognised three kinds of fossiliferous rocks and distinguished 3 fossiliferous levels in his collections: I – "Weicher Mergelschiefer" (whitish, thin-bedded soft marl), II – "Harter gelber Mergelschiefer" (hard, light brown, often reddish marlstone) and III – "Harter hellgrauer Mergelschiefer" (very hard, light grey, also often reddish marlstone to ironstone). Most of the plant fossils studied are preserved as dark compressions / impressions in reddish ironstone (levels II and III). They are partly covered with fossilised tissue but our attempts to prepare leaf cuticles mostly failed. Most of the carbonised mass bears traces of pyrite. Moreover, venation patterns are only poorly visible. The preservation of venation details on light yellow-brownish impressions from ETTINGSHAUSEN’s level I is often more satisfactory. Massive fruit remains, e.g. of Liquidambar, which occur in ironstone beds have been often destroyed by pyritisation. Before decomposition, they were probably more common in the collections. The material investigated in this revision is recognisable by collection file numbers. The quantity of coll. file nos listed under a taxon does not necessarily reflect its true abundance in the Parschlug flora because not all material is numbered and not all numbers of the specimens studied are included in the text. In the systematic part we restrict ourselves to complementing the descriptions given by UNGER and ETTINGSHAUSEN with diagnostic features that they were unaware of or misinterpreted, and we give full descriptions in the cases of newly characterised taxa. Only the published records from Parschlug are included into synonym lists under the respective taxa. The following abbreviations are used throughout the text to designate the respective collections: GBA Geologische Bundesanstalt, Wien IBUG Institut für Botanik der Karl-Franzens-Universität, Graz LMJ Landesmuseum Joanneum, Graz MMG Staatliche Naturhistorische Sammlungen Dresden, Museum für Mineralogie und Geologie Dresden NHMW Naturhistorisches Museum, Wien KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 51 Fig. 3: Geological section of Parschlug at the gallery of the mine in 1843 (according to UNGER 1848: 7, adapted and translated into English). 1- underlying quartzose fine-grained sandstone; 2 - ca. 1.9 m ("1 Klafter") black, not pure clayey coal with 2.5-5 cm thick inter-beds of sand in the lower part and dark grey claystone highepure coal 5-7.5 cm thick only; 3 - ca. 21 cm ("8 Zoll") dark brown hard rock with fragments of molluscs; 4 - up to ca. 65 cm ("2 Fuss") pure coal; 5 - ca 2.8 m ("1.5 Klafter") claystone; 6 - a thin inter-bed of hard rock; 7 - ca 1.9 m ("1 Klafter") marlstone; 8 - ca. 0.9 m ("3 Fuss") black brown coal; 9 - a thin inter-bed of fuller’s earth; 10 - ca. 2.1 m ("7 Fuss") almost pure pitch coal and slate coal; 11 - ? thickness - coal and marl transition above the coal seam into dark brown finegrained mica claystone with leaf impressions, easily weathered out; 12 - ca. 2.1 m ("7 Fuss") soft grey claystone; 13 - ca. 13 cm ("5 Zoll") harder broken up claystone to ironstone with best preserved plant impressions; 14 - more than 10 m ("mehrere Klafter") yellowish marlstone to marl with transition into the following; 15 – earthwork. 52 Annalen des Naturhistorischen Museums in Wien 105 A Systematics Pteridophyta Osmundaceae Osmunda L. Osmunda parschlugiana (UNGER) ANDREÁNSZKY Plate 1, Figures 1, 2 1847 Pteris parschlugiana UNGER, p. 122, pl. 36, fig. 6 (LMJ 76520, holotype). Holotype: LMJ 76520, figured by UNGER (1847: 122, pl. 36, fig. 6) - refigured on pl. 1, fig. 1. Additional material: NHMW 1878/6/6795. Detached pinnules, almost parallel-sided, 8-11 mm wide and more than 40 mm long, finely crenulate, semicordate. Secondary veins simple or forked, parallel, dense, originating under the angle of 45-55°. The attachment to the rhachis is damaged, contrary to UNGER’s figure. Actually we doubt that the axis below the pinnule is in fact the rhachis. The holotype pinnule corresponds exactly to the numerous specimens described in detail from the Early Miocene Most Formation by BŮZEK (1971). The semicordate base in combination with fine marginal crenulations distinguishes this fern from similar pinnules of Blechnum dentatum (see KVACEK & HURNÍK 2000: 6). The affinities to extant species of Osmunda must remain open pending a detailed study of co-occurring spores. These can be specific for different species (TRYON & LUGARDON 1990), while leaf morphological features of the Osmunda regalis-type are partly not distinctive enough to help in this respect. Thelypteridaceae Pronephrium K. PRESL CEK Pronephrium stiriacum (UNGER) KNOBLOCH & Z. KVAC Plate 1, Figure 3 Material: IBUG: Ett. coll. 111 (level III) This poorly preserved fern fragment shows the goniopterid venation and shares other morphological features, as shape and size, with other records of this species (cf. e.g. KVACEK & HURNÍK 2000). Pteridaceae Adiantum L. Adiantum renatum UNGER Plate 1, Figure 5 1847 Adiantum renatum UNGER, p. 122, pro parte, pl. 37, fig. 1. 1850a Adiantites renatus (UNGER) UNGER, p. 106. Material: IBUG: Ett. coll. 344 (level II) KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 53 A small fan-shaped leaf with flabellate venation, about 10 mm long including a short petiole. Judging from the published illustration of the missing holotype in UNGER (1847), we suspect this poorly preserved specimen to be conspecific. A similar fern occurs also at the Randeck Maar (GREGOR 1986). Salviniaceae Salvinia SÉGUIER Salvinia cf. mildeana GOEPPERT Plate 1, Figure 4 Material: IBUG: Ett. coll. 112 + 113 (part + counterpart, level II). Isolated floating leaves, elliptical, ca. 10 x 15 mm, showing a primary vein giving off straight secondaries under the angle of 90-45° and oblique cross veins forming oblique quadrangular meshes. Leaf surface typically tuberculate. The relatively small size of the specimen led us to compare the material from Parschlug with S. mildeana, which is typified by similar specimens from the Late Miocene site Sośnica. According to the recent revision (COLLINSON et al. 2001), not only size differences exist between this species and its probable ancestor, S. reussii (Late Oligocene to Early Miocene). At its type locality S. mildeana is accompanied by megaspores of the S. intermedia-type, while S. reussii is accompanied by those of the S. cerebrata-type. No megapores have been found at Parschlug and the size of floating leaves alone is an insufficiently reliable diagnostic character in view of the fact that the material from Parschlug is limited to a single specimen and its counter-impression. We found several specimens at IBUG identified as Salvinia microphylla ETTINGSHAUSEN, which are partly dubious plant remains (IBUG Ett. coll. 114) or wing-cases of beetles (IBUG Ett. coll. 115 et 116 level III). Gymnospermae Pinaceae Pinus L. Pinus sp. div. Plate 1, Figures 6-13 1850a 1852 1850a 1852 1850a 1852 1850a 1852 1850a 1852 1850a Pinites balsamodes UNGER, p. 357. Pinites balsamodes UNGER – UNGER, p. 95, pl. 35, fig. 7 (LMJ 76496, syntype), fig. 8. Pinites centrotos UNGER, p. 362. Pinites centrotos UNGER – UNGER, p. 98, pl. 37, fig. 1 (LMJ 76486, syntype), figs. 2, 3, 4 (LMJ 76500, syntype). Pinites furcatus UNGER, p. 363. Pinites furcatus UNGER - UNGER, p. 99, pl. 37, figs. 7-9. Pinites goethanus UNGER, p. 361. Pinites goethanus UNGER – UNGER, p. 96, pl. 35, fig. 18 (LMJ 76491, syntype), figs. 19-22. Pinites hepios UNGER, p. 362. Pinites hepios UNGER – UNGER, p. 97, pl. 35, figs. 6-8, 9 (LMJ 76501, syntype). Pinites leuce UNGER, p. 358. 54 Annalen des Naturhistorischen Museums in Wien 105 A 1852 1850a 1852 1852 1878a 1878a 1878a 1878a 1878a 1878a 1878a 1878a 1878a Pinites leuce UNGER – UNGER, p. 95, pl. 35, figs. 9-16. Pinites oceanines UNGER, p. 357. Pinites oceanines UNGER – UNGER, p. 94, pl. 35, figs. 1-4. Pinites taedaeformis UNGER, p. 97, pl. 36, fig. 4. Pinus hepios (UNGER) HEER - ETTINGSHAUSEN, p. 74, pl. 7, figs. 12, 13. Pinus laricio POIR. - ETTINGSHAUSEN, p. 75, pl. 7, 5, 6. Pinus palaeo-strobus (ETTINGSHAUSEN) ETTINGSHAUSEN , p. 74, pl. 1 figs. 1 b (NHMW 1878/6/9689), 2 b (NHMW 1878/6/9690), 3, 4, 7, 11, 12, 15, 16 (NHMW 1878/6/9687). Pinus post-taedaeformis ETTINGSHAUSEN, p. 77, pl. 4, figs. 3-5. Pinus prae-cembra ETTINGSHAUSEN, p. 77, pl. 3, figs. 2, 3. Pinus prae-pumilio ETTINGSHAUSEN, p. 75, pl. 9, figs. 5, 7, 8, pl. 10, fig. 1 a, 15 b (NHMW 1878/6/9765, syntype), 14 b (NHMW 1878/6/9762, syntype). Pinus prae-silvestris ETTINGSHAUSEN, p. 75, 76, pl. 1 figs. 5 (NHMW 1878/6/9744, syntype), 6 (NHMW 1878/6/9745, syntype), pl. 7, figs. 20 (NHMW 1878/6/9746, syntype), pl. 10, fig. 9 (NHMW 1878/6/9743, syntype). Pinus prae-taedaeformis ETTINGSHAUSEN, p. 77, pl. 2, fig. 3 (NHMW 1878/6/9779, syntype). Pinus rigios UNGER - ETTINGSHAUSEN, p. 79, pl. 4, fig. 6 (NHMW 1878/6/9797). Additional material: GBA 2002/01/26; IBUG Ett. coll.195; NHMW 1878/6/2479, 9706, 9780. The material, which comprises various kinds of dispersed male cones, seed cone scales, seeds and foliage, but not complete seed cones, does not seem to be identifiable to the species level. We refrain from establishing various morpho-taxa and we restrict ourselves to reproducing different morpho-types of these organs and do not attempt to differentiate them in detail. In our opinion, the number of natural species is certainly lower than the binomina listed in the synonymy. The single cone scale studied shows only poorly preserved details of the umbo. MAI (1986, 1994) has not recognised any entity listed above in his survey of Tertiary pines of Europe. ? Cathaya W.Y. CHUN & K.Z. KUANG ? Cathaya sp. Plate 1, Figures 20-23 Material: IBUG: Ett. coll. 317, 318, 335, 343, 6977 ( level II); NHMW 1878/6/9684. Several small cone scales rounded-rhomboidal, about 8-9 mm wide and ca. 10 mm long, obviously slightly convex before fossilisation, with hairy periphery and striate surface. One detached flat needle (2 x 60 mm in size) with rounded apex and slightly enlarged base. The shape of the cone scales matches well with those of a disintegrated seed cone of Cathaya. The bract has not been preserved in any of the specimens studied. The extant species endemic to China exceeds in size of cone scales the fossils, which are smaller, like the specimens from the Upper Miocene of Santa Barbara (MAI 1994). ETTINGSHAUSEN (in his catalogue, IBUG) suspected these remains to be allied to the Pinaceae, designating them as Pinus ciliata. The needle (IBUG Ett. coll. 343) may belong to the same plant. Cupressaceae sensu lato Glyptostrobus ENDL. Glyptostrobus europaeus (BRONGNIART) UNGER Plate 1, Figures 14-16 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 1845 1847 1847 1850a 1850a 1850a 1852 1852 1968 55 Juniperites baccifera UNGER, p. 80, pl. 21, figs.1 (NHMW 2001B0017/0001, syntype), 2. Widdringtonites ungeri ENDLICHER - UNGER, p. 7 (271), pro parte. Taxodites oeningensis ENDLICHER - UNGER, p. 15 (279), pro parte. Widdringtonites ungeri ENDLICHER - UNGER, p. 342. Taxodites dubius STERNBERG - UNGER, p. 351, pro parte. Taxodites oeningensis ENDLICHER -UNGER, p. 351, pro parte. Glyptostrobus oeningensis A. BRAUN - UNGER, p. 20. Taxodites dubius STERNBERG - UNGER, p. 20, pro parte. Widdringtonia baccifera (UNGER) KNOBLOCH, p. 126. Additional material: IBUG Ett. coll. 162 (level II), seed cones, 190a twigs det. by ETTINGSHAUSEN as Taxodium distichum miocenicum; NHMW 1878/6/2658a. Twigs with helically disposed and appressed scale leaves are rather common, contrary to the characteristic seed cones, which are rare probably due to decay through pyritisation. Rare are twigs with taxodioid foliage (of young shoots), which were determined by ETTINGSHAUSEN (in IBUG) as Taxodium distichum miocenicum. Besides seed cones, pollen cones attached terminally on the twigs also occur. The latter were misinterpreted as juniper-like seed cones (UNGER 1845, as Juniperites baccifera). ? Cupressus L. ? Cupressus sp. Plate 1, Figures 17-19 Material: GBA 2002/01/23-25; NHMW 1878/6/2554 b, 1845/0039/0003. Delicate twigs, straight, 1-1.5 mm thick and widely branched under an angle of about 45°, densely covered by almost isomorphic, decussately disposed scale leaves, broadly trigonal-deltoidal, blunt, with an indistinct gland on the facial leaves. These specimens discovered in the collections at GBA and NHMW in Vienna were all incorrectly determined by UNGER and ETTINGSHAUSEN as Widdringtonia ungeri or Juniperites baccifera, respectively, which are synonyms of Glyptostrobus europaeus. Similar foliage accompanies Cupressus seed cones at the Early Miocene site of Kymi and Late Miocene of Vegora (KVACEK et al. 2002b). Angiospermae Lauraceae Daphnogene UNGER Daphnogene polymorpha (A. BRAUN) ETTINGSHAUSEN Plate 2, Figure 8 1847 Ceanothus subrotundus A. BRAUN – UNGER, p. 144, pl. 49, fig. 7 (LMJ 76530). Additional material: GBA 2002/01/116; NHMW 2001B0017/0002. Cinnamomoid leaves are extremely rare at Parschlug, being represented mostly by broader forms typical of Miocene populations. The designation of foliage is maintained under this morpho-taxon for pragmatic reasons, although the associated fruits at Kreuzau are apparently allied to a camphor tree (PINGEN et al. 1994). 56 Annalen des Naturhistorischen Museums in Wien 105 A Berberidaceae Berberis L. CEK comb. nov. Berberis teutonica (UNGER) KOVAR-EDER & Z. KVAC Plate 2, Figures 9, 10 1850a Clethra teutonica UNGER, Gen. spec. pl. foss., p. 439 - basionym. 1866 Crataegus teutonica (UNGER) UNGER, p. 60, pl. 19, figs. 24, 25. Neotype designated here: 1878/6/2153 det. by ETTINGSHAUSEN as Celastrus sp. nov. - figured on pl. 2, fig. 9. Additional material: NHMW 1878/6/2442 + 9372 det. by ETTINGSHAUSEN as Quercus myrsinaefolia and Quercus mediterranea, respectively. Obovate subsessile leaves inconspicuously widely toothed, differing from similar Berberis berberidifolia (HEER) PALAMAREV & PETKOVA by a shorter petiole and denser secondaries, sharing irregular pinnate semicraspedodromous venation, which forms several rows of loops along the margin. The figures published by UNGER (1866) show two incomplete leaves of different size. Both differ slightly from the neotype by the narrowly and shortly decurrent base and denser pinnately semicraspedodromous venation. They do correspond with the neotype in their complicated looping along the margin and finely toothed margins. Another available specimen (pl. 2, fig. 10) has a rounded apex, whereas in the neotype and in one of the missing syntypes (UNGER 1866: pl. 19, fig. 25) the apex is bluntly acute. Leaf size varies in length from 35 to probably more than 50 mm and in width from 15 to about 35 mm. Only few specimens are available to circumscribe this newly recognised barberry of the European Neogene. The type of venation and marginal teeth are traits leading to the proposed alliance with Berberis. The syntypes are unfortunately missing and thus the protologue was the only basis for the new typification. Extant barberries having similar foliage are confined mostly to East Asia (e.g. B. pruinosa FRANCH., B. centiflora DIELS). Similar fossil leaf impressions occur elsewhere in the European Neogene, e.g. in the Middle Miocene flora of South Bohemia (KNOBLOCH & KVACEK 1996, as cf. ? Berberis sp.). CEK comb. nov. Berberis (?) ambigua (UNGER) KOVAR-EDER & Z. KVAC Plate 2, Figure 11 1847 Ilex ambigua UNGER, Chlor. prot., p. 149, pl. 50, fig. 14 (LMJ 76519, holotype) – basionym. 1850a Ilex ambigua UNGER – UNGER, p. 461. Holotype: LMJ 76519 – figured by UNGER (1847: 149, pl. 50, fig. 14), refigured in pl. 2, fig. 11. The spiny margin and the obovate shape of the only available specimen suggest Berberis. The secondary and higher order venation is not preserved to confirm this assumption. B. teutonica described above differs in a broader leaf lamina and less conspicuous marginal teeth (like in B. berberidifolia). Several extant species from China have similar leaves with thorny teeth. The scarcity and poor preservation of the available specimens prevent us from characterizing this obviously independent fossil species more precisely. Further leaf impressions of this species were described from Kymi, Greece (UNGER 1867 – as Ilex ambigua). Prinsepia serra described below differs in non-spiny marginal teeth. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 57 ? Mahonia NUTT. CEK comb. nov. Mahonia (?) aspera (UNGER) KOVAR-EDER & Z. KVAC Plate 13, Figures 1-8 1847 ? ? 1847 1850a 1850a 1850a 1850a 1864 1864 1864 1866 Quercus aspera UNGER, Chlor. prot., p. 108, pl. 30 figs. 1 (LMJ 76532 right, syntype), 2 (LMJ 76529 top right, lectotype), fig. 3, pl. 31, figs, 1, 3 - basionym. Ilex sphenophylla UNGER, p. 148, pl. 50, fig. 9 (LMJ 76515 lower left, holotype). Quercus aspera UNGER – UNGER, p. 400. Ilex sphenophylla UNGER - UNGER, p. 461 ("phenophylla"). Ilex cyclophylla UNGER, p. 461. Styrax boreale UNGER, p. 436. Ilex sphenophylla UNGER – UNGER, p. 12, pl. 3, figs. 3 (LMJ 76571), 4 (LMJ 76538), 5, 6. Ilex neogena UNGER, p. 13, pl. 3, figs. 9, 10, 11? (LMJ 76572). Ilex cyclophylla UNGER – UNGER, p. 13, pl. 3, figs. 7 (LMJ 76537), 8 (LMJ 76579). Styrax boreale UNGER - UNGER, p. 33, pl. 11, figs. 11-13. Lectotype designated here: LMJ 76529, figured by UNGER (1847: pl. 30, fig. 2 top right) – refigured in pl. 13, fig. 3. Additional material: GBA 2002/01/42-46, 79 b, 80-92a, 93; NHMW 1878/6/2081, 2758, 9140, 9474, 9489, 9492, 9498, 9917. These leaflets are typical due to their basal acrodromous venation, which went unnoticed by UNGER, and an irregularly spiny margin. They are further characterized by short petiolules and sometimes slightly asymmetric bases. In our concept this distinct morpho-species comprises both spiny and entire-margined leaflets. The true generic affinity remains unclear. The leaf margin, venation, and the sometimes slightly asymmetric base recall leaflets of Mahonia, e.g. M. nervosa (PURSH) NUTT. from western North America. Quercus and Ilex differ in their venation (no basal acrodromous veins). A very similar leaf from the Lower Pannonian of the Valea Crişului (Romania) was recently described by GIVULESCU (1998) as Mahonia sp. We hesitate to definitively include Styrax boreale because these specimens are rather large, partly recall legumes and the specimens themselves are missing. Cercidiphyllaceae Cercidiphyllum SIEB. & ZUCC. Cercidiphyllum crenatum (UNGER) R. BROWN Plate 2, Figure 7 Material: NHMW 1878/6/6510. In all the Parschlug material studied, we discovered only one specimen. The incomplete leaf is roundish, typically crenulate, with the palmate venation. It fits well within the variation known from other records of this species. Altingiaceae Liquidambar L. 58 Annalen des Naturhistorischen Museums in Wien 105 A Liquidambar europaea A. BRAUN Plate 2, Figures 1-5 1847 1847 1850a 1851a 1852 1850a 1852 1878b Acer parschlugianum UNGER, p. 132, pl. 43, fig. 5 (LMJ 76517, holotype). Liquidambar europaeum A. BRAUN - UNGER, p. 120, pl. 35, figs. 1 (LMJ 76523), 2 (LMJ 76867), 3 top (LMJ 76516), 4, 5. Liquidambar protensum UNGER, p. 415. Liquidambar europaeum A.BRAUN - ETTINGSHAUSEN, p. 15, pl. 2, figs. 20, 22. Liquidambar protensum UNGER – UNGER, p. 116, pl. 43, fig. 27 (LMJ 76508, holotype). Liquidambar acerifolium UNGER, p. 415. Liquidambar acerifolium UNGER - UNGER, p. 116, pl. 43, fig. 28 (LMJ 76492, holotype). Liquidambar europaeum A. BRAUN - ETTINGSHAUSEN, p. 86, pl. 2, figs. 3 (NHMW 1878/6/2406), 4, pl. 3, fig. 7 (NHMW 1878/6/2453 bearing Xylomites liquidambaris ETTINGSHAUSEN), 4. Additional material: numerous specimens, e.g. GBA 2002/01/79a, 94a; NHMW 1878/6/9052, 9542, 9546, 7738. Remains of this fossil representative of sweet-gum tree are common at Parschlug, mainly as foliage. The typical leaf form varies from trilobate to quinquelobate, the latter rarely with additional lobes on the margin ("protensum" form). Contrary to the opinion of UNGER we believe that only one natural species with a wider variation in foliage occurred at Parschlug. The studies undertaken so far on the leaf cuticles indicate a close relationship of most Neogene leaf records of Europe to extant L. styraciflua of North America. Not having better preserved material at hand, we share this opinion, although L. orientalis is not distinguishable based solely on gross morphology. According to the nomenclatural rules, the ending of the epithet must be corrected to "europaea" to agree with the gender of the genus (feminine). Liquidambar sp. – fructus Plate 2, Figure 6 1847 Liquidambar europaeum A. BRAUN - UNGER, p. 120, pl. 35, fig. 3 bottom (LMJ 76516). Additional material: GBA 2002/01/96; NHMW 1878/6/9538. Contrary to the leaves, the fruiting heads are rather rare (see Material and Methods). The surface and details of the fruits are poorly preserved and do not show necessary details to decide the specific affinities (Liquidambar magniloculata CZECZOTT & SKIRGIELLO versus L. wutzleri GREGOR). Platanaceae Platanus L. Platanus leucophylla (UNGER) KNOBLOCH Plate 3, Figure 10, Plate 4, Figure 17 1850a Populus gigas UNGER, p. 417. 1852 Populus gigas UNGER - UNGER, p. 117, pl. 44, fig. 1. 1866 Acer productum A. BRAUN - UNGER, p. 46, pro parte, pl. 15, fig. 1. Material: IBUG Ett. coll. 1140 (level II), NHMW 1878/6/7713. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 59 Although the figured specimens listed in the synonymy were not found in the collections, several others can be safely identified as this plane tree with pedate-palmately lobed foliage common during middle-late Neogene times in Europe. They differ from the extant P. orientalis of southern Europe and the Near East in having broader lobes and match in this respect another extant species of Europe, the London Plane (P. hispanica) of uncertain origin. Betulaceae Betula L. Betula cf. dryadum BRONGNIART Plate 3, Figure 1 1847 1852 Betula dryadum BRONGNIART - UNGER, p. 117. Betula dryadum BRONGNIART - UNGER, p. 33, pl. 16, fig. 10 (LMJ 76497). Additional material: IBUG Ett. coll. 725. Winged fruitlets of birch are very rare and in principle correspond to the basic type assigned usually to Betula dryadum. Due to poor preservation, any more precise identification is out of the question. Betula vel Alnus sp. Plate 3, Figures 3, 4 1847 Fagus deucalionis UNGER, p. 101, pro parte. 1850a Fagus deucalionis UNGER – UNGER, p. 405, pro parte. 1852 Fagus deucalionis UNGER – UNGER, p. 38, pro parte, pl. 18, fig. 24 (LMJ 76489). Additional material: NHMW 1878/6/2490, 6499, 2001B0017/0004. Betulaceous foliage is not easily distinguishable to the genus level. Several leaf impressions of this kind were encountered in the Parschlug assemblage. Those that were better and more completely preserved are similar in the shape of the lamina and the marginal teeth to Alnus adscendens (GOEPPERT) ZASTAWNIAK & WALTHER (1998). In our opinion, this entity may also include birch foliage. This leaf type is extremely rare at Parschlug and in our case may belong either to a birch or an alder. Although UNGER identified this type of foliage as Fagus deucalionis, the latter was based on fruits (UNGER 1847) and cannot be used in this context. The leaf from Parschlug attributed by UNGER (1852) to F. deucalionis definitely belongs to the Betulaceae. Alnus Mill. CEK & HOLY Y Alnus julianiformis (STERNB.) Z. KVAC Plate 3 Figure 6 Material: IBUG Ett. coll. 284 (II). Only a single leaf fragment can be attributed – based on morphological traits (shape, marginal indistinct teeth, and craspedodromous venation) – to this species of alder, 60 Annalen des Naturhistorischen Museums in Wien 105 A widely distributed in Europe during the Miocene. It is one of the more thermophilic summergreen elements, corresponding best to the extant A. trabeculosa – A. formosana group of SE Asia. CEK Alnus gaudinii (HEER) KNOBLOCH & Z. KVAC Plate 3 Figure 5 Material: NHMW 1878/6/7508 + 9412 (part + counterpart) det. by ETTINGSHAUSEN as Castanea atavia and Quercus mediterranea. Lamina elliptic, 70 mm long (incomplete), 27 mm wide, base missing, apex acute, leaf margin finely double serrate, teeth slender and sharp; venation (semi)craspedodromous, midvein slender, straight, secondaries slender, densely spaced (6-10 mm), almost straight and parallel, occasionally forking, one branch entering the first order tooth, others either running marginally or entering second order teeth; tertiary veins percurrent, densely spaced, slightly oblique. This leaf resembles Alnus gaudinii, particularly those forms with relatively sharp teeth. Similar leaf forms occur in the Miocene to Pliocene in Europe (e.g. at Berga, MAI & WALTHER 1988). Fagaceae Fagus L. Fagus sp. - leaf Plate 3, Figures 7-9 1882 Fagus feroniae UNGER - ETTINGSHAUSEN, p. 99, pl. 17, fig. 2 (NHMW 1878/6/2491, counterpart 2492). Additional material: IBUG Ett. coll. 986, 989. Beech leaves are rare in the Parschlug assemblage. The limited number of complete specimens prohibits statistically evaluating the number of secondaries. The margin shows prominent teeth with craspedodromous endings of the secondaries, a feature of Oligocene F. saxonica, but this feature is developed also in the subsequent beech maximum of Europe starting with the latest Early Miocene. These leaf forms have been usually assigned to F. menzelii, F. kraeuselii or F. silesiaca. DENK (2002) plans to lump all local populations of this beech in Central and West Europe into a single variable species. All Tertiary fruits from Europe have been recently united in a single morphospecies Fagus deucalionis (DENK & MELLER 2001). It would be unwise to solve taxonomical problems of fossil beech foliage on the basis of the limited Parschlug material. Fagus sp. - cupule 1847 Fagus deucalionis UNGER, p. 101, pro parte. 1850a Fagus deucalionis UNGER - UNGER, p. 405, pro parte. 1852 Fagus deucalionis UNGER - UNGER, p. 38, pro parte, pl. 18, fig. 25 (LMJ 62667). The specimen is poorly preserved due to pyritisation. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 61 Fagus vel Alnus sp. Plate 3 Figure 2 Material: NHMW 1878/6/9137. We hesitate to assign this leaf unambiguously to Fagus because the leaf margin bears occasional secondary teeth between the main teeth above the secondary veins. Secondary teeth are extremely rare in foliage of extant beeches, e.g. Fagus pashanica YANG (KVACEK & WALTHER 1991). They occasionally occur even in fossil Fagus silesiaca WALTHER & ZASTAWNIAK (1991: Fig. 1:6). Quercus L. Quercus drymeja UNGER Plate 4, Figures 1-7 1847 1850a 1850a 1850b ? ? ? 1850a 1852 1852 1878b Quercus drymeja UNGER, p. 113, pl. 32, figs. 1 right (LMJ 76524 A, lectotype), 2, (non 3 = Myrica lignitum), 4. Quercus drymeja UNGER - UNGER, p. 400. Juglans hydrophila UNGER, p. 469. Juglans hydrophila UNGER - UNGER, p. 196, pro parte, pl. 53, figs. 7 (LMJ 76549), ? 8, 9 (LMJ 76541). Quercus urophylla UNGER, p. 403. Quercus mediterranea UNGER - UNGER, p. 35, pl. 18, fig. 4. Quercus urophylla UNGER - UNGER, p. 36, pl. 18, fig. 11. Quercus drymeja UNGER - ETTINGSHAUSEN, p. 87, pl. 3, fig. 10 (NHMW 1878/6/6557) bearing Xylomites drymejae ETTINGSHAUSEN. Lectotype designated here: LMJ 76524 A, figured by UNGER (1847: 113, pl. 32, fig. 1 right) – refigured in pl. 4, fig. 1. Additional material: GBA 2002/01/40, 105, 108, 110, 113; NHMW 1878/6/2447, 9388, 9399. Quercus drymeja is one of the most common sclerophyllous oaks of the Mediterranean area. It is distinguished from similar leaf forms from the Boreal Province (e.g. KNOBLOCH & KVACEK 1996, as Q. cf. drymeja) by a slender lamina with regular spiny teeth. A recent study of cuticle structure on the material from Vegora (KVACEK et al. 2002 b) showed that the abaxial leaf side of Q. drymeja bears occasional solitary, massive trichome bases typical of many sclerophyllous oaks, but certainly not of Quercus ilex, which has typically a densely hairy abaxial leaf surface. That is why this extant oak cannot serve as a living analogue for Q. drymeja, as has been traditionally maintained. On the other hand, the differences from the epidermal structure of Q. mediterranea are negligible. As is apparent in both the Parschlug and Vegora assemblages, transitions between these two species occur with regard to morphology of the leaf lamina. At Parschlug there are leaf forms similar to Myrica lignitum (narrow cuneate base) and to Quercus zoroastri (coarser toothed leaf margin and more rounded leaf base). The species can be easily distinguished in leaf assemblages whose cuticle structure is preserved. The nearest living relative of Q. drymeja should be sought among sclerophyllous oaks, most probably outside Europe. 62 Annalen des Naturhistorischen Museums in Wien 105 A Quercus mediterranea UNGER Plate 4, Figures 8-16 1847 1850a 1850a 1850a 1852 1852 1866 1878b Quercus mediterranea UNGER, p.114, pl. 32, figs. 1 top left (LMJ 76524B, lectotype), 5 ?, 6 ?, 7, 8 ?, 9 (NHMW 1845/0034/4, syntype). Quercus mediterranea UNGER – UNGER, p. 400. Quercus cyclophylla UNGER, p. 400. Prunus theodisca UNGER, p. 484. Quercus mediterranea UNGER - UNGER, p. 35, pl. 18, figs. 1 (LMJ 76507), 2, 3, 5, 6. Quercus cyclophylla UNGER - UNGER, p. 37, pl. 18, fig. 15. Prunus theodisca UNGER - UNGER, p. 61, pl. 18, fig. 31. Quercus mediterranea UNGER – ETTINGSHAUSEN, p. 83, pl. 1 fig. 6, 7 ?, 8 ? (bearing Sphaeria mediterranea ETTINGSHAUSEN). Lectotype: because the specimen selected by ILJINSKAYA (in TAKHTAJAN 1982: 102) is missing, a new lectotype is designated here: LMJ 76524 B, UNGER 1847, pl. 32, fig. 1 top left – refigured in pl. 4, fig. 8. Additional material: GBA 1864/01/5, 2002/01/19, 106, 107; IBUG Ett. coll. 908, 912, 914, 934, 943 + 944 (part + counterpart), 949 + 950 (part + counterpart); NHMW 1878/6/7532, 9374, 9381. This sclerophyllous oak was widely distributed during the Neogene in southern Europe and adjacent areas. Its limits towards the previous Q. drymeja, having slender leaves, are somewhat arbitrary because of highly variable foliage (KVACEK et al. 2002 b). This variation is well expressed at Parschlug, which is the type locality of this species and where this species was described under several binomina (see synonymy). The cuticle structure obtained from various Late Miocene sites of Greece suggests affinities to the group of extant Q. coccifera (KVACEK & WALTHER 1989: fig. 5a, b). Quercus zoroastri UNGER Plate 5, Figures 1-4 1850a Quercus zoroastri UNGER, p. 401, pro parte. 1850b Juglans hydrophila UNGER, p. 196, pro parte, pl. 53, fig. 6 (LMJ 76866). 1852 Quercus zoroastri UNGER - UNGER, p. 36, pro parte, pl. 18, figs. 7, 8. Neotype designated here: NHMW 1878/6/2401 – figured in pl. 5, fig. 1. Additional material: GBA 2002/01/42; IBUG Ett. coll. 932, NHMW 1878/6/6478, 9377. Leaves broad elliptic to ovate, long petiolate, coriaceous, cuneate to rounded at the base, coarsely simple toothed except the entire base, venation craspedodromous, secondaries straight to slightly bent, dense, never forked, entering regular, ± closely spaced, apically directed teeth. Tertiary veins inconspicuous. In our opinion this is also a sclerophyllous oak; it differs in the long petiole from Q. mediterranea and is characterised by the shape of lamina, which is broad elliptic to ovate (contrary to Q. drymeja and Q. kubinyii), and by the shape of teeth, which resembles some forms of Q. kubinyii. The occurrence of occasional secondary teeth between primary teeth above the secondary veins is a feature characteristic of this species and suggestive of the morphology of Prinsepia (see P. serra below). The base of the leaf is usually symmetric, contrary to UNGER’s description (1852: 36), which incorrectly includes a leaflet of Sapindus pythii (UNGER 1852: pl. 18, fig. 8). Some leaf forms of Q. sosnowskyi KOLAKOVSKII may be similar in tooth form and lamina shape, but there are KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 63 also pronounced differences between the two species - Q. sosnowskyi has very typical, distinct percurrent tertiary venation and often forked secondaries. No very similar living relative of Q. zoroastri has been recovered. Theaceae ? Gordonia ELLIS cf. ? Gordonia oberdorfensis KOVAR-EDER Plate 5, Figures 5-8 Material: NHMW 1878/6/2004 + 2005 (part + counterpart), 2006 + 2007 (part + counterpart) determined by ETTINGSHAUSEN as Ficus troglodytarum UNGER, 1878/6/2009, 20025 + 2026 (part + counterpart), 2038 as Diospyros sp. nov. in sched., 2021 + 2022 as Daphne sp. nov. in sched.; probably NHMW 1878/6/2701. These elongate and rather large leaves with entire margin have a very thick and straight midvein. The secondaries are very dense, originating at very steep angles; they initially run aside the midvein and then diverge towards the leaf margin. The secondaries run an irregular course, sometimes fork and the branches of neighbouring secondaries join each other. Although we lack information about the epidermal structure of the leaves from Parschlug, the venation pattern is very similar to that of ? Gordonia oberdorfensis (KOVAR-EDER & MELLER 2001: 79) and extant Gordonia axillaris (KVACEK & WALTHER 1984a: pl. 25, fig. 3 – as Polyspora axillaris). These leaves derive from level I with the exception of the specimen NHMW 1878/6/2701, which is from level III. Ternstroemites BERRY emend. HICKEY CEK comb. nov. Ternstroemites pereger (UNGER) KOVAR-EDER & Z. KVAC Plate 6, Figures 1-7 1850a 1850a 1850a 1850b 1852 1866 Carpinus oblonga UNGER, p. 409, pro parte. Amygdalus pereger UNGER, Gen. spec. pl. foss., p. 483, pro parte, basionym. Crataegus orionis UNGER, p. 481. Amygdalus pereger UNGER - UNGER, p. 184, pro parte, pl. 55, figs. 11, 13, 14. Carpinus oblonga UNGER - UNGER, p. 10, pro parte, pl. 20 fig. 16. Crataegus oreonis UNGER - UNGER, p. 59, pl. 18, fig. 15 (LMJ 76593). Neotype designated here: NHMW 1878/6/8169 det. by ETTINGSHAUSEN as Fraxinus intermedius ETTINGSHAUSEN - figured on pl. 6, fig. 1. Additional material: GBA 2002/01/39 (part + counterpart), 41, 111, 112, 114, NHMW 1853/26/473 det. as Ceratopetalum parschlugianum ETTINGSHAUSEN, 1878/6/7451 + 9516 (part + counterpart) det. by ETTINGSHAUSEN as Carpinus oblonga and Quercus serra, 1878/6/7452 det. by ETTINGSHAUSEN as Carpinus oblonga UNGER, 1878/6/8171 det. by ETTINGSHAUSEN as Fraxinus intermedia. Leaves lanceolate to elongate, long petiolate, leaf margin crenulate, except at the very base, with more or less distinct apical glands; secondary veins semicraspedodromous. The morpho-genus Ternstroemites BERRY as emended by HICKEY (1977: 141) comprises foliage of the Theaceae, which are characterised by having simple glandular teeth on 64 Annalen des Naturhistorischen Museums in Wien 105 A the margin. Although we are not well informed about the detailed venation of the studied specimens due to coriaceous texture of the leaves, the species as emended above fits well into this group of leaf forms. Unfortunately, we are unable to corroborate this assignment with the epidermal structure, and it is therefore difficult to make comparisons with the previously described Theaceae leaves of the European Tertiary, which are mostly based on both gross morphology and epidermal anatomy (e.g. KVACEK & WALTHER 1984b). Myricaceae Myrica L. Myrica lignitum (UNGER) SAPORTA Plate 7, Figures 1-6, 8, 9 1847 1850a 1850a 1850a 1850b 1851b 1852 Quercus lignitum UNGER, p. 113, pl. 31, figs. 5-7. Quercus lignitum UNGER - UNGER, p. 402. Comptonia laciniata UNGER, p. 394. Prinos hyperboreus UNGER, p. 462. Comptonia laciniata UNGER, p. 161, pl. 29 fig. 2. Dryandroides lignitum (UNGER) ETTINGSHAUSEN, p. 741, pl. 34, fig.5 (GBA 1851/04/10). Quercus lignitum UNGER, p. 34, pro parte, pl. 17, figs. 1 and 2 (LMJ 76504), 3, 4 (LMJ 76503), 5, 6 (LMJ 76510 right), 7 (LMJ 76485). 1852 Quercus commutata UNGER, p. 35, pl. 17, figs. 8, 9, 10 (LMJ 76510 left). 1864 Prinos hyperboreus UNGER - UNGER, p. 14, pl. 3, fig. 37. 1878b Myrica lignitum (UNGER) SAPORTA - ETTINGSHAUSEN, p. 82, pl. 1 figs. 1, 2 (bearing Phyllerium lignitum ETTINGSHAUSEN). 1880 Myrica lignitum (UNGER) SAPORTA - ETTINGSHAUSEN, pl. 12, figs. 1-17 (fig.1 - NHMW 1878/6/9260, fig. 5 - NHMW 1878/6/7376, fig. 10 - NHMW 1878/6/9270, fig. 12 - NHMW 1879). 1888 Myrica lignitum (UNGER) SAPORTA - ETTINGSHAUSEN & STANDFEST, pl. 1, figs. 2, 3 (694), 5 (477), 6 (482), 7, 8 (488), 9 (489), 10, 11 (492), 12 (500),13 (509), 15 (517), 16 (524), 17 (531), 18, 19 (559), pl. 2, figs. 20 (568), 21 (581), 23 (591), 24 (598), 25 (605), 26 (608), 27 (612), 28 (615), 29 (617), 32 (620), 34 (631, 632), 36 (647), 37 (653), 38 (661+662), 39 (666), 40 (718), 41 [non 33 (630), 35 (633)] (all specimen nos in brackets ex IBUG Ett. coll.). 1976 Myrica lignitum (UNGER) SAPORTA – KNOBLOCH & KVACEK, p. 20-21, pl. 8, figs. 1-3. 1982 Myrica lignitum (UNGER) SAPORTA - KOVAR, p. 80, pl. 12 figs. 1-8 (coll. file nos see there). Lectotype designated here: LMJ 76503, figured by UNGER (1852: pl. 17, fig. 4) - refigured on pl. 7, fig. 6. Additional material: GBA 1847/03/11, 2002/01/1, 92b; IBUG Ett. coll. 1083 (II), 1084 (III); NHMW 1878/6/2324, 2339c, 2367, 2426, 2563, 7382, 8841, 9309, 9312. This species was revised at an earlier date by the first author (KOVAR 1982), who circumscribed M. lignitum including cuticular anatomy and the variation in its leaf morphology. Leaf anatomical features have been obtained from a number of other Miocene sites in Europe (see KNOBLOCH & KVACEK 1976), making this quite variable species now well recognisable. In addition, leaf forms identified as Comptonia laciniata UNG. (an additional specimen in NHMW 1878/6/7382) represent, in our opinion, extreme morphological variations of M. lignitum at Parschlug. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 65 Myrica oehningensis (A. BRAUN) HEER Plate 7, Figure 7 1850a Comptonia oeningensis A. BRAUN - UNGER, p. 394. 1850b Comptonia oeningensis A. BRAUN - UNGER, p. 161, pl. 29, fig. 3 (LMJ 76546). Leaves designated under this species recall Comptonia and so were identified by UNGER (1850a, b) and ETTINGSHAUSEN (1851a – as Comptonia vindobonensis). Their lamina, however, differs from that of the extant Comptonia peregrina by much more irregular dissection of the margin tending from deeply lobed to coarsely crenate-toothed. At Parschlug, such a leaf form is exceptional, but elsewhere, e.g. in Bavaria (RIEDERLE & GREGOR 1997, as Comptonia oehningensis and Myrica ungeri vel Comptonia oeningensis) is very typical and common (see tab. 2). The taxonomic position of such leaf remains is to be solved on the basis of cuticular anatomy. Myrica sp. - fructus 1888 Myrica lignitum (UNGER) SAPORTA – ETTINGSHAUSEN & STANDFEST, pl. 1, figs. 1 a-c. This single specimen described and figured by ETTINGSHAUSEN & STANDFEST (1888) is missing. It was possibly lost due to pyritisation. Juglandaceae Engelhardia LESCH. Engelhardia orsbergensis (WESSEL & WEBER) JÄHNICHEN, MAI & WALTHER Plate 6, Figures 10-12 Material: GBA 2002/01/22, 100; IBUG Ett. coll. 723, 841; NHMW 1878/6/2053a, 2816, 8951 det. by ETTINGSHAUSEN as Hakea parschlugiana, 9123. Parschlug is a new site for this species (known also as Palaeocarya orsbergensis (WESSEL & WEBER ) JÄHNICHEN, FRIEDRICH & TAKÁC), which was widely distributed in the Tertiary of Europe. The leaflet morphology and its association with the fruits described below make this record unequivocal. Engelhardia macroptera (BRONGNIART) UNGER Plate 6, Figures 8, 9 1850b Carpinus producta UNGER, p. 164, pl. 32, fig. 6 (LMJ 76540 - UNGER erroneously mentioned the locality of Socka instead of Parschlug). Additional material: NHMW 1878/6/2697, 2698 (part + counterpart), NHMW 1879/610. Involucra of this species are rare at Parschlug. They do not differ from the standard form typified by the specimens from Armissan, France (JÄHNICHEN et al. 1977); they share triveined lobes of the involucrum and occasionally bear an impression of the fruit. Also, as at other occurrences in Europe, the fruits are associated with the leaflets and with the compound leaves as mentioned above. 66 Annalen des Naturhistorischen Museums in Wien 105 A Tiliaceae Tilia L. Tilia longebracteata ANDRAE Plate 6, Figures 13-15 1869 Tilia lignitum ETTINGSHAUSEN - p. 15, pro parte, pl. 42, fig. 6 (IBUG Ett. coll. 1541). Additional material: IBUG Ett. coll. 1663, fructus (det. by ETTINGSHAUSEN as Celastrus europaeus), Ett. coll. 2049, fructus (det. by ETTINGSHAUSEN as Prunus paradisiaca); GBA 2002/01/31, fructus. The few impressions of detached, globose, slightly angular fruits found at Parschlug recall those of linden. They are in our opinion not identifiable to the species level because of poor preservation. They are not attached to the bract described from Parschlug in the flora of Bílina by ETTINGSHAUSEN (1869), but may belong to the same plant. This isolated bract shows a fragment of the peduncle, which is attached at one point. Hence the bract is not adnate, as incorrectly drawn by ETTINGSHAUSEN. This type of bract is known to occur in several Tertiary species of linden in the Northern Hemisphere (type B sensu MANCHESTER 1994). The above designation is employed here as a morpho-species. Its holotype from Daia, Sarmatian (ANDRAE 1861: pl. 1, fig. 2), seems to represent the same kind of bract, devoid of fruits. No leaves of Tilia co-occur at Parschlug (impressions identified by ETTINGSHAUSEN as Tilia milleri – IBUG Ett. coll. 1542-1544 are clearly betulaceous fragments). Craigia W.W. SMITH & EVANS CEK, BŮZ ZEK & MANCHESTER Craigia bronnii (UNGER) Z. KVAC Plate 6, Figures 16, 17 1847 Ulmus bronnii UNGER, p. 100, pro parte. 1850a Ulmus bronnii UNGER - UNGER, p. 410, pro parte. Material: GBA 2002/01/35; IBUG Ett. coll. 1167, 2804a det. by ETTINGSHAUSEN as Ulmus bronnii; NHMW 1878/6/7581, 1878/6/7582, 1878/6/9676, 1878/6/9677. A few detached valves found at Parschlug are of the same kind as from other European sites. Such fruit remains have been assigned to several genera, recently to Craigia, an extant endemite of China (KVACEK et al. 1991, 2002a). Ulmaceae The various ulmaceous leaves found at Parschlug include transitional forms where we are unable to decide between Cedrelospermum, Zelkova, and Ulmus plurinervia, e.g. LMJ 76488, pl. 8, fig. 12. In the following text we document our views on the character of foliage in the respective species and we figure the most typical forms. Ulmus L. Ulmus plurinervia UNGER Plate 6, Figures 18-22 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 67 1847 Ulmus plurinervia UNGER, p. 95, pl. 25, figs. 1-4. 1850a Ulmus plurinervia UNGER - UNGER, p. 411. 1851a Planera ungeri ETTINGSHAUSEN, p. 14, pro parte, pl. 2, figs.11, 12. Neotype designated here: NHMW 1878/6/9667 - figured on pl. 6, fig. 19. Additional material: GBA 2002/01/101-104; IBUG Ett. coll. nos. 1031-1034 (twigs), 994, 1108, 1114, 1116; NHMW 1878/6/2650, 7592, 9082, 9155, 9154 + 9665 (part + counterpart). This elm species based on leaves from Parschlug was established by UNGER (1847) and later generally employed for small, more or less asymmetrical leaves with simple teeth. Only ILJINSKAYA (in TAKHTAJAN 1982: 16) doubted its affinity to Ulmus, suggesting that most of the figured syntypes were more likely foliage of Zelkova or Hemiptelea. As stated above, the limits of U. plurinervia are indeed partly uncertain towards both Zelkova and Cedrelospermum. However, the selected collection shown in pl. 6 is typical and almost indistinguishable from U. braunii HEER s.s. (Oehningen), except for the double serrate margin in most specimens of the latter. Without repeating the endless discussions about European Tertiary elms, we merely intend to clearly delimit U. plurinervia for further studies. The fruits described below are other organs of the same plant, adding fruit characters of this elm, which is typical of mesic assemblages of the European Neogene (e.g. Erdőbénye). CEK sp. nov. Ulmus parschlugiana KOVAR-EDER & Z. KVAC Plate 6, Figures 23-26 1843 1845 1847 1850a Ulmus zelkovaefolia UNGER, pro parte, pl. 24, fig. 7 left, fig. 8. Ulmus zelkovaefolia UNGER, pro parte, pl. 26, fig. 8. Ulmus zelkovaefolia UNGER, p.95 pro parte. Ulmus zelkovaefolia UNGER - UNGER, p. 411, pro parte. Holotype designated here: IBUG Ett. coll.1100 – figured on pl. 6, fig. 23. Paratypes designated here: NHMW 1878/6/ 9651, 9658 - figured on pl. 6, figs. 24, 26. Additional material: GBA 2002/01/95; IBUG Ett. coll. 1102-1104; NHMW 1878/6/2118, 9081, 9651. Elm samaras in bundles, stalked (stalk about 5 mm long), with persistent perianth, winged, broadly oval, typically 8-10 mm long and 6-9 mm wide, with an elliptic endocarp (ca. 2 x 3.5 mm in size) surrounded in the centre by moderately wide wings (the rim of the same width or slightly wider than the endocarp body) that broadly extend into a pair of styles. Wings with strong marginal vein and a prominent axial vein laterally deflected from the stipe towards the endocarp (pl. 6, fig. 26). As the leaves of Ulmus plurinervia are quite common and belong to the only elm species at Parschlug, we suspect that these fruits belong to the plant that produced U. plurinervia foliage. ETTINSGHAUSEN (1851a) was the first to arrive at this conclusion. Because the fruits occur detached from the leaves, they deserve in our opinion a separate binomen. Similar fruits were merged with Ulmus pyramidalis by BŮZEK (1971) because, in the Early Miocene Most Formation of North Bohemia, they regularly accompany leaves indistinguishable from U. pyramidalis. At Schrotzburg two kinds of fruits occur – one indistinguishable from our U. parschlugiana (HANTKE 1954: pl. 6, fig. 18) and the other (HANTKE 1954: pl. 6, fig. 17) of the Chaetoptelea-type (sensu MANCHESTER 1989). At Sośnica, the type locality of U. pyramidalis and U. carpinoides, 68 Annalen des Naturhistorischen Museums in Wien 105 A three kinds of fruits occur, one of the Chaetoptelea-type (GOEPPERT 1855: pl. 14, figs. 15-18) and two others of different size with broader wings (GOEPPERT 1855: pl. 14, figs 18-20 and 21). In the latter cases, the fruits have very short stipes so that the perianth adheres directly to the fruit base, in contrast to U. parschlugiana. Fruits similar to U. parschlugiana are produced for example by extant U. americana (cf. MANCHESTER 1989: fig. 12.1 c). A more extensive study of the discussed material is necessary to clarify the taxonomy of the whole group. Cedrelospermum SAPORTA emend. MANCHESTER CEK comb. nov. - foliage Cedrelospermum ulmifolium (UNGER) KOVAR-EDER & Z. KVAC Plate 8, Figures 1-5 ? ? 1850a 1850a 1850b 1851a 1850a 1860 1866 Comptonia ulmifolia UNGER, Gen. spec. pl. foss., p. 394 - basionym. Rhus triphylla UNGER, p. 474. Comptonia ulmifolia UNGER - UNGER, p. 194, pl. 29 figs. 4, 5 (LMJ 76536, lectotype). Planera ungeri ETTINGSHAUSEN, p. 14, pro parte, pl. 2, figs. 15, 17, ? 18. Prunus euri UNGER, p. 485. Rhus triphylla UNGER - UNGER, p. 44, pl. 20, fig. 13. Prunus euri UNGER - UNGER, p. 61, pl. 18 fig. 30. Lectotype designated here: LMJ 76536, figured by UNGER (1850b: 194, pl. 29, fig. 5) – refigured on pl. 8, fig. 5. Additional material: GBA 2002/01/38; IBUG Ett. coll. 135, 1079, 1085, 1087, 1088, etc. NHMW 1878/6/2053b, 7634, 9572, 9573, 9575, 9619, 9622, 9630 (det. by ETTINGSHAUSEN Planera ungeri in sched.), 1878/6/7520 (det. by ETTINGSHAUSEN Quercus lonchitis in sched.). The variability of this species is similar to that encountered particularly in the Middle Miocene localities of Europe (e.g. Randeck Maar, RÜFFLE 1963, as Tremophyllum tenerrimum). Contrary to similar leaves of Ulmus plurinervia and Zelkova zelkovifolia, the asymmetrical, slender forms with a narrow, elongate upper part towards the apex are typical of this species. Transitional forms occur to both mentioned taxa. Larger leaves of Cedrelospermum are particularly difficult to differentiate from Ulmus. Ulmus parvifolia A. BR. sensu UNGER (1852: 43, pl. 20, fig. 22 - LMJ 76488) may be either Cedrelospermum or Ulmus plurinervia. ETTINGSHAUSEN (1851a), who was unaware of the existence of Cedrelospermum at Parschlug, established his Planera ungeri in a broad sense to include foliage of Zelkova and Cedrelospermum into a single, unnatural entity. HABLY & THIÉBAUT (2002) established C. flichei (SAPORTA) HABLY & THIÉBAUT for the Palaeogene and Miocene morphotypes of foliage. In our opinion, those from Magyaregregy coincide with C. ulmifolium in all respects including wider angles of the base (and the absence of intersecondaries - ? due to poor preservation). In view of the slight size difference of associated fruits from the type locality in the Palaeogene of southern France (see below) also these leaves warrant a separate taxon. CEK Cedrelospermum stiriacum (ETTINGSHAUSEN) KOVAR-EDER & Z. KVAC comb. nov. - fructus Plate 8, Figure 6 1888 Embothrium stiriacum ETTINGSHAUSEN, Denkschr. k. Akad. Wiss. math.-nat. Cl. 54, p. 316, pl. 4, fig. 32 (lectotype, NHMW 1878/6/3583) - basionym. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 69 Lectotype designated here: NHMW 1878/6/3583, figured by ETTINGSHAUSEN 1888: pl. 4, fig. 32 (Moskenberg). Additional material: IBUG Ett. coll. 1359, 1360+1361 (part + counterpart), 1362 det. by ETTINGSHAUSEN as Embothrium megalopterum ETTINGSHAUSEN in sched., 1364, 2899 as Embothrium stiriacum; NHMW 1878/6/8045, 8046 det. by ETTINGSHAUSEN as Embothrium giganteum ETTINGSHAUSEN in sched. Samaras with a single wing innervated by about 6 sub-parallel veins and asymmetrically positioned cleft of two persistent styles, 5-8 mm wide and typically 14-17 (- 23) mm long, seed part oblique to the wing, mostly beak-like narrowed on the base. Fruits of Cedrelospermum have been encountered at Parschlug for the first time. They conform in morphology and wing venation to the populations from the Oligocene to Middle Miocene of Europe (HABLY & THIÉBAUT 2002) but notably match in a bigger mean size and a narrower base of the seed part the population from Magyregregy. THIÉBAUT (personal communication) is going to separate the latter into a new species C. hablyae. The binomen suggested here has the priority. Records from Leoben (ETTINGSHAUSEN 1888), Schönegg (ETTINGSHAUSEN 1890) and the Randeck Maar (RÜFFLE 1963) belong to the same species. Because this plant has not yet been found as twigs with attached fruits (contrary to other species of the genus), the associated leaves must be given a separate binomen (see above). Zelkova SPACH Zelkova zelkovifolia (UNGER) BŮZZEK & KOTLABA Plate 8, Figures 8-11 1843 1845 1847 1850a 1850a 1851a 1852 1852 Ulmus zelkovaefolia UNGER, pl. 24 figs. 7 (right), 9-13. Ulmus zelkovaefolia UNGER, pl. 26, fig. 7 (NHMW 1987/57, lectotype). Ulmus zelkovaefolia UNGER, p. 94 (here valid diagnosis). Ulmus zelkovaefolia UNGER - UNGER, p. 411, pro parte. Ulmus praelonga UNGER, p. 411. Planera ungeri ETTINGSHAUSEN, p. 14, pro parte, pl. 2 figs. 7, 13, 16. Zelkova ungeri KOVATS - UNGER, p. 42, pl. 20, fig. 19. Ulmus praelonga UNGER - UNGER, p. 43, pl. 20, fig. 20 (LMJ 76487, holotype). Lectotype designated here: NHMW 1987/57, figured by UNGER (1845: pl. 26, fig. 7) - refigured on pl. 8, fig. 9; previous lectotypification by ILJINSKAYA (in TAKHTAJAN 1982: p. 18) is invalid because the selected specimen is missing and a mere illustration is not accepted by the current nomenclatural rules for the typification. Additional material: GBA 2002/01/18; NHMW 1878/6/9590, 9642. Parschlug is the type locality of this common member of Tertiary floras of Eurasia. Besides coarsely simple-toothed large leaves of sterile twigs (pl. 8, figs. 10, 11), smaller forms also co-occur, the latter mostly attached to twigs, sometimes still bearing the fruits. This dimorphy, known also in extant Zelkova, led some authors to accept another independent species, Z. praelonga, (e.g. BERGER 1953). Some leaf records described as Zelkova ungeri from the Bavaria Molasse (e.g. RIEDERLE & GREGOR 1997) and Oehningen (HANTKE 1954) do not belong to Zelkova but represent probably leaflets of the Vitaceae (Parthenocissus). 70 Annalen des Naturhistorischen Museums in Wien 105 A Celtidaceae Celtis L. Celtis japeti UNGER Plate 8, Figure 7 1850a Celtis japeti UNGER, p. 412. 1852 Celtis japeti UNGER - UNGER, p. 44, pl. 20, figs. 25-26. Neotype designated here: NHMW 1878/6/7654 – figured on pl. 8, fig. 7. Additional material: NHMW 1878/6/7691. Because neither of the two type specimens is available, the lectotypification done by KUTUZKINA (in TAKHATAJAN 1982) is currently unacceptable. Of two topotypical specimens, which were discovered in the collections, the one with better preserved venation is selected here to serve as the neotype. We doubt that the population from Parschlug can be definitively distinguished from the records of Erdőbénye (Celtis trachytica ETTINGSH.) and Sośnica (C. begonioides GOEPP.), but more numerous collections of these entities are necessary to compare their variation. If merged (e.g. by KRISHTOFOVICH & BAYKOVSKAYA 1965), C. japeti has priority over C. trachytica. Salicaceae Populus L. Populus populina (BRONGNIART) KNOBLOCH Plate 8, Figure 18, Plate 14, Figure 1 1850a 1850a 1852 1852 Populus latior A. BRAUN - UNGER, p. 416. Populus aeoli UNGER, p. 416. Populus latior A. BRAUN – UNGER, p. 45, pl. 21, figs. 3 (LMJ 76509), 4 (LMJ 76505), 5. Populus aeoli UNGER - UNGER, p. 45, pl. 21, fig. 2 (LMJ 76506, holotype). Additional material: GBA 2002/01/29, 78. The few specimens studied from Parschlug fit well within the morphological variation of this poplar, which was widely spread during the Neogene of Europe. Because only one species of Populus has been demonstrated at Parschlug, the fruits described below probably belong to the same plant. Populus sp. - fructus Plate 8, Figures 19-21 1850a Celastrus europaeus UNGER, p. 459, pro parte. 1866 Macreightia germanica HEER - UNGER, p. 26, pl. 8, figs. 12 bottom, 13. Additional material: GBA 2002/01/32, 34, 115; IBUG Ett. coll. 1665, 1666, 1693; NHMW 1878/6/9896, 2387. Capsules open, shortly stalked, trivalvate, partly incompletely preserved, valves elliptical, 3-5 mm wide and 5 – 10 mm long, clearly convex, flattened by fossilisation, slightly rugulate on the outer surface. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 71 UNGER (1850a: 459) initially combined some leaves and the above-described capsules under Celastrus europaeus, but later (UNGER 1864: 10, 1866: 26) decided to separate the fruits and determined them as Macreightia germanica HEER (calyx). We, however, have no doubts that these remains are fruits (partly opened capsules) belonging to Populus populina. Although we did not find the original specimens, we discovered topotypical material. The trivalvate capsules figured by HEER (1859: pl. 103, figs. 1,2) as Macreightia germanica definitely correspond to the remains from Parschlug and also co-occur in Oehningen with numerous leaves of P. populina (as P. latior HEER). Similar remains have been attributed to Populus in the Bavarian Neogene (GREGOR 1982: pl. 6, figs. 12-17). In our opinion these detached fruits express no diagnostic features that permit identification to the species level. Very similar capsules also co-occur with the leaves of the P. zaddachii HEER – type in North Bohemia within fluvial facies of the Most Formation, Late Oligocene to Early Miocene (personal observation Z. KVACEK). Buxaceae Buxus L. CEK, BŮZ ZEK & HOLY Y Buxus cf. egeriana Z. KVAC Plate 8, Figures 15 (?), 16, 17 (?) ? ? 1850a Quercus myrtilloides p. 404, pro parte. 1852 Quercus myrtilloides UNGER - UNGER, p. 38, pro parte, pl. 18, fig. 17 (LMJ 76502). Material: GBA 2002/01/14, LMJ 76524C (reverse side). The leaves assigned to this entity have a dense, complicated venation found in Buxus. They exceed in size Buxus pliocenica SAPORTA & MARION, which commonly occurred during the Late Neogene in Europe, but do not attain the length of typical, much more slender leaves of B. egeriana (type locality Habartov, Sokolov Basin, Ottnangian – KVACEK et al. 1982). The elliptical shape of the above-listed specimens recalls more the bigger leaves of B. pliocenica but differ partly by a long petiole. Some other leaf impressions identified as Myrsine formosana (BERGER 1955) and Buxus pliocenica (BERGER & ZABUSCH 1953) may represent the same type of foliage from the Middle Miocene from Lavanttal and the Vienna Basin. None of these impressions has been studied anatomically. Rosaceae Rosa L. cf. Rosa sp. Plate 8, Figure 14 1850a Spiraea zephyri p. 482. 1866 Spiraea zephyri UNGER - UNGER, p. 60, pl. 18, figs. 22, 23. 1866 Pyrus mini UNGER, p. 58, pl. 18, fig. 20. Material: IBUG Ett. coll. 1059 (level III). 72 Annalen des Naturhistorischen Museums in Wien 105 A Although none of the type specimens of Spiraea zephyri and Pyrus mini survived, we suspect that these illustrations represent leaflets of a rose. One terminal leaflet of the same kind is housed at IBUG and is illustrated here. It certainly cannot serve as a basis to sufficiently circumscribe a fossil species. Prinsepia ROYLE CEK comb. nov. Prinsepia serra (UNGER) KOVAR-EDER & Z. KVAC Plate 13, Figures 9-17 1847 Quercus serra UNGER, Chlor. prot. p. 109, pro parte, pl. 30, fig. ? 4 left (LMJ 76527, syntype), fig. 5 (LMJ 76528, lectotype), fig. 6 - basionym. 1850a Quercus serra UNGER - UNGER, p. 400, pro parte. 1852 Ulmus quercifolia UNGER, p. 43, pl. 20, fig. 24 (non UNGER 1847). 1852 Quercus serra UNGER - UNGER, p. 38, pl. 18, fig. 16 (LMJ 76495). 1878b Quercus serra UNGER – ETTINGSHAUSEN, p. 86, pl. 4, fig. 4 (NHMW 1878/6/6554, bearing Xylomites quercus serrae ETTINGSHAUSEN) Lectotype designated here: LMJ 76528, figured by UNGER (1847: pl. 30, fig. 5) - refigured on pl. 13, fig. 9. Epitype designated here: LMJ 76495, figured by UNGER (1852: pl. 18, fig. 16) - refigured on pl. 13, fig. 10. Additional material: GBA 2002/01/15-17, 75-78, 79b. NHMW 1878/6/2095, 2096, 2341a, 2423a, 2505, 2778, 2818, 7538 + 9528 (part + counterpart), 7539 + 9671 (part + counterpart det. by ETTINGSHAUSEN as Quercus serra and Ulmus plurinervia), 9502, 9505, 9509, 9511, 9514, 9517, 9519, 9521, 9525, 9527. Contrary to the original description the petioles measure at least 18 mm. The leaf base is sometimes asymmetric, the margin is densely, sharply, but irregularly toothed almost along the whole leaf length. The secondary veins are thin and densely spaced and interspaced with thin intersecondaries, semicraspedodromous. Especially near the leaf margin the higher order venation has a rather irregular pattern (exmedially ramified sensu ASH et al. 1999). Occasionally, when the marginal teeth are less elongated and less sharply pointed (pl. 13, fig. 17), these leaves resemble Ternstroemites pereger. In the latter, however, the teeth are more or less distinctly glandular and therefore apically rounded. We suspected a rosaceous affinity of these leaves and ultimately compared them with those of Prinsepia on E. ZASTAWNIAK’s suggestion. Indeed, the fossil leaves match particularly the large-leafed P. sinensis (OLIV.) OLIV. (subgen. Plagiospermum), notably in venation. We also cannot rule out a possibility that some of the entire-margined leaves from Parschlug with a similar dense venation pattern (e.g. "Quercus" daphnes shown on pl. 12, figs. 12 and 13) may belong to the same plant, because Prinsepia develops at the same time entire and toothed leaves. Fruits of Prinsepia have been recovered in the Miocene – Early Pliocene deposits of Central Europe and compared also to deciduous subgen. Plagiospermum (MAI 1984 b), which includes thorny shrubs distributed from Himalayas to E Asia. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 73 ? Prinsepia sp. Plate 8, Figure 13 Material: NHMW 1878/6/9747. One spiny twig fragment, the spines broadened at the base, curved, apex sharp. This twig fragment possibly belongs to the leaves described above. Fabaceae Among the many remains attributed by UNGER to Leguminosae, only few can be characterized as independent entities. It is impossible to definitively combine the fruits and foliage into a single species except in the case of Podocarpium. Therefore, the next suite of morpho-taxa deals with these two organs independently. In principle we apply a similar approach as HABLY (1992) except that we prefer to assign epithets to the morphogenus Leguminosites BOWERBANK as emended by SCHIMPER (1874), i.e. including both fruits, seeds and foliage, over Leguminocarpon GOEPPERT (= Leguminocarpos GOEPPERT). We partly preserve the binomina given to foliage by UNGER because there is no possibility to check the affinity by cuticular studies. Some of UNGER’s species of alleged legume foliage fall into the category of "Angiosperms incertae sedis" (p. 84 f.). Leguminosites BOWERBANK emend. SCHIMPER CEK comb. nov. Leguminosites hesperidum (UNGER) KOVAR-EDER & Z. KVAC Plate 9, Figures 2-4 1850a Robinia hesperidum UNGER, Gen. spec. pl. foss., p. 245, pro parte - basionym. 1850a Acacia parschlugiana UNGER, p. 494, pro parte. 1864 Robinia hesperidum UNGER - UNGER, p. 21, pro parte, pl. 4, figs. 11, 12, 13 (GBA 1864/01/21, lectotype). 1864 Acacia parschlugiana UNGER - UNGER, p. 35, pro parte, pl. 11, fig. 19 (LMJ 77653). Lectotype designated here: GBA 1864/01/21, figured by UNGER (1864: pl. 4, fig. 13) - refigured on pl. 9, fig. 4. Additional material: NHMW 1878/6/8783, 9109. Pods 10 mm wide and less than 100 mm long, slightly to distinctly contracted between the seeds (up to 6-8 per pod), irregularly densely or widely spaced, even within one pod, rounded to flattened where neighbouring seeds meet, seeds max. 9 mm in diameter. L. hesperidum corresponds with Leguminocarpon type I sensu HABLY (1992: 173). These fruits are rather widely distributed at certain time intervals, e.g. in Hungary and at Oehningen (HEER 1859, pl. 140, fig. 11). We refrained from applying the binomen Acacia parschlugiana for these pods because we reserve it for the compound leaves that have been described together with the pods although never found attached to each other (see below). UNGER (1864: 21) hesitated between Robinia and Acacia in the assignment of these fruits. 74 Annalen des Naturhistorischen Museums in Wien 105 A CEK comb. nov. Leguminosites dionysi (UNGER) KOVAR-EDER & Z. KVAC Plate 9, Figure 5 1850a Cytisus dionysi UNGER, Gen. spec. pl. foss., p. 486 - basionym. 1864 Cytisus dionysi UNGER - UNGER, p. p. 19, pl. 4, fig. 1 (LMJ 76577, lectotype). Lectotype designated here LMJ 76577, figured by UNGER (1864: 19, pl. 4, fig. 1) - refigured on pl. 9, fig. 5. A short slender pod with several (?) seeds. No further material is available. CEK comb. nov. Leguminosites palaeogaeus (UNGER) KOVAR-EDER & Z. KVAC Plate 9, Figure 1 1850a Mimosites palaeogaea UNGER, Gen. spec. pl. foss., p. 494 - basionym. 1864 Mimosa palaeogaea (UNGER) UNGER, p. 34, pl. 11, fig. 12. Neotype designated here: NHMW 2001B0017/3 - figured on pl. 9, fig. 1. Long-stalked pods with several seeds, margins parallel-sided (without contractions). L. palaeogaeus corresponds with Leguminocarpon type VI sensu HABLY (1992: 182, pl. 3, fig. 3) from Magyaregregy. CEK comb. nov. Leguminosites parschlugianus (UNGER) KOVAR-EDER & Z. KVAC Plate 9, Figures 6-7 1850a Bauhinia parschlugiana UNGER, Gen. spec. pl. foss., p. 493 - basionym. 1864 Bauhinia parschlugiana UNGER – UNGER, p. 31, pl. 11, fig. 3. Neotype designated here: NHMW 1878/6/9895 - figured on pl. 9, fig. 7. Material: GBA 2002/01/59, 60, 61; NHMW 1878/6/2820, 2821, 8889, 8890. Pods with two (occasionally three) seeds, sometimes slightly contracted. L. parschlugiana corresponds with Leguminocarpon type IV sensu HABLY (1992: pl. 4, figs. 1-6) and L. mecsekense (ANDREÁNSZKY) HABLY. Similar fruits have been compared with Dalbergia (ANDREÁNSZKY 1955) and with Cladrastis (HERENDEEN 1992 c). This type of pods occurs also in Bavaria (e.g. RIEDERLE & GREGOR 1997: pl. 3, fig. 2). Podocarpium A. BRAUN Podocarpium podocarpum (A. BRAUN) HERENDEEN Plate 9, Figures 8-11 ? 1851a Cassia ambigua UNGER - ETTINGSHAUSEN, p. 27, pl. 5, figs. 12, 13. Material: fruits: GBA 2002/01/27; IBUG Ett. coll. 2245 (II); NHMW 1878/6/8876 (part) + 8877 (counterpart), 9894; leaflets: GBA 2002/01/28, 62-68; NHMW 1878/6/8349, 8878, 8884. Pods of this completely known plant (HERENDEEN 1992 a, b, LIU et al. 2001) are so characteristic that the record at Parschlug is unequivocal. Several leaflets with a characteristic, more prominent basal vein on one side also co-occur. In Europe, the maximal distribution of P. podocarpum falls into Early-Middle Miocene. This legume mostly inhabited gallery forests under subtropical and warm-temperate climates. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 75 Phaseolites UNGER Phaseolites securidacus UNGER Plate 9, Figures 13, 14 1850a Phaseolites securidacus UNGER, p. 488. 1864 Phaseolites securidacus UNGER - UNGER, p. 24, pl. 5, figs. 9 (LMJ 76569, lectotype), 10. Lectotype designated here: LMJ 76569, figured in UNGER (1864: pl. 5, fig. 9) – refigured on pl. 9, fig. 14. Additional material: NHMW 1878/6/2517 (part) + 2518 (counterpart) det. by ETTINGSHAUSEN as Chrysophyllum parschlugianum ETTINGSHAUSEN in sched.; GBA 2002/01/12. Leaflets broadly elliptic, bluntly acuminate, cuneate, sessile. Secondaries numerous, dense, eucamptodromous, distinctly impressed. Leaflets of this kind are common among various representatives of the Papilionoideae. gen. indet. "Acacia" parschlugiana UNGER Plate 9, Figure 12 1850a Acacia parschlugiana UNGER, p. 494 pro parte. 1852 Comptonia laciniata UNGER, p. 33, pl. 16, fig. 8. 1864 Acacia parschlugiana UNGER - UNGER, p. 35, pro parte, pl. 11, fig. 20. Neotype designated here: NHMW 1878/6/9117 - figured on pl. 9, fig. 12. Additional material: GBA 2002/01/69-72, 94b; NHMW 1878/6/9859. Complete, partly disintegrated leaves of this legume probably belong to the Mimosoideae. The fragmentary evidence afforded by this type of foliage prevents us from assigning it to a particular extant genus. The specimen determined by UNGER (1852: 33, pl. 16, fig. 8) as Comptonia laciniata is not available. However, in contrast to UNGER, who believed this fossil to be catkins, we think it constitutes a fragment of "Acacia" parschlugiana. Similar compound leaves also occur at the Randeck Maar (RÜFFLE 1963). "Juglans" parschlugiana UNGER Plate 9, Figures 15, 16 1850a Juglans acuminata A. BRAUN - p. 468, pro parte (non Oehningen = J. acuminata A. BRAUN ex UNGER = Cedrela acuminata (A. BR.) ILJINSKAYA). 1860 Juglans parschlugiana UNGER, p. 37, pro parte, pl. 19, figs. 1, 2 (LMJ 76559, lectotype), 3, 4 (LMJ 76560, syntype), 5, 6, (non 7). Lectotype designated here: LMJ 76559, figured by UNGER (1860: pl. 10, fig. 2) – refigured on pl. 8, fig. 15. Additional material: NHMW 1878/6/9119 det. ETTINGSHAUSEN as Ficus tenuinervis; 1878/6/2569 det. as Juglans parschlugiana UNGER; GBA 2002/01/2. Because of the smaller leaflet size and the rather coriaceous texture of Juglans parschlugiana, we doubt its relationship to the Juglandaceae. Legumes are in our opinion a more appropriate group within which to search for affinities. The fruit, which UNGER (1860: pl. 19, fig. 7) combined with this species, must be excluded because it is an indeterminable fragment. 76 Annalen des Naturhistorischen Museums in Wien 105 A Rhamnaceae Paliurus MILL. Paliurus tiliifolius (UNGER) BŮZZEK Plate 11, Figure 1 ? ? ? ? 1847 1847 1850a 1850a 1850a 1850a 1850a 1864 1864 1864 Paliurus favonii UNGER, p. 147, pro parte, pl. 50, figs. 7, 8 (non 6 left). Ceanothus europaeus UNGER, p. 144, pl. 49, fig. 8. Paliurus favonii UNGER - UNGER, p. 463, pro parte. Ceanothus europaeus UNGER - UNGER, p. 466. Ziziphus tremula UNGER, p. 463. Bauhinia parschlugiana UNGER, p. 493, pro parte. Ziziphus protolotus UNGER, p. 463. Ziziphus tremula UNGER - UNGER, p. 16, pl. 3 fig. 39 (LMJ 76566). Ziziphus renata UNGER, p. 16, pl. 3, figs. 40, 41. Ziziphus protolotus UNGER - UNGER, p. 17, pl. 3, fig. 43. Additional material: GBA 2002/01/36; NHMW 1878/6/8581 + 8582 (part + counterpart), 8584. Leaves of this species, which are regularly found in association with the Paliurus fruits (e.g. in the Lower Miocene of North Bohemia – BŮZEK 1971), are usually bigger and finely serrate: those from Parschlug partly differ, due to local environmental conditions, in their smaller size and sub-entire margins. Judging from the figure of Ziziphus protolotus UNGER (1864), this leaf impression is also an atypical specimen of much smaller size due to ecological conditions. Similar entire leaves of "Quercus" aspera differ in having basal secondaries that do not run that far towards the apex. The type locality of this species is Bílina-Brest’any (UNGER 1847). Because the typification was omitted in the revision by HABLY et al. (2001), we designate here as the lectotype the specimen no. BP 64.306.1 (Natural History Museum Budapest) figured by UNGER (1847) on pl. 49, fig. 2, and refigured by ETTINGSHAUSEN (1869) on pl. 50, fig. 17 and by HABLY et al. (2001) on pl. 85, fig. 1. Paliurus favonii UNGER Plate 11, Figures 2, 3, 7 1847 Paliurus favonii UNGER, p. 147, pro parte, pl. 50, fig. 6 left (LMJ 76518, lectotype) (non 7, 8 = Paliurus tiliaefolius (A. BR.) BŮZEK). 1850a Paliurus favonii UNGER - UNGER, p. 463, pro parte. Lectotype designated here: LMJ 76518, figured by UNGER (1847: pl. 50, fig. 6 left) - refigured on pl. 11, fig. 7. Epitype designated here: NHMW 1878/6/8583 - figured on pl. 11, fig. 3. Additional material: IBUG Ett.coll. 1841, 1842. Parschlug is the type locality of this species, which was widely distributed in Europe during the Tertiary. Due to pyritisation the specimens from Parschlug usually do not show the characteristic trilocular pattern. In all other respects they match the other records of this kind in Europe. Contrary to the treatment of UNGER (1847) and the recommendation of BŮZEK (1971: 74), we separate fruits and leaves into two species because they have never been found attached. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 77 Berchemia DC. Berchemia multinervis (A. BRAUN) HEER Plate 11, Figures 4, 5 Material: NHMW 1878/6/2078, 9107, 9108. Two leaves morphologically corresponding to the type material from Oehningen (HEER 1859: 77, pl. 128, figs. 9-18). The true generic affiliation remains uncertain because the corresponding leaf morphology occurs in different genera among the Rhamnaceae (BŮZEK 1971: 74). The only leaves available all derive from ETTINGSHAUSEN’s level I. Sapindaceae Contrary to the other species, the specimen lists of Acer species are rather complete due to the monographic studies of Acer by STRÖBITZER-HERMANN (2003). Acer L. Acer tricuspidatum BRONN Plate 10, Figures 10-12 1847 1847 1850a 1850a Acer trilobatum ALEX. BRAUN – UNGER, p. 130, pro parte, pl. 41, fig. 6. Acer productum ALEX. BRAUN – UNGER, p. 131, pro parte, pl. 42, fig. 8 (LMJ 76526). Acer trilobatum ALEX. BRAUN – UNGER, p. 450, pro parte. Acer productum ALEX. BRAUN – UNGER, p. 451, pro parte. Additional material: GBA 2002/01/52-54, 55 (part + counterpart); IBUG Ett. coll. 1552-1554, 2809; LMJ 77889, 77892 + 77900A (part + counterpart); NHMW 1845/39/16 + 17 (part + counterpart), 1878/6/2112, 2647A + 2648A (part + counterpart), 8421. This species is rather rare at Parschlug. Leaf forms called "forma tricuspidatum" and "forma productum" occur there along with leaves of intermediate morphology. Leaves having the characteristic morphology of "Acer pyrenaicum" RÉROLLE, which are common in the Late Miocene and Early Pliocene floras (e.g. KVACEK et al. 2002 b), are absent at Parschlug. Acer pseudomonspessulanum UNGER emend. STRÖBITZER-HERMANN Plate 10, Figures 7-9 ? ? 1847 1847 1850a 1850a 1878b 1972 Acer pseudomonspessulanum UNGER, p. 132, pro parte, pl. 43, fig. 2 (LMJ 76522, lectotype). Acer pseudocampestre UNGER, p. 133, pro parte, pl. 43, fig. 6. Acer pseudomonspessulanum UNGER – UNGER, p. 449, pro parte. Acer pseudocampestre UNGER – UNGER, p. 450, pro parte. Acer decipiens A. BRAUN – ETTINGSHAUSEN, p. 89, pro parte, pl. 5, fig. 5 (with Rhytisma aceris ETTINGSHAUSEN; NHMW 1878/6/6295). Acer decipiens AL. BRAUN 1851 sensu novo – WALTHER, p. 121, pro parte, pl. 2, figs. 6 (MMG Parsch. 144/2), 8, pl. 54, fig. 7 (MMG Parsch. 144/2). 78 Annalen des Naturhistorischen Museums in Wien 105 A Lectotype designated here: LMJ 76522, figured by UNGER (1847: pl. 43, fig. 2) - refigured on pl. 10, fig. 8. Additional material: GBA 2002/01/56, 57? (vel Acer integrilobum), 58; IBUG Ett. coll. 1559? (vel Acer integrilobum), one specimen without number; LMJ 77896, 77897? (vel Acer integrilobum), 77899, 77903; NHMW 1878/6/2068, 2399? (vel Acer integrilobum), 3248 (counterpart to 6295), 9156, 9311. Leaves small, palmate, 3-lobed, base usually rounded, lobes deeply incised, rather narrow, nearly of the same length, narrowing continuously towards the acute or obtuse apex, margin entire or sometimes with single small teeth. From the two leaves assigned by UNGER (1847) to Acer pseudomonspessulanum, only that on pl. 43, fig. 2 is accepted as the lectotype. The leaf on pl. 43, fig. 1 belongs to Acer integrilobum. Against UNGER’s opinion, Acer pseudomonspessulanum is not the most common species at Parschlug; this distinction goes to A. integrilobum. Acer integrilobum WEBER sensu WALTHER Plate 10, Figures 1-6 1847 Acer pseudomonspessulanum UNGER, p. 132, pro parte, pl. 43, fig. 1 (LMJ 76531) {non fig. 2 = Acer pseudomonspessulanum UNGER emend. STRÖBITZER-HERMANN] 1850a Acer pseudomonspessulanum UNGER – UNGER, p. 449, pro parte. 1878b Acer decipiens A. BRAUN – ETTINGSHAUSEN, p. 85, 88, 89, pro parte, pl. 2, figs. 1, 2 (bearing Hysterium parschlugianum ETTINGSHAUSEN), pl. 4, fig. 11 (bearing Xylomites aceris decipientis ETTINGSHAUSEN), pl. 5, fig. 1 (bearing Rhytisma aceris ETTINGSHAUSEN; NHMW 1878/6/6594). Within the natural variation of this species, two morphological groups can be distinguished among the studied specimens: Group A (Plate 10, Figures 1-4) Additional material: GBA 2002/01/47 (part + counterpart), 49-51, 57? (vel Acer pseudomonspessulanum); IBUG Ett. coll. 83 + 84 (part + counterpart), 1559? (vel Acer pseudomonspessulanum), 1565 + 1566 (part + counterpart), 1567 + Nr. 195 (part + counterpart), 1568, 2813 + Nr. 190 (part + counterpart); LMJ 77888, 77894, 77897? (vel Acer pseudomonspessulanum); NHMW 1878/6/2327, 2399? (vel Acer pseudomonspessulanum), 2533 + 2534 (part + counterpart), 2585, 6596, 8440? + 8441? (part + counterpart), 8444, 8702. Leaves palmate, 3-lobed; base usually rounded, sometimes obtuse or very rarely cordate (similar to certain leaves of Acer pseudomonspessulanum), the basal part of the centrallobe is quite broad, it narrows abruptly at the upper third and has a characteristic acuminate apex; the side-lobes are nearly as long as the central-lobe, their apex is acuminate or acute; sinus between the lobes usually rounded and wide; entire margin, very seldom single, small teeth. Group B (Plate 10, Figures 5-6) Additional material: GBA 2002/01/48; IBUG Ett. coll. 1259, 1560, 1561 (counterpart to NHMW 1878/6/8445); LMJ 77895, 77898; NHMW 1878/6/2411 + 9157 (part + counterpart), 2412, 2451 (counterpart to 6594), 2544 + 9251 (part + counterpart), 2545, 8445 (counterpart to IBUG Ett. coll. 1561). Leaves smaller than in group A, palmate, 3-lobed; base usually rounded, very rarely obtuse; central lobe much longer and broader than the very small side lobes; apex of the KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 79 lobes acute or attenuate, the tip sometimes rounded; sinus between the lobes rounded and wide; margin entire, sometimes one or two pairs of small teeth on the central lobe and/or on the basal side of the side lobes. A few leaves show an intermediate position between groups A and B and are therefore difficult to group. Acer sp. div. - fructus Plate 10, Figures 13-16 1847 1847 1850a 1850a Acer pseudomonspessulanum UNGER, p. 132, pro parte, pl. 43, figs. 3 (LMJ 76514), 4. Acer pseudocampestre UNGER, p. 133, pro parte, pl. 43, figs. 8, 9. Acer pseudomonspessulanum UNGER - UNGER, p. 449, pro parte. Acer pseudocampestre UNGER - UNGER, p. 450, pro parte. Binomina that were originally applied to leaves cannot be used for detached fruits in the same way, contrary to the opinion of, e.g. UNGER (1847), BŮZEK (1971), TANAI & OZAKI (1977), or TANAI (1983). Classification on the species level of these winged fruits, which are preserved as impressions only, is problematic because the combination with leaf taxa based on reliable feature complexes remains unsolved and the classification of fruits requires preservation of internal structures (cf. MAI 1983, 1984a). Based on differences in nutlet shape and size, on the course of veins in the wings, and on the distance that the nutlets are enclosed by the wing on the ventral side, 3 different formal groups are distinguish within the material from Parschlug. A few fruits fit none of these groups. It remains to be determined whether the aforementioned features are really diagnostic for segregating natural species, since the observed variation in fruits of extant maple species is very large. Form-group 1 (Plate 10, Figure 16): Additional material: IBUG 187, Ett. coll. 1549 + 2873 (part + counterpart), 1550 + 1551 (part + counterpart); NHMW 1853/26/468, 1878/6/8419, 9099, 9242. Nutlet quadrate or roundish, rarely oval, 6-10 mm long; 6-9 mm wide, wing 15-28.5 mm long, max. 6-12 mm wide, at the contact to the nutlet 3-7 mm wide, length-ratio wing : nutlet = 2.5-2.9; end of the wing cut or, towards the dorsal side, rounded, dorsal side straight, ventral side convex, a few veins initially run in a narrow zone parallel to the dorsal side and then approx. upright towards the ventral side, several times dichotomously ramified, interconnected by several cross-anastomoses, divergence-angle 28°50°; wing nearly not enclosing the nutlet on the ventral side. Form-group 2 (Plate 10, Figures 14, ?15): Additional material: IBUG 104, 188, 189, Ett. coll. 1881, 1980, 2803?, 2902, one specimen without number; LMJ 77891, 77902 (part + counterpart); NHMW 1853/26/476, 1878/6/2582, 2708? + 2709? (part + counterpart), 8416, 8420 + 9243 (part + counterpart), 8551, 9097, 9100?, 9254, 9891. Nutlet mostly oval, rarely approx. triangular; 7-12 mm long, 3.5-8 mm wide, wing 1222 mm long, often incomplete, max. 4.5-10 mm wide, often incomplete, at the contact to the nutlet 3.5-5 mm wide, length-ratio wing : nutlet = 1.7-3; end of the wing round- 80 Annalen des Naturhistorischen Museums in Wien 105 A ed, dorsal side straight or slightly convex, ventral side more or less convex, several veins initially run parallel to the dorsal side and then approx. upright or <90° towards the ventral side, several times dichotomously ramified; divergence-angle 25°-58°, wing enclosing about one third of the nutlet's ventral side. Form-group 3 (Plate 10, Figure 13): Additional material: IBUG Ett. coll. 1122 + 1564 (part + counterpart), one specimen without number (counterpart to NHMW 1878/6/8447); NHMW 1878/6/8447 (counterpart to IBUG without number), 9158, 9253. Nutlet roundish, 4-4.5 mm long; 3.5 mm wide, wing 11.5-19 mm long, max. 7.5-8.5 mm wide, sometimes incomplete; at the contact to the nutlet 3-3.5 mm wide; length-ratio wing : nutlet = 2.9-4.8; end of wing rounded, dorsal side mostly straight or rarely somewhat convex, ventral side more or less convex, several veins initially run parallel to the dorsal side and then approx. upright or <90° towards the ventral side, several times dichotomously ramified; divergence-angle (measurable only in one specimen) 60°; wing nearly not enclosing the nutlet on the ventral side. Specimens that do not fit in any of the groups: IBUG Ett. coll. 1557 + 1558 (part + conterpart); LMJ 76514. Anacardiaceae Toxicodendron MILL. CEK & WALTHER Toxicodendron herthae (UNGER) Z. KVAC Plate 9, Figures 17-19 1850a Rhus herthae UNGER – UNGER, p. 473. 1850a Juglans melaena UNGER, p. 470. 1860 Rhus herthae UNGER – UNGER, p. 42, pl. 20, figs. 7, 8 (LMJ 76562, lectotype), 9 (LMJ 76551, syntype). 1860 Juglans melaena UNGER - UNGER, p. 38, pl. 19, figs. 8-10. Lectotype designated here: LMJ 76562, figured by UNGER (1860: pl. 20, fig. 8) – refigured on pl. 9, fig. 17. Additional material: NHMW 1878/6/2027 det. by ETTINGSHAUSEN as Juglans parschlugiana, 1878/6/9252 + 2543 (part + counterpart) det. as Acer decipiens. The species concept for Fagus herthae (with the basionym of Rhus herthae UNGER 1849) that was proposed by ILJINSKAYA (1962 and 1964) is not based on the original diagnosis and material from Parschlug. It must be rejected along with the therein-selected type specimen. The nomenclature of this species has been discussed in detail by KVACEK & WALTHER (1998: 27). The nearest living relative has not yet been established. ? Cotinus MILL. CEK comb. nov. Cotinus (?) aizoon (UNGER) KOVAR-EDER & Z. KVAC Plate 11, Figures 6, 8-10 1847 Rhamnus aizoon UNGER, Chlor. prot., p. 146, pro parte, pl. 50, figs. 1 right, 2 - basionym. 1850a Rhamnus aizoon UNGER – UNGER, p. 464. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 1850a 1864 1864 1878b 81 Celastrus cuneifolius UNGER, p. 459. Rhamnus aizoon UNGER – UNGER, p. 17, pl. 3, figs. 44 (LMJ 76575, lectotype), 45, 46. Pittosporum cuneifolium (UNGER) UNGER, p. 6, pl. 1, figs. 14, 15. Rhamnus aizoon UNGER - ETTINGSHAUSEN, p. 86, pl. 3, fig. 9 (NHMW 1878/6/6553 bearing Xylomites rhamni aizoonis ETTINGSHAUSEN). Lectotype designated here: LMJ 76575, figured by UNGER (1864: pl. 3, fig. 44) – refigured on pl. 11, fig. 8. Additional material: GBA 2002/01/8, 11, 74, ? 75; LMJ 77607. Broadly obovate leaves, petiolate, leaf apex emarginate and slightly mucronate, secondary veins camptodromous, relatively densely spaced. SAPORTA (1865: 352 (208), pl. 12, fig. 6 (erronously 7 in the text)) described a similar leaf from the Oligocene of Armissan as Rhus palaeocotinus SAPORTA and mentioned that the leaf apex is not characteristic of the extant nearest relative Rhus cotinus L. (now Cotinus cogyggria SCOP.). Due to poor preservation we are unable to discern the marginal vein that joins all secondaries in the foliage of extant Cotinus, which would help to corroborate the suggested affinity. A similar but finely toothed and short, petiolate leaf was discovered under GBA 2002/01/73. This specimen weakens the probability that the entity as circumscribed above belongs to Cotinus. The specimen figured by UNGER (1847: pl. 3, fig. 44) should be in the collection LMJ but is at present missing. Simaroubaceae Ailanthus DESF. CEK comb. nov. Ailanthus pythii (UNGER) KOVAR-EDER & Z. KVAC Plate 14, Figures 2-5 1850a 1850a 1850a 1850a 1850b Sapindus pythii UNGER, Gen. spec. pl. foss., p. 457 - basionym. Juglans elaenoides UNGER, p. 469, pro parte. Quercus zoroastri UNGER, p. 401, pro parte. Rhus elaeodendroides UNGER, p. 474. Juglans elaenoides UNGER – UNGER, p. 179, pro parte, pl. 53 fig. 3 (LMJ 76542 + 77652, part + counterpart). 1852 Quercus zoroastri UNGER – UNGER, p. 36, pro parte, pl. 18, fig. 9, (non 7 and 8). 1860 Sapindus pythii UNGER – UNGER, p. 33, pl. 14, figs. 6, 7, 8 (LMJ 76557, lectotype), 9-17. 1860 Fraxinus primigenia UNGER – UNGER, p. 22, pro parte, pl. 8, figs. 3, 8 ?. 1860 Rhus elaeodendroides UNGER – UNGER, p. 45, pro parte, pl. 21, figs. 4, 5, 11. 1878b Sapindus pythii UNGER – ETTINGSHAUSEN p. 85, pl. 3, fig. 5 (bearing Sphaeria palaeo-sapindi ETTINGSHAUSEN, NHMW 1878/6/6484). Lectotype designated here: LMJ 76557, figured by UNGER (1860: pl. 14, fig. 8) - refigured in pl. 14, fig. 4 Additional material: NHMW 1878/6/2649 + 2650 (part + counterpart) determined by ETTINGSHAUSEN as Pistacia lentiscus, 1878/6/2031, 2525a, 2527, 2554 a. GBA 2002/01/9,10. Leaflets of the same morphology were described recently as Ailanthus mecsekensis HABLY (2001) from a fossiliferous layer at Magyaregregy with a mass occurrence of the Ailanthus confucii fruits. We believe that A. mecsekensis is conspecific with A. pythii. 82 Annalen des Naturhistorischen Museums in Wien 105 A The relationship to Ailanthus is also well documented by long petiolules, although fruits of the genus have been rarely documented at Parschlug. At Parschlug, Fraxinus fruits are also rarely preserved in association with the above leaflets and compound leaves, but the irregularly toothed to almost entire leaf margin and very long petiolules are certainly not typical for foliage of ash. Ailanthus confucii UNGER Plate 11, Figure 11 Material: NHMW 1878/6/2121, 2606. The specimens available are both incomplete fruits of Ailanthus. They correspond, as far as we can judge, to the populations from Erdőbénye and Magyaregregy (HABLY 2001). Oleaceae Fraxinus L. Fraxinus primigenia UNGER Plate 11, Figures 12-15 1850a Fraxinus primigenia UNGER, p. 431, pro parte. 1860 Fraxinus primigenia UNGER - UNGER, p. 22, pro parte, pl. 8, fig. 1. Neotype designated here NHMW 1878/6/8155 – figured on pl. 11, fig. 13. Additional material: NHMW 1878/6/8156, 9889; IBUG Ett. coll. 1384, 1385 + 1386 (part + counterpart) det. by ETTINGSHAUSEN as Fraxinus pachyptera, 1387 as Fraxinus praeexcelsior. Ash fruits are recorded at many sites of the European Tertiary, particularly in riparian assemblages, e.g. KOVAR-EDER & KRAINER (1991). In our opinion, their morphology does not allow separation to the species level. The associated leaves have not yet been recognised among the leaf fossils of Parschlug. Ash leaflets or complete leaves have often been misinterpreted for the Juglandaceae (KVACEK & HURNÍK 2000), mostly as Juglans bilinica UNG. Apocynaceae Nerium L. Nerium sp. Plate 11, Figures 16-18 Material: NHMW 11878/6/7177, 8173, 8175. Slender, coriaceous leaves, width 10-25 mm, incomplete length max. 70 mm, base cuneate decurrent; petiole stout, straight, leaf apex missing, margin entire at the base, in the upper parts slightly wavy, midvein stout, straight, secondaries very densely spaced, thin, of almost the same thickness, originating at wide angles, running parallel, looping each other near the margin. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 83 Similar leaves are known from the European Neogene and are usually compared with or assigned to Nerium oleander, e.g. RIEDERLE & GREGOR (1997: pl. 11, figs. 1-5 – as aff. Nerium sp.) from Kirrberg, Bavaria, Upper Freshwater Molasse, Early-Middle Miocene, PALAMAREV & PETKOVA (1987: 141, pl. 36, fig. 6) from Rucinzi, Bulgaria, Sarmatian (Central Paratethys stage), Middle Miocene; GIVULESCU (1962: 165, fig. 180) - Valea Neagra, Pannonian, Late Miocene. BERGER (1952: 105) described foliage as Nerium bilinicum ETTINGSHAUSEN from Vösendorf (Vienna, Pannonian E, Late Miocene). The latter binomen is inappropriate because the type specimen of N. bilinicum derives from Kuclín (Late Eocene) and its venation does not correspond to that of Nerium (HABLY et al. 2001: pl. 23, fig. 6). The specimens from Vösendorf were not available for reinvestigation. Although the venation pattern seems to be very similar to that of Nerium, we are unable to decide whether an intramarginal vein, which would point towards the Lythraceae (Decodon, see KVACEK & SAKALA 1999), exists. Although sometimes resembling "Quercus" daphnes due to the coriaceous lamina and the dense secondary venation, Nerium can be distinguished by the narrow cuneate decurrent leaf base and the almost equally thick secondaries, whereas in "Quercus" daphnes the intersecondaries between the secondaries can be clearly distinguished. In the ETTINGSHAUSEN collection IBUG, a cylindrical capsule narrow elongate and slightly bent in shape (Ett. coll. 1405) has been determined by ETTINGSHAUSEN as Catalpa europaea n. sp. It may belong to Nerium as well. Monocotyledoneae Smilacaceae Smilax L. Smilax sagittifera HEER emend. HANTKE Plate 11, Figures 19, 20 1847 Smilacites sagittata UNGER, p. 129, pl. 40, fig. 4. 1850a Smilacites sagittata UNGER - UNGER, p. 317. 1851 Smilax sagittata (UNGER) A. BRAUN in STIZENBERGER, p. 75 (non Smilax sagittata HAMILTON). Material: GBA 1847/03/20; IBUG Ett. coll. 400; NHMW 1878/6/7190 + 7191 (part + counterpart) det. by ETTINGSHAUSEN as Smilax grandifolia; IBUG Ett. coll. 399 as Smilax cf. sagittifera. Most specimens from Parschlug match in lamina shape the morpho-species as crrcumscribed from Oehningen (HANTKE 1954). Although the cuticle was separated by a routine maceration from specimen NHMW 1878/6/ 7190 + 7191, diagnostic structures including the cell structure and stomata are not preserved. The specimen IBUG Ett.coll. 399 can be assigned to Smilax based on the venation pattern. However, the characteristic cordate base is not developed. Although the type specimen is not available, we see no reason why ILJINSKAYA & SHTEFYRTSA (1971: 180) hesitated to assign Smilacites sagittatus UNGER (1847: pl. 40, fig. 4) directly to the genus Smilax. 84 Annalen des Naturhistorischen Museums in Wien 105 A Monocotyledoneae gen. et sp. indet. 1850a 1850a 1852 1852 Cyperites tertiarius UNGER, p. 313. Sparganium acheronticum UNGER, p. 327. Cyperites tertiarius UNGER - UNGER, p. 14, pl. 5, fig. 5 (LMJ 76511). Sparganium acheronticum UNGER - UNGER, p. 17, pl. 7, fig. 2. Additional material: NHMW 1878/6/2675. These monocotyledonous leaf fragments lack diagnostic features. The scarcity of monocotyledonous leaves at Parschlug is remarkable. Angiosperms incertae sedis fam. et gen. indet. "Celastrus" europaea UNGER Plate 12, Figures 1, 2 1850a Celastrus europaeus UNGER, p. 459. 1864 Celastrus europaeus UNGER - UNGER, p. 10, pl. 2, figs. 10 (LMJ 76576, lectotype), 11 (LMJ 76581, syntype), 12 (LMJ 76563, syntype), 13. Lectotype designated here: LMJ 76576, figured by UNGER (1864: pl. 2 fig. 10) - refigured on pl. 12 fig. 1. Morphologically, these leaves correspond with Dicotylophyllum deichmuelleri Z. KVACEK & WALTHER (1998: 14). As the epidermal structures are not preserved on the specimens from Parschlug, we refrain from merging them with this species from the Lower Oligocene of North Bohemia. Smaller leaves of "Celastrus" europaea are even more reminiscent of narrower leaves of "Evonymus" latoniae UNGER mentioned below. "Cornus" ferox UNGER Plate 12, Figures 6, 7 1850a 1851a 1864 1866 1878b Cornus ferox UNGER, p. 441, pro parte. Pterospermum ferox ETTINGSHAUSEN, p. 22, pl. 4, fig. 4. Hardenbergia orbis veteris UNGER, p. 23, pl. 5, fig. 5. Cornus ferox UNGER - UNGER, p. 76, pl. 24, fig. 21. Aristolochia parschlugiana ETTINGSHAUSEN, p. 88, pl. 4, fig. 9 (NHMW 1878/6/6566 + 8109 part + counterpart, bearing Xylomites aristolochiae) (nomen). Neotype designated here: NHMW 1878/6/6566 + 8109 (part + counterpart), figured by ETTINGSHAUSEN (1878 b: pl. 4, fig. 9) - refigured on pl. 12, fig. 6. Although no exact affinities can currently be suggested, we reject the affinity to the Cornaceae because of the slightly cordate leaf base and the basal origin of the first pair of secondaries. Aristolochia is also rather unlikely. So-called Aristolochia aesculapi HEER has been transferred by BŮZEK (1971) to Diversiphyllum (? Convolvulaceae). KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 85 "Evonymus" latoniae UNGER Plate 12, Figures 3-5 1850a Evonymus latoniae UNGER, p. 460. 1864 Evonymus latoniae UNGER - UNGER, p. 11, pl. 2, fig. 25 (LMJ 76574 + 76573, part + counterpart, lectotype). Lectotype designated here: LMJ 76574 + 76573, (part + counterpart), figured by UNGER (1864: pl. 2, fig. 25) - refigured on pl. 12 fig. 3. Additional material: NHMW 1878/6/2063 det. by ETTINGSHAUSEN as Rhus elaeodendroides UNGER, 1878/6/2742 as Celastrus sp. nov., 1878/6/9176 as Celastrus europaeus UNGER. Marginal teeth glandular, regularly spaced, of almost equal size. Superficially, these leaves resemble "Celastrus" europaeus, but they differ in the leaf margin as indicated above. "Quercus" daphnes UNGER Plate 12, Figures 10-15 ? ? ? 1847 1847 1847 1850a 1850a 1850a 1850a 1850a 1866 1866 1866 1878b Quercus chlorophylla UNGER, p. 111, pl. 31, fig. 1. Quercus daphnes UNGER, p. 112, pl. 31 figs. 2, 3 (LMJ 76525, lectotype). Quercus elaena UNGER, p. 112, pl. 31, fig. 4. Quercus daphnes UNGER - UNGER, p. 402. Quercus chlorophylla UNGER – UNGER, p. 402. Quercus elaena UNGER - UNGER, p. 402. Achras lycobroma UNGER, p. 435. Rhododendron flos saturni UNGER, p. 440. Achras lycobroma UNGER, p. 23, pro parte, pl. 8, fig. 1 (LMJ 76591). Rhododendron flos saturni UNGER - UNGER, p. 38, pl. 12 fig. 15 (LMJ 76590 counterpart). Myrsine doryphora UNGER - UNGER, p. 19, pl. 6, fig. 10. Quercus daphnes UNGER - ETTINGSHAUSEN, p. 86, pl. 3, fig. 8 (NHMW 1878/6/6549 + 9457 part + counterpart), pl. 4, fig. 5 (NHMW 1878/6/6550 bearing Xylomites daphnes ETTINGSHAUSEN). Lectotype designated here: LMJ 76525, figured by UNGER (1847: pl. 31, fig. 3) – refigured on pl. 12, fig. 15. Epitype designated here: LMJ 76590 - counterpart of the specimen figured by UNGER (1866: pl. 12, fig. 15) - refigured on pl. 12, fig. 11 a, b. Additional material: GBA 1847/03/10, 2002/01/13; IBUG Ett. coll. 966; NHMW - all det. by ETTINGSHAUSEN - NHMW 1845/39, 1878/6/2375, 7557 as Quercus chlorophylla UNGER, NHMW 1878/6/2774, 9425, 9455, 9459 as Quercus daphnes UNGER, 1878/6/9460; 1878/6/8234 as Sapotacites longepetiolatus ETT., 1878/6/7850 as Laurus palaeo-benzoin ETTINGSHAUSEN. Leaves with characteristically densely spaced secondaries and intersecondaries. The characteristic venation is poorly visible in the specimen of Q. chlorophylla (UNGER 1847: 111, pl. 31, fig. 1), which is not available. Leaves with similar venation pattern can be found in different families such as Apocynaceae, Rosaceae, Myrsinaceae, and Sapotaceae but not in the Fagaceae. 86 Annalen des Naturhistorischen Museums in Wien 105 A Antholithes BRONGNIART CEK sp. nov. Antholithes stiriacus KOVAR-EDER & Z. KVAC Plate 15, Figures 13-15 1850a Celastrus elaenus UNGER, p. 459, pro parte. 1864 Prinos hyperboreus UNGER - UNGER, p. 14, pro parte, pl. 3, fig. 34 a, b. Holotype designated here: NHMW 1878/6/9870 det. by ETTINGSHAUSEN as Smilax grandifolia – figured on pl. 15, fig. 15. Additional material: IBUG Ett. coll. 415, 427, 432 det. by ETTINGSHAUSEN as Asterocalyx stiriacus. Inflorescence (? partial) loosely botryoidal, flower short stalked, actinomophic, octomeric, half-epigynous (?), calyx shortly synsepalous, cup-like, 3-5 mm long, free tips of sepals very narrow elliptic, blunt at the apex, about 1-2 mm long and 0.3-0.5 mm wide, corolla very shortly sympetalous (?) or choripetalous, 5-6 mm in diameter, petals nearly lineal to narrowly obovate, about 2.5 mm long and 0.5 mm wide. UNGER (1850a) assigned these flower remains to Celastrus and later changed his mind and associated them with Prinos (i.e. Ilex) hyperboreus, which he initially based on leaves only (UNGER 1850a: 462, 1864: 14). ETTINGSHAUSEN (in his catalogue IBUG) supposed these remains to belong to Smilax. He described as Asterocalyx stiriacus (ETTINGSHAUSEN 1888) a heterogenous entity based on leaves and flowers from Leoben-Münzenberg (type material is missing). The leaf figured there on pl. 3, fig. 4 is Smilax. The inflorescence and flower remains on figs. 1 to 3 belong to the species described above. ETTINGSHAUSEN (1890: 83) described the same type of flowers from Schönegg and compared them with the Dioscoreaceae. None of the suggested groups are similar in the composition of flowers. The Celastraceae usually have 4- or 5-merous flowers, the Smilacaceae and Dioscoreaceae differ also in the flower diagram * P 3+3 or P (3+3) A 3+3, as is often the case among monocotyledons. Of the dicotyledonous families that come into question, the Aquifoliaceae sometimes have more than 4-5-merous flowers. The Sapotaceae contain genera with small octomeric flowers, e.g. * K (4)+(4) C (4)+(4) A 8+8+8 G (8) in Madhuca GMEL. (PENNINGTON 1991), which may correspond to A. stiriacus. Antholithus sp. from the Randeck Maar (GREGOR 1986: pl. 4, fig. 12) and Kirrberg near Balzhausen (RIEDERLE & GREGOR 1997: pl. 3, fig. 11, pl. 5, fig. 5) belongs to the same species. Also at Magyaregragy the same type of flowers is abundant (HABLY, personal communication). We lack evidence of pollen in situ, which would certainly help to decipher the affinities. The generic name Asterocalyx proposed by ETTINGSHAUSEN (1888) endangers the name in current use of the genus Astrocalyx MERRIL (Melastomataceae), being its early orthographic variant. In our opinion it should be proposed into the list of nomina rejicienda. We suggest to typify the genus Asterocalyx ETTINGSHAUSEN by the leaf (A. stiriacus ETTINGSHAUSEN 1888: pl. 3, fig. 4) and to include it into the synonymy of Smilax. Cypselites HEER Cypselites sp. Plate 15, Figure 16 Material: IBUG Ett. coll. 1374. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 87 Narrow elongated seed, 6 mm long, longitudinally striate, with a slender, 1-mm-long stalk. Coma not preserved. Such seeds were previously described as fruits of Compositae (HEER 1859: 2). Later, REID & CHANDLER (1926) recognised in such remains the seeds of the Apocynaceae. The generic name Cypselites HEER has priority over Apocynospermum E.E.M. REID & CHANDLER. Saportaspermum MEYER & MANCHESTER Saportaspermum sp. Plate 15, Figures 6-8 1850a Robinia hesperidum p. 487, pro parte. 1864 Robinia hesperidum UNGER - UNGER, p. 21, pro parte, pl. 4, fig. 14. Additional material: GBA 2002/01/30, 33, 97-98; IBUG Ett. coll. 1343, 1346, 1349 +1350 (part+counterpart) det. by ETTINGSHAUSEN as Embothrium parschlugianum ETTINGSHAUSEN, 1357 as Embothrium postsotzkianum ETTINGSHAUSEN, 1358 as Embothrium subboreale ETTINGSHAUSEN. NHMW 1878/6/2796, 2797, 8018, 8023 + 8024 (part + counterpart), 8025, 8028, 8029, 8033, 8034, 8035, 8036, 9904 det. by ETTINGSHAUSEN as Embothrium parschlugianum ETTINGSHAUSEN, 1878/6/8002 and 8003 as Hakea parschlugiana ETTINGSHAUSEN, 1878/6/8014, 8017 as Embothrium affine ETTINGSHAUSEN. Winged seeds of this kind are widespread in Europe, starting from the Eocene. This morpho-genus was established in North America and typified by Saportaspermum occidentale MEYER & MANCHESTER (1997). The population from Parschlug is represented by numerous specimens and is morphologically more variable; it clearly belongs to another species. The same kind of seeds was described by ETTINGSHAUSEN (1890) from Schönegg (and Parschlug) as Embothrium parschlugianum. ETTINSGAUSEN (1888) has referred to some other seeds of similar morphology and slightly different size and shape as Embothrium sotzkianum, E. salicinum and other binomina from the Leoben area. The whole set of these morpho-taxa requires a broader study within Europe to set limits between the species. The affinities of these enigmatic winged seeds remain doubtful. ? Chaneya WANG & MANCHESTER ? Chaneya sp. Plate 12, Figures 8, 9 Material: IBUG Ett. coll. 2983, 2984, (both level II); NHMW 1878/6/8741, 8742. Pentamerous calyces, partly incomplete, without fruits preserved, obviously epigynous. Sepals entire, sessile, only shortly fused, narrow elliptic, parallel-sided in some cases, 3-6 mm wide and 14 to 20 mm long. Venation reticulate, consisting of narrow elongate meshes between several closely spaced primaries. In Europe, such fossils have usually been assigned to Porana (e.g. HEER 1859, ETTINGSHAUSEN 1888: pl. 9, fig. 19) and Monotes (WEYLAND 1937). A detailed study of more complete material from the Tertiary of Asia and North America (WANG & MANCHESTER 2000) revealed basic discrepancies between the morphology of the fossil fruits and the mentioned genera, which resulted in the erection of a new fossil genus 88 Annalen des Naturhistorischen Museums in Wien 105 A Chaneya WANG & MANCHESTER. These authors found notable similarities with certain members of the Simaroubaceae, namely Picrasma BL. (W. Himalayas to Japan and Fiji). The fossils described above may belong to Chaneya, but a more detail study of the European Chaneya-like fossils is required, including the type material of "Porana" from Oehningen (HEER 1859). Dicotylophyllum SAPORTA Dicotylophyllum sp. 1 Plate 15, Figure 1 Material: NHMW 1878/6/2091. A twig with three leaves, two of them basal fragments, the third 52 mm long and 17 mm wide, short-petiolate, base acute/cuneate, shape of lamina slender elliptic to slightly obovate, lamina basally entire-margined, in the upper part widely, simply, bluntly serrate, sinus acute; venation semicraspedodromous, poorly preserved, secondaries densely spaced, indistinct, originating at wide angles, looping near the leaf margin; loops connected with the teeth by veinlets. Affinities doubtful. Dicotylophyllum sp. 2 Plate 15, Figures 2, 3 1878b Quercus serra UNGER - ETTINGSHAUSEN, p. 86, pro parte, pl. 4, fig. 7 (NHMW 1878/6/6555). Additional material: GBA 2002/01/109. Narrow oblong, slender leaves (? leaflets), length 46 mm (complete specimen) and 60 mm (fragment), width 8 and 16 mm; base acute, slightly asymmetrical, probably sessile, apex attenuate; margin regularly, densely serrate, tooth apices rounded; midvein straight, secondaries poorly preserved, thin, relatively densely spaced, angles of origin 30- 40°. Only two specimens have been discovered among all the investigated material. The morphology is reminiscent of Sorbus species with compound leaves, but the rounded teeth of the fossils are different. They are less comparable with Prinsepia foliage. Dicotylophyllum sp. 3 Plate 15, Figures 9, 10 Material: GBA 2002/01/20; NHMW 1878/6/8571. Two obovate/spatulate leaves, shortly petiolate, base cuneate/decurrent, apex rounded, length 27 and 29 mm, width 11 and 13 mm, leaf margin basally entire, in the apical part shallowly crenulate; primary vein straight, secondaries indistinct, densely spaced, running rather steep across the lamina. These leaves designated here as Dicotylophyllum sp. 3 recall Celastrus noatica UNGER (1864: 7, pl. 2, figs. 2, 3), but the shape of the lamina is more elliptical and the secondary veins are more distinct in the latter. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 89 Dicotylophyllum sp. 4 Plate 15, Figure 11 Material: GBA 2002/01/21. Leaf lamina sub-orbiculate, 90 mm long (but base and apex missing), 72 mm wide; venation acrodromous (whether basal or subbasal cannot be decided), simple craspedodromous, margin simply to occasionally double serrate, teeth acuminate, from the (sub)basal lateral main veins numerous veins originate at the basal part and run towards the margin, ending in the primary teeth, occasional secondary teeth present; tertiaries forked-percurrent. This leaf cannot be assigned to Platanus leucophylla because the marginal teeth are not hooked, are relatively regularly spaced and the leaf is not trilobate. The teeth are reminiscent of Davidia, but the acrodromous venation is not characteristic of this genus. Also Vitaceae foliage comes into question. Similar large leaves, which clearly belong to Celtis, occur at Bílina (personal observation of Z. KVACEK). Dicotylophyllum sp. 5 Plate 15, Figure 12 Material: NHMW 1878/6/7507. Upper part of an elongate lamina, 90 mm long, 33 mm wide, apex attenuate (although not complete), leaf margin simple serrate, apically crenate rather than serrate, teeth widely but relatively regularly spaced and small, sometimes sharp; venation (semi) craspedodromous, midvein straight, secondaries widely spaced (up to 13 mm), running relatively steep across the lamina (35-45°), sometimes forking at variable distances from the midrib, their branches either forming loops near the margin or sending veinlets into the teeth or directly entering the teeth. It cannot be excluded that this fragment represents Fraxinus foliage. However, the true affinities are obscure. Dicotylophyllum sp. 6 Plate 15, Figures 4, 5 Material: IBUG Ett. Coll. 1083, 1084. Two apical fragments of elongate slender leaves, apex attenuate, margin regularly simply toothed, basal side of the teeth convex, apical side convex-concave, tooth apex acute or rounded; venation simple craspedodromous, secondaries originating at an angle of 45-55° from the midvein, tertiaries forked-percurrent. In our opinion the similarity of these fragments to Myrica lignitum is only superficial and their true affinity remains obscure. Dicotylophyllum sp. div. All those leaves which offer no distinct morphological characters to be safely recognized again elsewhere and still were given species names and described by UNGER, are united under this heading. In all these cases we were unable to discern useful morpho-types of foliage. 90 Annalen des Naturhistorischen Museums in Wien 105 A 1850a Amorpha stiriaca UNGER, p. 486, pro parte. 1864 Amorpha stiriaca UNGER – UNGER, p. 20, pro parte, pl. 4, fig. 5 (leaf). 1850a Andromeda glauca UNGER, p. 438. 1866 Andromeda glauca UNGER – UNGER, p. 35, pl. 12, fig. 1. 1850a Apocynophyllum lanceolatum UNGER, p. 434. 1866 Myrsine doryphora UNGER, p. 19, pl. 6, fig. 10. 1850a Azalea hyperborea UNGER, p. 440. 1866 Azalea hyperborea UNGER – UNGER, p. 40, pl. 12, figs. 21 (LMJ 76583), 22. 1850a Capparis ogygia UNGER, p. 443. 1864 Physolobium kennedyaefolium UNGER, p. 22, pl. 5, fig. 1. 1850a Cassia ambigua UNGER, p. 492. 1864 Cassia ambigua UNGER – UNGER, p. 29, pl. 10, fig. 9. 1850a Cassia memnonia UNGER, p. 492. 1864 Cassia memnonia UNGER – UNGER, p. 29, pl. 10, figs. 4, 5. 1850a Celastrus elaenus UNGER, p. 459. 1864 Celastrus elaenus UNGER – UNGER, p. 10, pl. 2, figs 16-19. The specimens shown in UNGER (1864: figs. 18, 19) may represent small leaves of Myrica lignitum. 1850a Celastrus cassinefolius UNGER, p. 459, pro parte. 1864 Celastrus cassinefolius UNGER, p. 7, pl. 2, fig. 1. 1850a Celastrus cassinefolius UNGER, p. 459, pro parte. 1864 Celastrus noaticus UNGER – UNGER, p. 7, pl. 2, figs. 2, 3 (LMJ 76539). 1850a Cotoneaster andromedae UNGER, p. 482. 1866 Cotoneaster andromedae UNGER – UNGER, p. 59, pl. 18, fig. 11 (LMJ 76586), 12 (LMJ 76585). 1866 Cotoneaster pusillus UNGER, p. 59, pl. 18, fig. 13. 1850a Glycyrrhiza blandusiae UNGER, p. 486, pro parte. 1864 Glycyrrhiza blandusiae UNGER – UNGER, p. 20, pro parte, pl. 4, figs. 8-10. 1864 Ilex simularis UNGER, p. 13, pl. 3, fig. 14. 1850a Ledum limnophilum UNGER, p. 441. 1866 Ledum limnophilum UNGER – UNGER, p. 40, pl. 12, figs. 25, 26. 1850a Myrica deperdita UNGER, p. 395. 1852 Myrica deperdita UNGER – UNGER, p. 32. 1850a Myrtus miocenica UNGER, p. 480. 1866 Myrtus miocenica UNGER, p. 57, pl. 18, fig. 6. 1850a Nemopanthes angustifolius UNGER, p. 462. 1864 Nemopanthes angustifolius UNGER – UNGER, p. 15, pl. 3, fig. 35 (LMJ 76567). 1850a Phaseolites orbicularis UNGER, p. 488. 1864 Physolobium orbiculare (UNGER) UNGER, p. 22, pl. 5, fig. 3. 1850a Phaseolites physolobium UNGER, p. 488. 1864 Physolobium antiquum UNGER, p. 21, pl. 5, fig. 4. 1850a Phaseolites serratus UNGER, p. 488. 1850a Pistacia lentiscoides UNGER, p. 473. 1860 Pistacia lentiscoides UNGER – UNGER, p. 46, pl. 21, fig. 14. 1847 Ilex parschlugiana UNGER, p. 148, pl. 50, fig. 8. KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 91 1850a Ilex parschlugiana UNGER – UNGER, p. 461. 1850a Ilex stenophylla UNGER, p. 461. 1864 Ilex stenophylla UNGER – UNGER, p. 14, pl. 3, figs. 15-19. The specimens shown in UNGER (1864: figs. 15, 16) may constitute Myrica lignitum. 1850a Prunus atlantica UNGER, p. 484, pro parte. 1866 Prunus atlantica UNGER – UNGER, p. 61, pro parte, pl. 18, fig. 26. 1850a Prunus paradisiaca UNGER, p. 484, pro parte. 1866 Prunus paradisiaca UNGER – UNGER, p. 62, pro parte pl. 18, fig. 29 (leaf). 1850a Pyrus euphemes UNGER, p. 481. 1850b Pyrus euphemes UNGER – UNGER, p. 183, pl. 59, fig. 10 (LMJ 76548). 1850a Pyrus minor UNGER, p. 481. 1850b Pyrus minor UNGER – UNGER, p. 183, pl. 59, fig. 16 (LMJ 76547). 1850a Pyrus theobroma UNGER, p. 481. 1850b Pyrus theobroma UNGER – UNGER, p. 183, pl. 59, fig. 5 (LMJ 76550). 1852 Quercus gmelini A. BRAUN – UNGER, p. 36, pl. 18, fig. 10. 1847 Quercus hamadryadum UNGER, p. 110, pl. 30, fig. 8. 1850a Quercus hamadryadum UNGER – UNGER, p. 400. 1852 Quercus myricaefolia UNGER, p. 37, pl. 18, fig. 12. 1850a Quercus myrtilloides UNGER, p. 404, pro parte. 1852 Quercus myrtilloides UNGER – UNGER, p. 38, pro parte, pl. 18, fig. 18 (LMJ 76490), 19, 20. 1850a Rhamnus aizoides UNGER, p. 464. 1864 Rhamnus aizoides UNGER – UNGER, p. 17, pl. 3, fig. 47 (LMJ 76565). 1850a Rhamnus degener UNGER, p. 464. 1864 Rhamnus degener UNGER – UNGER, p. 18, pl. 3, fig. 49. 1850a Rhamnus pygmaeus UNGER, p. 465. 1864 Rhamnus pygmaeus UNGER – UNGER, p. 18, pl. 3, fig. 48. 1850a Rhus cuneolata UNGER, p. 474. 1860 Rhus cuneolata UNGER – UNGER, p. 44, pl. 20, fig. 12 (LMJ 76553). 1850a Rhus elaeodendroides UNGER, p. 474. 1860 Rhus elaeodendroides UNGER – UNGER, p. 45, pro parte, pl. 21, figs. 1-3, 6-8 (LMJ 76558), 9, 10 (LMJ 76556). This taxon is a highly heterogeneous group. 1850a Rhus napearum UNGER, p. 474. 1860 Rhus napearum UNGER – UNGER, p. 43, pl. 20, fig. 11 (LMJ 76561). 1850a Rhus retine UNGER, p. 475. 1860 Rhus retine UNGER – UNGER, p. 43, pl. 20, fig. 10. Although this leaf is reminiscent of Lauraceae we think the secondaries are too regular and too densely spaced. 1850a Rhus zanthoxyloides UNGER, p. 474. 1860 Rhus zanthoxyloides UNGER – UNGER, p. 45, pl. 21, fig. 13 (LMJ 76552). This leaf may belong to Myrica lignitum. 1850a Robinia hesperidum UNGER, p. 487, pro parte. 1864 Robinia hesperidum UNGER – UNGER, p. 21, pro parte, pl. 4 figs. 15-17. 92 Annalen des Naturhistorischen Museums in Wien 105 A 1850a Sideroxylon hepios UNGER, p. 434. 1866 Sideroxylon hepios UNGER – UNGER, p. 24, pl. 8, fig. 4 (LMJ 76587). No venation is visible except the midvein. 1847 Ulmus quercifolia UNGER, p. 96, pl. 25, fig. 5. 1850a Ulmus quercifolia UNGER – UNGER, p. 411. 1850a Vaccinium chamaedrys UNGER, p. 439. 1866 Vaccinium chamaedrys UNGER, p. 36, pl. 12, fig. 1a. 1850a Vaccinium empetrites UNGER, p. 440. 1866 Vaccinium empetrites UNGER – UNGER, p. 37, pl. 12, figs. 2a (LMJ 76588), c. 1850a Vaccinium icmadophilum UNGER, p. 439. 1866 Vaccinium icmadophilum UNGER – UNGER, p. 37, pl. 12, fig. 5 a, b. 1850a Vaccinium myrsinefolium UNGER, p. 439. 1866 Vaccinium myrsinefolium UNGER – UNGER, p. 38, pl. 12, fig. 6 (LMJ 76589). 1850a Vaccinium vitis japeti UNGER, p. 439. 1866 Vaccinium vitis japeti UNGER – UNGER, p. 36, pl. 12, fig. 3 a-c. Besides the leaf remains listed above, there is a greater number of dubious and indeterminable objects referred to by UNGER as remains of various generative organs, which are not treated in detail in the present account. For the lists of taxa according to the previous publications (nomina nuda excluded) and current revisions see tables 4-11. Taphonomy SACHSENHOFER et al. (2002) determined analogies in the development of the basins along the Norian depression: basal alluvial deltaic sediments are typically followed by one coal seam which is overlain by fine-grained, lacustric or sometimes even brackish sediments (as in the Fohnsdorf basin); this is often followed by subsequent coarsing upward, topped by fluvial gravels. The Parschlug flora was deposited in lacustric facies, in rather homogeneous pelitic sediments that developed above the coal seam. Besides water, wind has probably played an essential role in transporting the plant material into the lake sediments because the diversity of winged fruits and seeds is rather high: Acer sp. div., Ailanthus confucii, Betulaceae, Cedrelospermum, Craigia bronnii, Engelhardia macroptera, Fraxinus primigenia, Paliurus favonii, Pinus sp., Saportaspermum, Tilia longebracteata, and Cypselites. Flower remains such as Antholithes stiriacus and ? Chaneya sp. or insect wings still preserved in pairs support this view. In this respect the flora of Parschlug resembles the Cypris Clay flora (BŮZEK et al. 1996) and the flora from the Randeck Maar (RÜFFLE 1963). The differentiation of vegetation due to properties of the substrate (oligotrophic soils with sclerophyllous oaks and pine forests versus gallery forests on fertile soils) has largely vanished in this way from the fossil assemblages. Palaeoecology, sociology, and climate In the assemblage of Parschlug (fig. 4), a few azonal elements are most common: Glyptostrobus europaeus, Myrica lignitum, and Liquidambar europaea. Others are KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 93 either not abundant such as Acer tricuspidatum and Populus populina or even rather rare such as Alnus julianiformis and Cercidiphyllum. Most taxa are clearly zonal. Among these, more or less humid mesophytic elements are of minor importance: Fagus, Betula, Zelkova, Acer, Platanus leucophylla, Fraxinus, Tilia, and Berchemia. Certain genera that today include species with chartaceous and rather large leaves (leaf size classes notophyllous/microphyllous) as well as species with coriaceous small leaves such as Ulmus and Acer are represented by the small-leafed species U. plurinervia and A. pseudomonspessulanum, respectively. Also, Smilax is represented by small-leafed forms of S. sagittifera and the Theaceae by the small-leafed Ternstroemites pereger. They suggest subhumid conditions, which is supported by several taxa with characteristic subhumid physiognomic appearance such as Cedrelospermum, Berberis, Mahonia (?), Leguminosae, Paliurus, Cotinus (?), and even by sclerophyllous plants such as Quercus mediterranea, Q. zoroastri, and Q. drymeja. Additionally, there are some taxa, partly of obscure systematic affinity, which recall subhumid conditions due to their coriaceous lamina (e.g., "Quercus" daphnes). Neither diverse nor abundant are taxa typical of humid subtropical forests, including the Lauraceae (only 3 leaves of Daphnogene polymorpha), Theaceae (cf. ? Gordonia oberdorfensis), and Engelhardia. Ailanthus may also belong to this group. Some taxa are difficult to evaluate in this respect, among them Pinus div. sp. or certain Leguminosae. The reconstruction of the vegetation profile from the basin to the upland (fig. 5) is based on the autecology of single taxa, their abundance and the whole association. Aquatic plants are extremely rare, being confined to a few specimens of Salvinia and monocots. Therefore, we have no reason to assume the presence of extensive shallow water in the area where the plants were collected. (This probably represents the southwestern area of the Parschlug basin, where mining has focussed due to favourable geological conditions - see Geography and Geological Frame.) Oligotypic wetland gallery forests composed mainly of Glyptostrobus europaeus, Myrica lignitum, Liquidambar europaea and possibly also Zelkova zelkovifolia were developed along the shores of the lake. The mesophytic forests on drier substrates were species-diverse and probably not uniform, depending on the soil and exposition to the sun. We may expect small patches of humid mesophytic habitats with mixed-mesophytic forests (Fagus, Betula, Engelhardia, Fraxinus, Ailanthus, Daphnogene, and Podocarpium podocarpum). More extensive were probably subhumid forests composed of sclerophyllous oaks in the canopy and plants of similar physiognomic character in the shrub storey, including Berberis, Mahonia (?), possibly Prinsepia, Cedrelospermum, and Paliurus. Legumes may have been present in both. Pine stands were dispersed or mixed with oaks. Oak and pine forests probably grew preferentially on southern slopes and poor substrate, while humid mesophytic forests developed on northern expositions and deeper soils. The climatic proxies are deduced from the taxonomic composition and the physiognomic character of the elements. Most important in this respect is the floristic spectrum, which we assign to zonal vegetation. The relatively high number of physiognomically evergreen sclerophyllous and small-leafed taxa suggests subtropical but relatively drier climatic conditions compared to the preceeding humid, probably frostless subtropical/warm temperate conditions documented from the Lower Miocene of Oberdorf (Ottnangian; MELLER et al. 1999: 169) and the humid warm-temperate conditions documented from the Pannonian of the Molasse zone north of the Alps (KOVAR-EDER 1988: 63). 94 Annalen des Naturhistorischen Museums in Wien 105 A KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 95 Parschlug in the context of other floras from the Norian depression The relatively low diversity and mostly low abundance of azonal components and the high diversity of probably zonal taxa clearly distinguish the Parschlug flora from all the others known along the Norian depression (Tamsweg, Leoben, Fohnsdorf ETTINGSHAUSEN 1888, KNOBLOCH & KVACEK 1982, STRÖBITZER 1999). Of the azonal taxa, Glyptostrobus europaeus, Myrica lignitum, and Liquidambar europaea are common in all the mentioned localities. Acer tricuspidatum is present at Parschlug but not very abundant. Alnus julianiformis, usually one of the most common azonal elements (e.g. Leoben), is documented in Parschlug by merely one specimen. The probably zonal elements from Parschlug, e.g. Cedrelospermum, Leguminosae, Rosaceae, Berberidaceae, are largely absent at the other sites. Actually, physiognomically sclerophyllous taxa are both diverse and abundant there in contrast to the other floras of this region. The scarcity of Lauraceae at Parschlug also contrasts with the other sites, where Daphnogene is rather common and often associated with other Lauraceae (e.g. at Lintsching). Among the Fagaceae, physiognomically sclerophyllous oaks (Q. drymeja, Q. mediterranea, Q. zoroastri) prevail at Parschlug but are almost absent in the other sites. Quercus kubinyii (KOVATS ex ETTINGSHAUSEN) CZECZOTT alias Castanea atavia UNGER is doubtful at Parschlug, but otherwise very abundant, e.g. at Leoben. In all the floras along the Norian depression, Fagus is rather rare if present at all. Beech obviously played a less important role in the vegetation here than later during the Late Miocene (KOVAR-EDER 1988). Moreover, the Parschlug flora is unique in several aspects: Although its diversity is reduced from formerly about 180 to 60 taxa, it is by far the most species-diverse Neogene flora in Austria. Among the verified species, the higher number of probably endemic taxa is remarkable, e.g. Prinsepia serra, Ternstroemites pereger, and Mahonia (?) aspera. The presence and diversity of the Fabaceae (fruits and leaves) is remarkable, and none of the other floras from the Norian depression compares with that of Parschlug. Fig. 4: Floral picture of Parschlug: (1) Osmunda parschlugiana, (2-4) Pinus sp. div., (5) Glyptostrobus europaeus, (6) ? Cupressus sp., (7) ? Cathaya sp., (8) Liquidambar europaea, (9) Cercidiphyllum crenatum, (10) Daphnogene polymorpha, (11, 12) Berberis teutonica, (13) Betula vel Alnus sp., (14) Alnus gaudinii, (15) Fagus sp., (16) Quercus drymeja, (17) Quercus mediterranea UNGER, (18) Platanus leucophylla, (19) Quercus zoroastri UNGER, (20) cf. Gordonia oberdorfensis, (21) Ternstroemites pereger, (22) Engelhardia macroptera, (23) Engelhardia orsbergensis, (24) Tilia longebracteata, (25) Craigia bronnii, (26) Ulmus plurinervia, (27) Ulmus parschlugiana, (28) Myrica lignitum, (29) Myrica oehningensis, (30) Cedrelospermum ulmifolium, (31) Celtis japeti, (32) Zelkova zelkovifolia, (33) cf. Rosa sp., (34) Buxus cf. egeriana, (35) Populus sp., (36) Leguminosites palaeogaea, (37) Leguminosites hesperidum, (38) Leguminosites parschlugianus, (39, 40) Podocarpium podocarpum, (41) "Acacia" parschlugiana, (42) Phaseolites securidacus, (43) "Juglans" parschlugiana, (44) Toxicodendron herthae, (45) Acer integrilobum, (46) Acer pseudomonspessulanum, (47) Acer tricuspidatum, (48-50) Acer sp. div., (51) Paliurus tiliifolius, (52) Paliurus favonii, (53) Berchemia multinervis, (54) Cotinus (?) aizoon, (55) Ailanthus confucii, (56) Fraxinus primigenia, (57) Nerium sp., (58, 59) Smilax sagittifera, (60) "Celastrus" europaea, (61) "Evonymus" latoniae, (62) "Cornus" ferox, (63) ? Chaneya sp., (64) "Quercus" daphnes, (65) Mahonia (?) aspera, (66) Prinsepia serra, (67) Populus populina, (68) Ailanthus pythii, (69) Dicotylophyllum sp. 2, (70) Saportaspermum sp., (71) Antholithes stiriacus, (72) Cypselites, (73) Cedrelospermum stiriacum. 96 Annalen des Naturhistorischen Museums in Wien 105 A Fig. 5: Reconstructed vegetation of Parschlug. Gallery forest with Glyptostrobus europaeus, Myrica lignitum, Liquidambar europaea, and Zelkova zelkovifolia along a stream; small patches of humid mesophytic forests with Fagus, Betula, Engelhardia, Fraxinus, Ailanthus, Daphnogene, and Podocarpium on deeper soils and/or north-exposed slopes (foreground); on drier substrates and possibly south-exposed slopes subhumid (more open) forests with sclerophyllous oaks in the canopy and Ulmus plurinervia, Berberis, Mahonia (?), possibly Prinsepia, Cedrelospermum, and Paliurus; pine stands and ? Cupressus dispersed or mixed with oaks. The flora of Parschlug in the Central European context Southern Germany (Randeck Maar and Upper Freshwater Molasse) The flora from the Randeck Maar (RÜFFLE 1963), which has been dated to MN 5 (HEIZMANN 1983), compares in many aspects quite well with Parschlug. From the Upper Freshwater Molasse (OSM) several macrofloras are correlated by mammals and/or regional geology to the late Early Miocene/Middle Miocene (MN 5/6), (tabs. 1, 2). The publications dealing with Burtenbach (SCHMID 1983), Entrischenbrunn (SCHMITT & BUTZMANN 1997), Gallenbach (SCHMID & GREGOR 1983), and Kirrberg (RIEDERLE & GREGOR 1997) need many taxonomic corrections (see tab. 2). For other sites, only lists of unfigured taxa have been published (Ursberg, RIEDERLE 1997). Systematic treatments of the macrofloras from Pfaffenzell and Eberstetten do not exist. This situation hinders a more detailed comparison than given below. Nonetheless, a comparison is possible because these floras have been recently studied in the collection of the Bayerische Staatssammlung by the first author. The floras from the OSM are of relatively distal fluvial origin, while the Randeck Maar and Parschlug represent lacustric facies connected in the latter to lignite-forming facies. 97 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Tab. 1: Macrofloras from southern Germany relevant for the comparison with Parschlug and their stratigraphic position. Abbreviation used: BSM = Bayerische Staatssammlung Munich. locality Randeck Maar Burtenbach Eberstetten Entrischenbrunn Derching Gallenbach 1 Kirrberg Pfaffenzell 1 reference for macro flora RÜFFLE 1963 SCHMID 1983, and pers. studies of the first author in the BSM unpub., pers. studies of the first author in the BSM SCHMITT & BUTZMANN 1997 SCHMIDT 1976, 1980, and pers. studies of the first author in the BSM SCHMID & GREGOR 1983, and pers. studies of the first author in the BSM RIEDERLE & GREGOR 1997 unpub., pers. studies of the first author in the BSM age MN 5 OSM, sedimentary cycle 3, correlated MN 5 OSM, sedimentary cycle 6, correlated MN 5 by regional geology around Ries impact, correlated MN5/6 OSM, sedimentary cycle 8 reference for age HEIZMANN 1983 BÖHME et al. 2002 OSM, sedimentary cycle 8 BÖHME et al. 2002, and pers. comm. HEISSIG, 2002 pers. comm. BÖHME 2002 pers. comm, HEISSIG and BÖHME 2002 upper MN 6 by mammals around 14.6 m.a., radiometric date of a tuffite BÖHME et al. 2002 pers. comm. BÖHME, HEISSIG 2002 pers. comm. BÖHME 2002 In the OSM floras, the absence of conifers is remarkable. The Randeck Maar is poor in conifers as well. At Parschlug the abundance of Glyptostrobus europaeus is certainly linked to the nearby lignite-forming facies. Finally, conifers are neither diverse nor abundant there. The floras of the OSM, the Randeck Maar, and Parschlug share the presence and sometimes abundance of Podocarpium podocarpum (leaves and fruits). Moreover, different legume pods do occur in Eberstetten, Pfaffenzell, Kirrberg, and the Randeck Maar, as they do in Parschlug. The Randeck Maar and Parschlug share the characteristic pinnate leaves of "Acacia" parschlugiana ("unbestimmbarer leguminosenartiger Blattrest", in RÜFFLE 1963: pl. 9, figs. 23, 24). In the Randeck Maar, Parschlug, and Pfaffenzell some entire-margined, obviously coriaceous leaves possibly represent Leguminosae. This rather rich and diverse representation of Leguminosae distinguishes these floras from those of the late Middle/Late Miocene floras from the Styrian, Vienna, and Molasse basins. Populus is represented in all compared floras and sometimes even rather common, while at Parschlug, only P. populina has been recorded. Populus balsamoides is present in all south German localities, while P. populina is present only in few. In contrast, Salix records are relatively scarce and not abundant at the individual sites; the genus is completely absent at Parschlug. Platanus leucophylla is often associated in the OSM, as it is at Parschlug. Generally, maples are rare and not species-diverse compared to the record from the Late Miocene. Among the compared localities, Parschlug is the most species-diverse (3 maple species). The OSM floras and Parschlug share the presence of probable endemites: 98 Annalen des Naturhistorischen Museums in Wien 105 A - Prinsepia serra from Parschlug is unknown so far from any other site. - In the OSM, leaves that have been incorrectly determined as Pterocarya castaneifolia (GÖPPERT) MENZEL by RIEDERLE & GREGOR (1997) from Kirrberg and by SCHMID & GREGOR (1983) from Gallenbach and as Salix sp. by SCHMITT & BUTZMANN (1997, pl. 4, fig. 13) constitute a taxon of yet unknown affinity (? Rosaceae), but are present in most of the floras (except Burtenbach). - The identification of Zelkova ungeri (ETTINGSHAUSEN) KOVATS from Kirrberg (RIEDERLE & GREGOR 1997) and also from Oehningen (HANTKE 1954: pl. 8, figs. 1-2) is taxonomically (as well as nomenclaturally) incorrect. These leaves clearly do not represent the genus Zelkova. They are of yet unknown affinity but may rather represent leaflets of the Vitaceae (Parthenocissus). From Gallenbach, SCHMID & GREGOR (1983) also list Z. ungeri. However, the material studied in the Bayerische Staatssammlung Munich yielded no Zelkova, besides the above-mentioned peculiar leaf type, which seems to be almost endemic to the OSM. In fact, Zelkova zelkovifolia (correct synonym of Z. ungeri) is rarely encountered in the discussed OSM floras, but is abundant in the Randeck Maar and Parschlug. The flora from the Randeck Maar shares even more taxa with Parschlug. Noteworthy are Adiantum renatum (Adiantum sp., GREGOR 1986: pl. 1, fig. 4), Cupressus, Cedrelospermum (see tab. 2), Craigia bronnii (Pteleaecarpum europaeum (BRONN) BŮZEK & KNOBLOCH in GREGOR 1986), Engelhardia, Acer integrilobum, Antholithes stiriacus (Antholithus sp. sensu GREGOR 1986: pl. 4, fig. 12), Celtis, Berchemia, and Ailanthus confucii, of which the latter three are more abundant in the Randeck Maar. On the other hand, elements characteristic and abundant in Parschlug, such as Mahonia (?) aspera, Quercus mediterranea, and Ulmus plurinervia, have not been recorded in the Randeck Maar flora. However, the specimens figured as Zelkova praelonga BERGER (RÜFFLE 1963: pl. 4, fig. 3 and possibly fig. 4) may represent U. plurinervia. At the same time, some elements, e.g. Sideroxylon salicites (WEBER) WEYLAND and Koelreuteria macroptera (KOVATS) EDWARDS, are rather common in the Randeck Maar flora but absent at Parschlug. Cypris Shale flora, western Bohemia The Early Miocene (Ottnangian-Karpatian) flora of the Cypris Shale in western Bohemia belongs to the typical "Younger Mastixioid" plant assemblages, although the Mastixiaceae are infrequent (BŮZEK et al. 1996). The flora is dominated by thermophilic ("Palaeotropic") elements of a humid subtropical climate – Tetraclinis salicornioides, diverse Lauraceae, Theaceae, Symplocaceae, Trigonobalanopsis, Platanus neptuni, Engelhardia. Deciduous ("Arctotertiary") elements are also present, but of low diversity. A limited number of taxa are shared with the flora of Parschlug: Myrica lignitum, M. oehningensis, Acer integrilobum, Liquidambar, Podocarpium, Craigia, Tilia, Ulmus, Zelkova, Cedrelospermum, Fraxinus, Populus populina, Ailanthus, and Betulaceae. The striking difference between the two assemblages is in the humid (Cypris) versus subhumid (Parschlug) aspects. A number of endemic plants of Parschlug are not known in western Bohemia. Instead of P. leucophylla, Platanus netpuni occurs in the Cypris Shale flora. 99 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Pinus Cupressus Platanus leucophylla Quercus drymeja Quercus mediterranea Ulmus plurinervia Cedrelospermum Zelkova zelkovifolia type "Zelkova ungeri" sensu RIEDERLE & GREGOR 1997 - non Zelkova but ? Toxicodendron or Vitaceae Comptonia oeningensis/ Myrica vindobonensis Daphnogene Leguminocarpon div. (several seeds) Acacia parschlugiana Podocarpium podocarpum Populus balsamoides Populus populina Salix type "-Pterocarya castaneifolia" sensu RIEDERLE & GREGOR 1997- non Pterocarya but ? Rosaceae Smilax Parschlug Burtenbach Derching Eberstetten Entrischenbrunn Gallenbach 1 Kirrberg Pfaffenzell 1 Randecker Maar Glyptostrobus europaeus Tab. 2: Occurrences of selected taxa from the Parschlug plant assemblages compared with similar floras from southern Germany with notes and references: 11 RIEDERLE & GREGOR (1997), pl. 11, figs. 8-10, det. as Ulmus pyramidalis. 12 RIEDERLE & GREGOR (1997), pl. 3, fig. 9. 13 SCHMITT & BUTZMANN (1997), pl. 4, fig. 9, det. as Ulmus pyramidalis. 14 SCHMITT & BUTZMANN (1997), pl. 3, fig. 5 det. as cf. Populus mutabilis. 15 SCHMITT & BUTZMANN (1997) pl. 3, figs. 7, 8, pl. 4, fig. 2 det. as cf. Myrica sp. 16 SCHMITT & BUTZMANN (1997), det. as Salix sp. 17 SCHMID & GREGOR (1983), pl. 3, fig. 3. 18 None of the specimens figured by RIEDERLE & GREGOR (1997) belongs to Zelkova. 19 RÜFFLE (1963), pl. 9, figs. 23, 24 "unbestimmbarer leguminosenartiger Blattrest". 10 RÜFFLE (1963), pl. 5, figs. 16-26, pl. 20, figs. 4, 5 det. as Tremophyllum tenerrimum, pl. 12, figs. 1-17, pl. 25, fig. 6 det. as Embothrites borealis. 11 RÜFFLE (1963), pl. 1, fig. 14. Abbreviations used: v = very, a = abundant, r = rare, pr = poor remains, n = non. X va _ _ _ _ _ _ _ X pr X _ _ _ _ _ _ _ _ X _ X X X X _ X _ X X _ _ 3 X _ 1 X _ _ X _ X (r) X (r) _ _ 8 _ _ X _ _ X? _ 4 X X Xa X _ _ X _ X? _ _ X X _ X vr X Xa X X X Xa Xr X X _ _ X _ _ X X X _ X X _ X? X _ X _ _ _ X X 6 X X X X _ X _ _ _ _ _ X X _ X (r) _ _ _ _ _ _ _ 11 X? X _ _ _ 5 X _ _ _ _ X _ _ X? _ _ 2 n _ _ X _ _ _ _ _ _ X? 10 X X _ _ _ _ _ _ _ 9 X X X X X X X X X X _ X X X X X X X X X _ X _ _ 7 X? X _ _ 100 Annalen des Naturhistorischen Museums in Wien 105 A Tab 3: Revised floral list of Parschlug (prefixed symbols of approximate abundance: D – dominant, C – common, R – rare or single). R - Osmunda parschlugiana (UNGER) ANDREÁNSZKY R - Pronephrium stiriacum (UNGER) KNOBLOCH et Z. KVAČEK R - Adiantum renatum UNGER R - Salvinia cf. mildeana GOEPPERT C - Pinus sp. div. R - ? Cathaya sp. D - Glyptostrobus europaeus (BRONGNIART) UNGER R - ? Cupressus sp. R - Daphnogene polymorpha (A. BRAUN) ETTINGSHAUSEN R - Berberis teutonica (UNGER) KOVAR-EDER et Z. KVAČEK comb.nov. R - Berberis (?) ambigua (UNGER) KOVAR-EDER et Z. KVAČEK comb.nov. C - Mahonia (?) aspera (UNGER) KOVAR-EDER et Z. KVAČEK comb. nov. R - Cercidiphyllum crenatum (UNGER) R. BROWN D - Liquidambar europaea A. BRAUN R - Liquidambar sp. – fructus R - Platanus leucophylla (UNGER) KNOBLOCH R - Betula cf. dryadum BRONGNIART R - Betula vel Alnus sp. R - Alnus julianiformis (STERNB.) Z. KVAČEK et HOLÝ R - Alnus gaudinii (HEER) KNOBLOCH et Z. KVAČEK R - Fagus sp. - leaf R - Fagus sp. - cupule R - Fagus vel Alnus sp. C - Quercus drymeja UNGER C - Quercus mediterranea UNGER R - Quercus zoroastri UNGER R - cf. ? Gordonia oberdorfensis KOVAR-EDER R - Ternstroemites pereger (UNGER) KOVAR-EDER et Z. KVAČEK comb. nov. D - Myrica lignitum (UNGER) SAPORTA R – Myrica oehningensis (A.BRAUN) HEER R - Myrica sp. - fructus R - Engelhardia orsbergensis (WESSEL et WEBER) JÄHNICHEN, MAI et WALTHER R - Engelhardia macroptera (BRONGNIART) UNGER R - Tilia longebracteata ANDRAE R - Craigia bronnii (UNGER) Z. KVAČEK, BŮŽEK et MANCHESTER C - Ulmus plurinervia UNGER R - Ulmus parschlugiana KOVAR-EDER et Z. KVAČEK sp. nov. R - Cedrelospermum ulmifolium (UNGER) KOVAR-EDER et Z. KVAČEK comb. nov. -foliage R - Cedrelospermum stiriacum (ETTINGSHAUSEN) KOVAREDER et Z. KVAČEK comb. nov. - fructus C - Zelkova zelkovifolia (UNGER) BŮŽEK et KOTLABA R - Celtis japeti UNGER R - Populus populina (BRONGNIART) KNOBLOCH R - Populus sp. – fructus R - Buxus cf. egeriana Z. KVAČEK, BŮŽEK et HOLÝ R - cf. Rosa sp. C - Prinsepia serra (UNGER) KOVAR-EDER et Z. KVAČEK comb. nov. R - ? Prinsepia sp. R - Leguminosites hesperidum (UNGER) KOVAR-EDER et Z. KVAČEK comb. nov. R - Leguminosites dionysi (UNGER) KOVAR-EDER et Z. KVAČEK comb. nov. R - Leguminosites palaeogaea (UNGER) KOVAR-EDER et Z. KVAČEK comb. nov. R - Leguminosites parschlugianus (UNGER) KOVAR-EDER et Z. KVAČEK comb. nov. R - Podocarpium podocarpum (A. BRAUN) HERENDEEN R - Phaseolites securidacus UNGER R - "Acacia" parschlugiana UNGER R - "Juglans" parschlugiana UNGER R - Paliurus tiliifolius (UNGER) BŮŽEK R - Paliurus favonii UNGER R - Berchemia multinervis (A. BRAUN) HEER R - Acer tricuspidatum BRONN R - Acer pseudomonspessulanum UNGER emend. STRÖBITZER-HERMANN R - Acer integrilobum WEBER sensu WALTHER R - Acer sp. div. - fructus R - Toxicodendron herthae (UNGER) Z. KVAČEK et WALTHER R - Cotinus (?) aizoon (UNGER) KOVAR-EDER et Z. KVAČEK comb. nov. C - Ailanthus pythii (UNGER) KOVAR-EDER et Z. KVAČEK comb. nov. R - Ailanthus confucii UNGER R - Fraxinus primigenia UNGER R - Nerium sp. R - Smilax sagittifera HEER emend. HANTKE R - Monocotyledoneae gen. et sp. indet. R - "Celastrus" europaea UNGER R - "Cornus" ferox UNGER R - "Evonymus" latoniae UNGER C - "Quercus" daphnes UNGER R - Antholithes stiriacus KOVAR-EDER et Z. KVAČEK sp. nov. R - Cypselites sp. C - Saportaspermum sp. R - ? Chaneya sp. R - Dicotylophyllum sp. 1-6 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 101 Miocene of Hungary Among numerous Sarmatian floras of Hungary (ERDEI in press), those from the areas of the Tokaj Mountains (e.g. Erdőbénye and Tallya) are comparable with Parschlug due to their mesophytic character. Taken together, they share a higher number of common elements including Fagus, Quercus mediterranea, Q. drymeja, Betulaceae, Ulmaceae and Celtidaceae, Engelhardia, Acer, Platanus leucophylla, Ailanthus, and some Leguminosae including Podocarpium. Noteworthy is the same scarcity of the Lauraceae as in Parschlug. Contrary to Parschlug, younger elements typical of the Late Miocene and Pliocene are already present: Ginkgo, Quercus pseudocastanea/pseudorobur group, Acer jurenakyi/subcampestre. Also lacking at Parschlug, but probably for palaeoclimatic reasons, are several "throughrunners", e.g. Tetraclinis salicornioides, Parrotia, Liriodendron, Koelreuteria, and Pungiphyllum - elements that are widespread in the late Palaeogene and Neogene of Europe. Nowhere in the Hungarian Sarmatian do we encounter leaf fossils typical of Parschlug, e.g. Mahonia (?) aspera, Prinsepia serra, and greater accumulations of Myrica lignitum. Another group of similar floras, also with mesophytic features, is concentrated in southern Hungary in the Mecsek Mountains. The plant fossils there are bound to the facies of fish scale-rich shale and diatomite, such as at Magyaregregy (HABLY 1985). The dating is somewhat uncertain, but the sites are usually assigned to the latest Early and early Middle Miocene (HABLY 2001, 2002). These floras seem to share most characters, both in physiognomy and composition, although local differences also occur. Azonal wetland vegetation is composed of Glyptostrobus, and Myrica lignitum, while Liquidambar is lacking. Because the site is under study (HABLY in prep.), a more precise comparison is premature. We may only note a rich representation of Leguminosae, Ailanthus, and Cedrelospermum, and the occurrence of many elements in common with Parschlug, including Fagus, Acer integrilobum (as mecsekense), A. pseudomonspessulanum (as decipiens), Myrica lignitum, Antholithes stiriacus, Engelhardia, Zelkova, Celtis, Craigia, and Populus populina. Some other elements are differently represented at Magyaregregy – common Lauraceae, Ziziphus paradisiaca (HABLY 2002, personal communication). In summary, the mentioned late Early/Middle Miocene floras of Hungary are well comparable with Parschlug both in composition and the aspects described above. The lack of typical endemites of Parschlug, however, suggests an even more subhumid climate of the Parschlug assemblage. Miocene of Greece In the area of southern Europe, the flora of Kymi, Evia, Greece (UNGER 1867, KVACEK in VELITZELOS, ed. 2002) shares common aspects and some elements with Parschlug. Both floras are dominated by sclerophyllous oaks, myricas (including Myrica oehningensis), Leguminosae, and Glyptostrobus. Noteworthy common accessory elements are Cupressus, Tilia, Populus, Cedrelospermum, Saportaspermum, Mahonia (?) cyclophylla, and Berberis (?) ambigua. The Kymi flora is older, middle Early Miocene in age, and distinct by a number of different plants – "Encephalartos" goerceixianus, Calocedrus suleticensis, Tetraclinis, Berberis kymeana, Diospyros rugosa and others. 102 Annalen des Naturhistorischen Museums in Wien 105 A Tab. 4: List of taxa from Parschlug published in UNGER (1841-47) with current revisions. UNGER, 1841-7, Chloris protogaea – reference A taxon reference collection/no. revision Acer parschlugianum p. 132, pl. XLIII, fig. 5 H; LMJ 76517 Liquidambar europaea Acer productum p. 131, pl. XLII, fig. 8 LMJ 76526 Acer tricuspidatum Acer pseudocampestre p. 133, pl. XLIII, fig. 6 (folium) S ? Acer pseudomonspessulanum pl. XLIII, fig. 8, 9 (fructus) Acer sp. Acer pseudomonspessulanum new illustration pl. 10, fig. 10 p. 132, pl. XLIII, fig. 1 S; LMJ 76531 Acer integrilobum pl. 10, fig. 3 pl. XLIII, fig. 2 L; LMJ 76522 A. pseudomonspessulanum pl. 10, fig. 8 pl. XLIII, fig. 3 S; LMJ 76514 Saportaspermum sp. pl. XLIII, fig. 4 S; LMJ ? Saportaspermum sp. Acer trilobatum p. 130, pl. XLI, fig. 6 Acer tricuspidatum Adiantum renatum p. 122, pl. XXVII, fig. 1 Adiantum renatum pl. XXVII, fig. 2 indet. Betula dryadum p. 117 Betula cf. dryadum Ceanothus europaeus p. 144, pl. XLIX, fig. 8 Ceanothus subrotundus p. 144, pl. XLIX, fig. 7 Fagus deucalionis p. 101 Ilex ambigua p. 149, pl. L, fig. 14 H; LMJ 76519 Berberis (?) ambigua Ilex parschlugiana p. 148, pl. L, fig.8 H Dicotylophyllum sp. Ilex sphenophylla p. 148, pl. L, fig. 9 S; LMJ 76515 Mahonia (?) aspera Ilex stenophylla p. 149, pl. L, figs. 10-13 S Dicotylophyllum sp. Juniperites baccifera p. 80, pl. XXI, fig. 1 S; NHMW 2001B0017/1 Glyptostrobus europaeus pl. 1, fig. 14 pl. XXI, fig. 2 S Glyptostrobus europaeus LMJ 76523 Liquidambar europaea pl. XXXV, fig. 2 LMJ 76867 Liquidambar europaea pl. XXXV, fig. 3 (folium, infructescence) LMJ 76516 Liquidambar europaea and Liquidambar sp. Liquidambar europaeum p.120, pl. XXXV, fig. 1 ? Paliurus tiliifolius LMJ 76530 Betula vel Alnus sp. pl. XXXV, figs. 4, 52 Paliurus favonii p. 147, pl. L, fig. 6 (fructus) Daphnogene polymorpha pl. 2, fig. 1 Liquidambar europaea L; LMJ 76518 pl. L, figs. 7, 8 (folia) Paliurus favonii pl. 11, fig. 7 Paliurus tiliifolius Pteris parschlugiana p. 122, pl. XXXVI, fig. 6 H; LMJ 76520 Osmunda parschlugiana pl. 1, fig. 1 Quercus aspera p. 108, pl. XXX, fig. 1 right S; LMJ 76532 Mahonia (?) aspera pl. XXX, fig. 1 upper left, fig. 2 bottom S Mahonia (?) aspera pl. 13, fig. 4 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 103 Tab. 4: continued taxon reference collection/no. revision pl. XXX, fig. 1 bottom S Mahonia (?) aspera pl. XXX, fig. 2 top L; LMJ 76529 Mahonia (?) aspera new illustration pl. 13, fig. 3 pl. XXX, fig. 3 S Mahonia (?) aspera Quercus chlorophylla p. 111, pl. XXXI, fig. 1 H ? "Quercus" daphnes Quercus daphnes p. 112, pl. XXXI, fig. 2 S "Quercus" daphnes pl. XXXI, fig. 3 L; LMJ 76525 "Quercus" daphnes pl. 12, fig. 15 p. 113, pl. XXXII, fig. 1 L; LMJ 76524 Quercus drymeja pl. 4, fig. 1 pl. XXXII, fig. 2 S Quercus drymeja pl. XXXII, fig. 3 S Myrica lignitum pl. XXXII, fig. 4 S Quercus drymeja Quercus elaena p. 112, pl. XXXI, fig. 4 H "Quercus" daphnes Quercus hamadryadum p. 110, pl. XXX, fig. 8 H Dicotylophyllum sp. Quercus lignitum p. 13, pl. XXXI, figs. 5-7 S Myrica lignitum Quercus mediterranea p. 114, pl. XXXII, fig. 1 top left L; LMJ 76524 Quercus mediterranea pl. 4, fig. 8 p. 114, pl. XXXII, fig. 7 Quercus mediterranea Quercus drymeja Quercus mediterranea S pl. XXXII, figs. 5, 6, 8 pl. XXXII, fig. 9 Quercus serra Rhamnus aizoon ? Quercus mediterranea NHMW 1845/34/4 Quercus mediterranea pl. 4, fig. 9 p. 109, pl. XXX, fig. 4 left L; LMJ 76528 Prinsepia serra pl. XXX, fig. 5 S; LMJ 76521 Prinsepia serra pl. XXX, fig. 6 S pl. XXX, fig. 7 S Dicotylophyllum sp. p. 146, pl. L, fig. 1 (folium) S Cotinus (?) aizoon pl. L, fig. 2 folium S Cotinus (?) aizoon pl. L, fig. 3 flos Smilacites sagittata p. 129, pl. XL, fig. 4 Ulmus bronnii p. 100 Ulmus plurinervia Ulmus quercifolia Ulmus zelkovaefolia pl. 13, fig. 9 indet. H; LMJ 76512 Smilax sagittifera p. 95, pl. XXV, figs. 1-4 S Ulmus plurinervia p. 96, pl. XXV, fig. 5 H Dicotylophyllum sp. p. 94, pl. XXIV, fig. 7 upper right, figs. 9-12 S Zelkova zelkovifolia pl. XXIV, fig. 7 left bottom, fig. 8 (fructus) S Ulmus parschlugiana pl. XXIV, fig. 13 S Zelkova zelkovifolia pl. XXVI, fig. 7 L; NHMW 1987/57 Zelkova zelkovifolia pl. XXVI, fig. 8 L; LMJ 76513 Ulmus parschlugiana pl. 8, fig. 9 104 Annalen des Naturhistorischen Museums in Wien 105 A Tab. 5: List of taxa from Parschlug published in UNGER (1850a) with current revisions. References A-F see tabs. 4, 6, 8-11 UNGER, 1850 a. Genera et species taxon UNGER's revision Acacia parschlugiana in figure reference E pl. XI, fig. 19 (fructus) A A F F A E E B B Leguminosites hesperidum pl. XI, fig. 20 (folium) "Acacia" parschlugiana pl. XLII, fig. 8 Acer tricuspidatum pl. XV, fig. 1 (folium) Platanus leucophylla pl. XV fig. 2 (fructus) Acer sp. pl. XLIII, fig. 6 ? Acer pseudomonspessulanum pl. XV fig. 3 (fructus) Acer sp. pl. XV fig. 4 Acer cf. pseudomonspessulanum pl. XV fig. 5 (leaf fragment) Dicotylophyllum sp. pl. XLIII, fig. 1 Acer integrilobum pl. XLIII, fig. 2 Acer pseudomonspessulanum pl. XLIII, fig. 3 Acer sp. pl. XLI, fig. 6 Acer tricuspidatum pl. VIII, fig. 1 (folium) "Quercus" daphnes pl. VIII, fig. 2 (? fructus) pl. XXXVII, fig. 1 Adiantum renatum pl. IV, fig. 4 (? fructus) Dicotylophyllum sp. pl. IV, fig. 5 Mahonia (?) aspera pl. LV, figs. 11-14 (folia) Ternstroemites pereger pl. LV, fig. 15 (fructus) indet. F F E pl. XII, fig. 8 pl. XII, fig. 21, 22 pl. XI, fig. 3 (fructus) E E pl. III, fig. 41 (folium) pl. V, fig. 1 C C E pl. XLIII, fig.16 (folium) pl. XLIII, fig. 17 (fructus) pl. X, fig. 9 E B A pl. X, figs. 4,5 pl. LXV, fig. 7 pl. XLIX fig. 8 E A F F A Acer productum Acer pseudocampestre F F F A A Acer pseudomonspessulanum Acer trilobatum Achras lycobroma Adiantites renatus Amorpha stiriaca Adiantum renatum Amygdalus pereger Amygdalus quercula Andromeda glauca Azalea hyperborea Bauhinia parschlugiana Bauhinia parschlugiana Capparis ogygia Carpinus oblonga Cassia ambigua Cassia hyperborea Cassia memnonia Cassia petiolata Ceanothus europaeus Zizyphus renata Physolobium kennedyaefolium determination now Dicotylophyllum sp. Dicotylophyllum sp. Leguminosites parschlugianus ? Paliurus tiliifolius Dicotylophyllum sp. Ternstroemites pereger Engelhardia macroptera Dicotylophyllum sp. Dicotylophyllum sp. Dicotylophyllum sp. Dicotylophyllum sp. ? Paliurus tiliifolius 105 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Tab. 5: continued taxon Ceanothus subrotundus Celastrus cassinefolius Celastrus cassinefolius Celastrus cuneifolius Celastrus elaenus Celastrus elaenus Celastrus europaeus Celtis japeti Clethra teutonica Comptonia laciniata Comptonia oeningensis Comptonia ulmifolia UNGER's revision p.p. Celastrus cassinefolius p.p. Celastrus noaticus Pittosporum cuneifolium Fraxinus primigenia Glyzyrrhiza blandusiae Ilex stenophylla determination now E pl. II, figs. 2, 3 Dicotylophyllum sp. E pl. I, figs. 14, 15 Dicotylophyllum sp. pl. II, figs. 16-19 (folia) pl. III, figs. 34, a, b (flores) pl. II, figs. 10-13 pl. VIII, figs. 12, 13 Dicotylophyllum sp. Antholithes stiriacus "Celastrus" europaea Populus sp. pl. XLIII, figs. 25, 26 pl. XIX, figs. 24, 25 pl. XXIX, fig. 2 pl. XXXIX, fig. 8 pl. XXIX, fig. 3 pl. XXIX, figs. 4, 5 Celtis japeti Berberis teutonica Myrica lignitum "Acacia" parschlugiana Myrica lignitum Cedrelospermum ulmifolium "Cornus" ferox Dicotylophyllum sp. Ternstroemites pereger Monocotyledonae indet. Leguminosites dionysi E Prinus hyperboreus E E Macreightia F germanica C Crataegus teutonica F B C B B Cornus ferox Cotoneaster andromedae Crataegus orionis Cyperites tertiarius Cytisus dionysi Daphnogene cinnamomifolia Equisetites braunii Evonymus latoniae Fagus deucalionis Ilex ambigua Ilex cyclophylla Ilex parschlugiana Ilex phenophylla in figure reference A pl. XLIX, fig. 7 E pl. II, fig. 1 Ilex sphenophylla Ilex sphenophylla F F F C E pl. XXIV, fig. 21 pl. XVIII, figs. 11, 12 pl. XVIII, fig. 15 pl. XXVIII, fig. 5 pl. IV, fig. 1 E C C D D D E E A E A A E A E pl. II, fig. 25 pl. XLI, fig. 24 (folium) pl. XLI, fig. 25 (fructus) pl. VIII fig. 1 (fructus) pl. VIII, figs. 3, 8 pl. VIII, figs. 4-7 pl. IV, figs. 6, 7 (fructus) pl. IV, figs. 8-10 (folia) pl. L, fig. 8 pl. III, fig. 7, 8 pl. L, fig. 8 pl. L, fig. 9 pl. III, figs. 3-6 pl. L, figs. 10-13 pl. III, figs. 15-19 Daphnogene polymorpha Dicotylophyllum sp. "Evonymus" latoniae Betula vel Alnus sp. Fagus sp. Fraxinus primigenia Ailanthus pythii Dicotylophyllum sp. Dicotylophyllum sp. Berberis (?) ambigua Mahonia (?) aspera Dicotylophyllum sp. Mahonia (?) aspera Mahonia (?) aspera Dicotylophyllum sp. Dicotylophyllum sp. 106 Annalen des Naturhistorischen Museums in Wien 105 A Tab. 5: continued taxon Isoetites brauni Juglans acuminata UNGER's revision Juglans melaena Juglans quercina Ledum limnophilum Liquidambar acerifolium Liquidambar europaeum Muscites schimperi Myrica deperdita Myrtus miocenica Nemopanthes angustifolius Mimosa palaeogaea Nemopanthes angustifolius Olea mediterranea Paliurus favonii Phaseolites orbicularis Phaseolites physolobium Phaseolites securidacus Phaseolites serratus Pinites balsamodes Pinites centrotos Pinites furcatus Pinites goethanus Pinites hepios determination now B pl. LIII, fig. 3 (folium) Ailanthus pythii B B D pl. LIII, fig. 6 pl. LIII, figs. 7-9 pl. XIX, figs. 8-10 Quercus zoroastri Quercus drymeja Toxicodendron herthae F C A pl. XII, figs. 24-26 pl. XLIII, fig. 28 pl. XXXV, figs. 1-5 C E pl. XLIII, fig. 27 pl. XI, fig. 12 (fructus) C C F E pl. XXVII, fig. 1, 2 Dicotylophyllum sp. Liquidambar europaea Liquidambar europaea and L. sp. (fruit) Liquidambar europaea Leguminosites palaeogaeus indet. pl. XVIII, fig. 6 pl. III, fig. 35 Dicotylophyllum sp. Dicotylophyllum sp. E pl. L, fig. 6 pl. L, figs. 7, 8 pl. V, fig. 3 Paliurus favonii Paliurus tiliifolius Dicotylophyllum sp. E pl. V, fig. 4 Dicotylophyllum sp. E pl. V, figs. 9, 10 Phaseolites securidacus C C C C C C C C C C pl. XXXV, fig. 7 (scale) pl. XXXV, fig. 8 (folia) pl. XXXVII, figs. 1-3 pl. XXXVII, fig. 4 (male cone) pl. XXXVII, fig.7 (semen) pl. XXXVII, fig. 9 (folia) pl. XXXV, figs. 18-21 (semina) pl. XXXV, fig. 22 (folia) pl. XXXV, figs. 6-8 (folia) pl. XXXV, fig. 9 (semen) Pinus sp. Pinus sp. Pinus sp. Pinus sp. Pinus sp. Pinus sp. Pinus sp. Pinus sp. Pinus sp. Pinus sp. Juglans parschlugiana p.p. Juglans elaenoides Juglans falcifolia Juglans hydrophila Liquidambar protensum Mimosites palaeogaea in figure reference C pl. XXVII, fig. 18 A Physolobium orbiculare Physolobium antiquum indet. "Juglans" parschlugiana 107 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Tab. 5: continued taxon Pinites leuce Pinites oceanines Pistacia lentiscoides Populus aeoli Populus gigas Populus latior Populus ovalifolia Prinos europaeus Prinos hyperboreus UNGER's revision in reference C C C C D C C C figure determination now pl. XXXV, fig. 22 (semina) pl. XXXV, fig. 16 (folium) pl. XXXV, fig. 1 (semen) pl. XXXV, figs. 2-4 (folia) pl. XXI, fig. 14 pl. XLIV, fig. 2 pl. XIV, fig. 1 pl. XLIV, figs. 3-5 Pinus sp. Pinus sp. Pinaceae Coniferae Dicotylophyllum sp. Populus populina Platanus leucophylla Populus populina E pl. III, fig. 37 (folium) pl. III, figs. 34 (flores) pl. XVIII, figs. 26 (folium), 27 (fructus) pl. XVIII, fig. 30 Myrica lignitum Antholithes stiriacus Dicotylophyllum sp. and indet. ? Cedrelospermum ulmifolium ? Myrica infructescence Prunus atlantica F Prunus euri F Prunus paradisiaca F Prunus theodisca Pteris parschlugiana Pyrus euphemes Pyrus minor Pyrus theobroma Quercus aspera Quercus chlorophylla Quercus cyclophylla Quercus daphnes Quercus drymeja Quercus elaena Quercus hamadryadum Quercus lignitum Quercus mediterranea F F A B B A A A C A A A A A A C A A C C pl. XVIII, fig. 28 (? infructescence) pl. XVIII, fig. 29 (folium) pl. XVIII, fig. 31 pl. XXXVI, fig. 6 pl. LIX, fig. 10 pl. LIX, fig. 5 pl. XXX, figs.1 right, upper left, fig. 2 bottom pl. XXX, fig. 1 bottom, 2 top pl. XXXI, fig. 1 pl. XLI, fig. 15 pl. XXXI, figs. 2, 3 pl. XXXII, figs. 1, 2 pl. XXXII, fig. 3 pl. XXXII, fig. 4 pl. XXXI, fig. 4 pl. XXX, fig. 8 pl. XLI, figs. 1-7 pl. XXXII figs.1 top left, 7, 9 pl. XXXII, figs. 5, 6, 8 pl. XLI, figs. 1-3, 5, 6 pl. XLI, fig. 4 Dicotylophyllum sp. Quercus mediterranea Osmunda parschlugiana Dicotylophyllum sp. Dicotylophyllum sp. Mahonia (?) aspera Mahonia (?) aspera ? "Quercus" daphnes Quercus mediterranea ? "Quercus" daphnes Quercus drymeja Myrica lignitum Quercus drymeja "Quercus" daphnes Dicotylophyllum sp. Myrica lignitum Quercus mediterranea ? Quercus mediterranea Quercus mediterranea ? Quercus drymeja 108 Annalen des Naturhistorischen Museums in Wien 105 A Tab. 5: continued taxon Quercus myrtilloides Quercus serra Quercus urophylla Quercus zoroastri Rhamnus aizoides Rhamnus aizoon Rhamnus degener Rhamnus pygmaeus Rhododendron flos-saturni Rhus cuneolata Rhus elaeodendroides Rhus herthae Rhus napaearum Rhus nitida Rhus retine Rhus triphylla Rhus zanthoxyloides Robinia hesperidum Rosa penelopes Salix angustissima Sapindus pythii Sideroxylon hepios Smilacites sagittata Sparganium acheronticum Sphaerites disciformis Sphaerites punctiformis UNGER's revision in reference C C A A C C C C E A A E E E E F D D D D D D figure determination now pl. XLI, fig. 17 pl. XLI, figs. 18-20 pl. III, fig. 4 left, 5 pl. III, figs. 6, 7 pl. XLI, fig. 16 pl. XLI, fig. 11 pl. XLI, figs. 7, 8 pl. XLI, fig. 9 pl. III, fig. 47 pl. L, fig. 1, 2 (folia) pl. L, fig. 3 (flos) pl. III, fig. 44 pl. III, fig. 45, 46 pl. III, fig. 49 pl. III, fig. 48 pl. XII, fig. 12, fig. 15 pl. XX, fig. 12 pl. XXI, figs. 1-3, 6-8 pl. XXI, fig. 7 pl. XXI, figs. 4, 5, 11 pl. XX, figs. 7-9 pl. XX, fig. 11 ? Buxus cf. egeriana Dicotylophyllum sp. "Quercus" serra Dicotylophyllum sp. "Quercus" serra ? Quercus drymeja Quercus zoroastri Ailanthus pythii Dicotylophyllum sp. Cotinus (?) aizoon indet. Cotinus (?) aizoon Cotinus (?) aizoon Dicotylophyllum sp. Dicotylophyllum sp. "Quercus" daphnes Dicotylophyllum sp. Dicotylophyllum sp. Quercus drymeja Ailanthus pythii Toxicodendron herthae Dicotylophyllum sp. D D pl. XX, fig. 10 pl. XX, fig. 13 E E Dicotylophyllum sp. Cedrelospermum ulmifolium pl. XXI, fig. 13 Dicotylophyllum sp. pl. IV, figs. 11-13 (fructus) Leguminosites hesperidum pl. IV, fig. 14 (semina) Saportaspermum sp. pl. IV, figs. 15-17 (folia) Dicotylophyllum sp. D F A C pl. XIV, figs. 6-17 pl. VIII, fig. 4 pl. XL, fig. 4 pl. XXX, fig. 2 D E Ailanthus pythii Dicotylophyllum sp. Smilax sagittifera Monocotyledonae indet. 109 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Tab. 5: continued taxon Spiraea zephyri Styrax boreale Taxodites dubius Taxodites oeningensis Ulmus bronnii Ulmus parvifolia Ulmus plurinervia Ulmus praelonga Ulmus quercifolia Ulmus zelkovaefolia UNGER's revision in reference F F F figure determination now pl. XVIII, figs. 22, 23 pl. XI, fig. 11 pl. XI, figs. 12, 13 cf. Rosa sp. Mahonia (?) aspera ? Leguminosae C pl. XLIII, fig. 22 A C A C A pl. XXV, figs. 1-4 pl. XLIII, fig. 20 pl. XXV, fig. 5 pl. XLIII, fig. 24 pl. XXIV, fig. 7, upper right, figs. 9-12 pl. XXIV, fig. 7 left bottom, fig. 8 (fructus) pl. XXIV, fig. 13 pl. XXVI, fig. 7 pl. XXVI, fig. 8 pl. XII, fig. 1 a pl. XII, fig. 2 a, c pl. XII, fig. 5 a, b pl. XII, fig. 6 pl. XII, figs. 3 a-c Cedrelospemum ulmifolium vel Ulmus plurinervia Ulmus plurinervia Zelkova zelkovifolia Dicotylophyllum sp. Prinsepia serra Zelkova zelkovifolia A Vaccinium chamaedrys Vaccinium empetrites Vaccinium icmadophilum Vaccinium myrsinefolium Vaccinium vitis-japeti Widdringtonites ungeri Xylomites maculatus Xylomites tuberculatus Zanthoxylum fraxinoides Ziziphus protolotus Ziziphus tremula A A A F F F F F Zizyphus protolotus E Zizyphus tremula E pl. III, fig. 43 pl. III, fig. 39 Ulmus parschlugiana Zelkova zelkovifolia Zelkova zelkovifolia Ulmus parschlugiana Dicotylophyllum sp. Dicotylophyllum sp. Dicotylophyllum sp. Dicotylophyllum sp. Dicotylophyllum sp. ? Paliurus tiliifolius Paliurus tiliifolius 110 Annalen des Naturhistorischen Museums in Wien 105 A Tab. 6: List of taxa from Parschlug published in UNGER (1850b) with current revisions. UNGER, 1850 b. Socka (Sotzka) – reference B taxon reference collection/No. revision Amygdalus pereger p. 184, pl. LV, figs. 11-14 (folia) S Ternstroemites pereger pl. LV, fig. 15 (fructus) S indet. p. 164, pl. XXXII, fig. 6 LMJ 76540 Engelhardia macroptera pl. 6, fig. 9 pl. XXXII, fig. 4 LMJ 76545 possibly Engelhardia macroptera from Sotzka Cassia petiolata p. 189, pl. LXV, fig. 7 LMJ 76543 possibly Dicotylophyllum sp. from Sotzka Comptonia laciniata p. 161, pl. XXIX, fig. 2 H Myrica lignitum LMJ 76546 Myrica lignitum p. 162, pl. XXIX, fig. 4 S Cedrelospermum ulmifolium pl. XXIX, fig. 5 L; LMJ 76536 Cedrelospermum ulmifolium Juglans elaenoides p. 179, pl. LIII, fig. 3 (folium) LMJ 76542, counter- Ailanthus pythii impression 77652 pl. LIII, fig. 4 (fructus) S indet. Juglans hydrophila p. 179, pl. LIII, figs. 8, 9 S Quercus drymeja pl. LIII, fig. 6 S; LMJ 76866 Quercus zoroastri pl. 5, fig. 2 pl. LIII, fig. 7 S; LMJ 76549 Quercus drymeja pl. 4, fig. 2 Pyrus euphemes p. 183, pl. LIX, fig. 10 LMJ 76548 Dicotylophyllum sp. Pyrus minor p. 183, pl. LIX, fig. 16 LMJ 76547 Dicotylophyllum sp. Pyrus theobroma p. 183, pl. LIX, fig. 5 LMJ 76550 Dicotylophyllum sp. Carpinus producta Comptonia oeningensis p. 161, pl. XXIX, fig. 3 Comptonia ulmifolia new illustration pl. 7, fig. 7 pl. 8, fig. 5 Tab. 7: List of taxa from Parschlug published in ETTINGSHAUSEN (1851a) with current revisions. ETTINGSHAUSEN 1851 a. Tertiär-Floren der österreichischen Monarchie taxon reference Cassia ambigua p. 27, pl. V figs. 12, 13 GBA ?? ? Podocarpium podocarpum Liquidambar europaeum p. 15, pl. II, figs. 20, 22 GBA ?? Liquidambar europaea p. 14, pl. II, figs. 7, 13, 16 S, GBA ?? Zelkova zelkovifolia pl. II, figs. 15, 17 S, GBA ?? Cedrelospermum ulmifolium pl. II, fig. 18 S, GBA ?? ? Cedrelospermum ulmifolium pl. II, figs. 11, 12 S, GBA ?? Ulmus plurinervia Planera ungeri Pterospermum ferox collection/no. revision pl. II, fig. 14 S, GBA ?? ? Ulmus plurinervia p. 22, pl. IV, fig. 4 S, GBA "Cornus" ferox new illustration KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 111 Tab. 8: List of taxa from Parschlug published in UNGER (1852) and current revisions. UNGER, 1852. Iconographia plantarum fossilium – reference C taxon reference collection/no. revision Betula dryadum p. 105, pl. XXXIX, fig. 10 (fructus) LMJ 76497 Betula sp. pl. XXXIX, fig. 9 (folium) Betulaceae, ? Betula vel Alnus pl. XXXIX, fig. 11 indet. pl. XXXIX, fig. 12 indet. H new illustration Carpinus microptera p. 113, pl. XLIII, fig. 18 (bract) Carpinus oblonga p. 112, pl. XLIII, fig. 16 Ternstroemites pereger indet. p. 112, pl. XLIII, fig.17 (fructus) Engelhardia macroptera Celtis japeti p. 116, pl. XLIII, figs. 25, 26 Celtis japeti Comptonia laciniata p. 105, pl. XXXIX, fig. 8 "Acacia" parschlugiana Cyperites tertiarius p. 86, pl. XXVIII, fig. 5 LMJ 76511 Monocotyledoneae indet. Fagus deucalionis p. 110, pl. XLI, fig. 24 (folium) LMJ 76492 Betula vel Alnus sp. pl., XLI, fig. 25 (fructus) LMJ 62667 Fagus sp. pl. 3, fig. 3 Glyptostrobus oeningensis p. 92, unfig. Isoetites brauni p. 85, pl. XXVII, fig. 18 Liquidambar acerifolium p. 116, pl. XLIII, fig. 28 LMJ 76492 Liquidambar europaea indet. pl. 2, fig. 1 Liquidambar protensum p. 116, pl. XLIII, fig. 27 LMJ 76508 Liquidambar europaea pl. 2, fig. 4 Muscites fontinaloides p. 82, pl. XXVII, figs. 3, 4 Muscites schimperi p. 82, pl. XXVII, figs. 1, 2 LMJ 76498 indet. Myrica deperdita p. 104, sine fig. Pinites balsamodes p. 95, pl. XXXV, fig. 7 (scale) LMJ 76496 Pinus sp. pl. XXXV, fig. 8 Pinites centrotos p. 98, pl. XXXVII, fig. 1 Pinus sp. LMJ 76486 pl. XXXVII, figs. 2, 3 pl. XXXVII, fig. 4 (male cone) Pinites furcatus Pinites hepios Pinites goethanus Pinites leuce Pinites oceanines Pinus sp. Pinus sp. LMJ 76500 Pinus sp. p. 99, pl. XXXVII, figs. 7, 8 (semen) Pinus sp. pl. XXXVII, fig. 9 (folia) Pinus sp. p. 97, pl. XXXV, figs. 6-8 (folia) pl. 1, fig. 4 Pinus sp. pl. XXXV, fig. 9 (semen) LMJ 76501 Pinus sp. pl. 1, fig. 12 p. 96, pl. XXXV, fig. 18 (semen) LMJ 76491 Pinus sp. pl. 1, fig. 11 pl. XXXV, figs. 19-21 (semina) Pinus sp. pl. XXXV, fig. 22 (folia) Pinus sp. p. 95, pl. XXXV, figs. 9-15 (semina) Pinus sp. pl. XXXV, fig. 16 (folium) Pinus sp. p. 94, pl. XXXV, fig. 1 (semen) Pinaceae ? pl. XXXV, figs. 2-4 (folia) Coniferae 112 Annalen des Naturhistorischen Museums in Wien 105 A Tab. 8: continued taxon reference collection/no. revision Pinites taedaeformis Populus aeoli p. 97, pl. XXXVI, fig. 4 H Pinus sp. p. 117, pl. XLIV, fig. 2 H; LMJ 76506 Populus populina Populus gigas p. 117, pl. XLIV, fig. 1 Populus latior p. 117, pl. XLIV, fig. 3 LMJ 76509 Populus populina pl. XLIV, fig. 4 LMJ 76505 Populus populina pl. XLIV, fig. 5 LMJ 76484 Populus populina new illustration pl. 8, fig. 18 Platanus leucophylla Potamogeton castaliae p. 89, pl. XXX, fig. 1 LMJ 76499 indet. Quercus commutata p. 107, pl. XL, figs. 8, 9 S Myrica lignitum pl. XL, fig. 10 S; LMJ 76510 Myrica lignitum Quercus cyclophylla p. 109, pl. XLI, fig.15 Quercus mediterranea Quercus gmelini p. 108, pl. XLI, fig. 10 Dicotylophyllum sp. Quercus lignitum p. 106, pl. XL, fig. 1 LMJ 76504 Myrica lignitum p. 106, pl. XL, fig. 2 LMJ 76504 Myrica lignitum pl. XL, fig. 4 LMJ 76503 Myrica lignitum pl. XL, figs. 3, 5 pl. 14, fig. 1 pl. 7, fig. 8 pl. 7, fig. 2 pl. 7, fig. 6 Myrica lignitum pl. XL, fig. 6 LMJ 76510 Myrica lignitum pl. XL, fig. 7 LMJ 76485 Myrica lignitum LMJ 76507 Quercus mediterranea pl. 7, fig. 1 pl. XLI, figs. 21-23 (catkins) Quercus mediterranea p. 107, pl. XLI, fig. 1 pl. XL figs. 2, 3, 5, 6 Quercus mediterranea pl. XL, fig. 4 Quercus myricaefolia p. 109, pl. XLI, fig. 12 Quercus myrtilloides ? Quercus drymeja H Dicotylophyllum sp. p. 110, pl. XLI, fig. 17 LMJ 76502 ? Buxus cf. egeriana pl. XLI, fig.18 LMJ 76490 Dicotylophyllum sp. E; LMJ 76495 "Quercus" serra pl. XLI, figs. 19, 20 Dicotylophyllum sp. Quercus serra p. 110, pl. XLI, fig. 16 Quercus urophylla p.108, pl. XLI, fig.11 ? Quercus drymeja Quercus zoroastri p. 108, pl. XLI, figs. 7, 8 Quercus zoroastri pl. XLI, fig. 9 Ailanthus pythii Sparganium acheronticum p. 89, pl. XXX, fig. 2 Taxodites dubius pl. 8, fig. 15 pl. 13, fig. 10 ? LMJ 76499 (right) Monocotyledoneae p. 92, sine fig. Ulmus praelonga p. 115, pl. XLIII, fig. 20 H; LMJ 76487 Zelkova zelkovifolia pl. 8, fig. 11 Ulmus parvifolia p. 115, pl. XLIII fig. 22 LMJ 76488 Cedrelospermum ulmifolium vel Ulmus plurinervia pl. 8, fig. 12 Ulmus quercifolia p. 115, pl. XLIII fig. 24 Prinsepia serra Zelkova ungeri p. 114, pl. XLIII, fig. 19 Zelkova zelkovifolia 113 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Tab. 9: List of taxa from Parschlug published in UNGER (1860) with current revisions. UNGER, 1860. Sylloge plantarum fossilium 1 – reference D taxon reference Fraxinus primigenia p. 22, pl. VIII, fig. 1 (fructus) Fraxinus primigenia pl. VIII, figs. 3, 8 Ailanthus pythii pl. VIII, figs. 4-7 Dicotylophyllum sp. Juglans parschlugiana p. 37 pl. XIX, fig. 2 pl. XIX, fig. 4 collection/no. revision S; LMJ 76559 "Juglans" parschlugiana S; LMJ 76560 "Juglans" parschlugiana pl. XIX, figs. 1, 3, 5, 6 "Juglans" parschlugiana pl. XIX, fig. 7 (? fructus) indet. Juglans melaena p. 38, pl. XIX figs. 8-10 Toxicodendron herthae Pistacia lentiscoides p. 46, pl. XXI, fig. 14 Dicotylophyllum sp. Rhus cuneolata p. 44, pl. XX, fig. 12 LMJ 76553 Rhus elaeodendroides p. 45, pl. XXI, figs. 1-3, 6, 9 pl. 9, fig. 2 Dicotylophyllum sp. Dicotylophyllum sp. p. 45, pl. XXI, figs. 4, 5, 11 Ailanthus pythii pl. XXI, fig. 7 Rhus herthae new illustration Quercus drymeja pl. XXI, fig. 8 LMJ 76558 pl. XXI, fig. 10 LMJ 76556 p. 42, pl. XX, fig. 7 Dicotylophyllum sp. Dicotylophyllum sp. Toxicodendron herthae pl. XX, fig. 8 L; LMJ 76 562 A Toxicodendron herthae pl. XX, fig. 9 S; LMJ 76551 Toxicodendron herthae Rhus napearum p. 43, pl. XX, fig. 11 LMJ 76561 Dicotylophyllum sp. Rhus retine p. 43 pl. XX, fig. 10 Rhus triphylla p. 44, pl. XX, fig. 13 LMJ 76554 Cedrelospermum ulmifolium Rhus zanthoxyloides p. 45, pl. XXI, fig. 13 LMJ 76552 Dicotylophyllum sp. Sapindus pythii p. 33, pl. XIV, figs. 6, 7, 9-17 S Ailanthus pythii pl. XIV, fig. 8 Ailanthus pythii pl. 9, fig. 7 Dicotylophyllum sp. L; LMJ 76557 pl. 14, fig. 4 114 Annalen des Naturhistorischen Museums in Wien 105 A Tab. 10: List of taxa from Parschlug published in UNGER (1864) with current revisions. UNGER, 1864. Sylloge – reference E taxon reference collection/no. revision Acacia parschlugiana p. 34, pl. XI, fig. 19 (fructus) LMJ 77653 Leguminosites hesperidum pl. XI, fig. 20 (folium compositum) Amorpha styriaca p. 20, pl. IV, fig. 4 "Acacia" parschlugiana LMJ 76568 Dicotylophyllum sp. pl. IV, fig. 5 ? Mahonia (?) aspera Bauhinia parschlugiana p. 29, pl. XI, fig. 3 (fructus) Leguminosites parschlugianus Cassia ambigua p. 29, pl. X, fig. 9 Cassia memnonia p. 29, pl. X, figs. 4, 5 Dicotylophyllum sp. Celastrus cassinefolius p. 7, pl. II, fig. 1 Dicotylophyllum sp. Celastrus elaenoides p. 10, pl. II, figs. 16-19 Dicotylophyllum sp. Celastrus europaeus LMJ 76570 Dicotylophyllum sp. p. 10, pl. II, fig. 10 L; LMJ 76576 "Celastrus" europaea pl. II, fig. 11 S; LMJ 76581 "Celastrus" europaea pl. II, fig. 12 S; LMJ 76563 "Celastrus" europaea pl. II, fig. 13 Celastrus noaticus pl. II, fig. 2 pl. 12, fig. 2 Dicotylophyllum sp. pl. II fig. 3 LMJ 76539 Dicotylophyllum sp. Leguminosites dionysi Cytisus dionysi pl. IV, fig. 1 H; LMJ 76577 p. 11, pl. II, fig. 25 L; LMJ 76574 (part), "Evonymus" latoniae 76573 (counter part) Glyzyrrhiza blandusia p. 20, pl. IV figs. 6, 7 (fructus) pl. IV, figs. 8-10 (folia) pl. 9, fig. 5 pl. 12, fig. 3 Dicotylophyllum sp. Hardenbergia orbis-veteris p. 23, pl. V, fig. 5 "Cornus" ferox Ilex ambigua p. 14 Ilex cyclophylla p. 13, pl. III, fig. 7 LMJ 76537 Mahonia (?) aspera pl. III fig. 8 LMJ 76579 Mahonia (?) aspera p. 13, pl. III, fig. 9 pl. 13, fig. 3 Mahonia (?) aspera pl. III, fig. 10 LMJ 76580 Mahonia (?) aspera pl. III, fig. 11 LMJ 76572 Mahonia (?) aspera Ilex simularis p. 13, pl. III, fig. 14 Ilex sphenophylla p. 12, pl. III, fig. 3 LMJ 76571 Mahonia (?) aspera pl. III, fig. 4 LMJ 76538 Mahonia (?) aspera Ilex stenophylla pl. 12, fig. 1 Dicotylophyllum sp. Evonymus latoniae Ilex neogena new illustration Dicotylophyllum sp. pl. III, figs. 5, 6 Mahonia (?) aspera p. 14, pl. III, figs. 15-19 Dicotylophyllum sp. pl. 13, fig. 1 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 115 Tab. 10: continued taxon reference Mimosa palaeogaea p. 34, pl. XI, fig. 12 (fructus) Nemopanthes angustifolius p. 15, pl. III, fig. 35 Phaseolites securidacus p. 24, pl. V, fig. 9 collection/no. revision Leguminosites palaeogaeus LMJ 76567 Dicotylophyllum sp. L; LMJ 76569 Phaseolites securidacus pl. 9, fig. 14 pl. V, fig. 10 Dicotylophyllum sp. Physolobium antiquum p. 21, pl. V, fig. 4 Dicotylophyllum sp. Physolobium kennedyaefolium p. 22, pl. V, fig. 1 Dicotylophyllum sp. Physolobium orbiculare p. 22, pl. V, fig. 3 Dicotylophyllum sp. Pittosporum cuneifolium p. 6, pl. I, figs. 14, 15 Cotinus (?) aizoon Prinus hyperboreus p. 14, pl. III, fig. 34 (flores), Antholithes styriacus pl. III, fig. 37 (folium) Myrica lignitum Rhamnus aizoides p. 17, pl. III, fig. 47 LMJ 76565 Dicotylophyllum sp. Rhamnus aizoon p. 17, pl. III, fig. 44 L; LMJ 76575 Cotinus (?) aizoon pl. III, figs. 45, 46 Cotinus (?) aizoon Rhamnus degener p. 18, pl. III, fig. 49 Dicotylophyllum sp. Rhamnus pygmaeus p. 18, pl. III, fig. 48 Robinia hesperidum p. 21, pl. IV, fig. 12 (fructus) LM 76868 pl. IV, fig. 11, 13 (fructus) L; GBA 1864/01/21 Leguminosites hesperidum Dicotylophyllum sp. Leguminosites hesperidum pl. IV, fig. 14 Saportaspermum sp. pl. IV, figs. 15, 17 Dicotylophyllum sp. pl. IV, fig. 16 Zizyphus protolotus p. 17, pl. III, fig. 43 Zizyphus renata p. 16, pl. III, fig. 40, 41 Zizyphus tremula p. 16. pl. III, fig. 39 new illustration LMJ 76578 Dicotylophyllum sp. ? Paliurus tiliifolius Paliurus tiliifolius LMJ 76566 Paliurus tiliifolius pl. 9, fig. 4 116 Annalen des Naturhistorischen Museums in Wien 105 A Tab. 11: List of taxa from Parschlug published in UNGER (1866) with current revisions. UNGER, 1866. Sylloge – reference F taxon reference Acer productum p. 46, pl. XV fig. 1 Platanus leucophylla pl.XV, fig. 2 (fructus) Acer sp. p. 46, pl. XV, fig. 3 (fructus) Acer sp. pl. XV, fig. 4 (folium) Acer cf. psudomonspessulanum pl. XV, fig. 5 (folium) Dicotylophyllum sp. Acer pseudo-campestre collection/ no. revision Achras lycobroma p. 23, pl. VIII, fig. 1 (folium) LMJ 76591 "Quercus" daphnes Andromeda glauca p. 35, pl. XII, fig. 8 LMJ 76592 Dicotylophyllum sp. Azalea hyperborea p. 40, pl. XII, fig. 21 LMJ 76583 Dicotylophyllum sp. new illustration pl. 12, fig. 10 pl. VIII, fig. 2 pl. XII, fig. 22 Cornus ferox p. 76, pl. XXIV, fig. 21 Cotoneaster andromedae p. 59, pl. XVIII, fig. 11 pl. XVIII, fig. 12 Cotoneaster pusillus p. 59, pl. XVIII, fig. 13 Crataegus oreonis p. 59, pl. XVIII, fig. 15 Crataegus teutonica p. 60, pl. XIX, figs. 24, 25 Dicotylophyllum sp. "Cornus" ferox LMJ 76585 Dicotylophyllum sp. LMJ 76586 Dicotylophyllum sp. Dicotylophyllum sp. LMJ 76593 Ternstroemites pereger Berberis teutonica Ledum limnophilum p. 40, pl. XII, figs. 24-26 Dicotylophyllum sp. Macreightia germanica p. 26, pl. VIII, figs. 12, 13 (flores) Populus sp. Myrsine doryphora p. 19, pl. VI, fig. 10 ? "Quercus" daphnes Myrtus miocenica p. 57, pl. XVIII, fig. 6 Dicotylophyllum sp. Prunus atlantica p. 61, pl. XVIII, fig. 26 (folium) Dicotylophyllum sp. pl. XVIII, fig. 27 (fructus) Prunus euri p. 61, pl. XVIII, fig. 30 ? Cedrelospermum ulmifolium Prunus paradisiaca p. 62, pl. XVIII, fig. 28 ? Myrica (infructescence) pl. XVIII, fig. 29 (folium) Dicotylophyllum sp. Prunus theodisca p. 61, pl. XVIII, fig. 31 Quercus mediterranea Pyrus mini p. 58, pl. XVIII, fig. 20 Rhododendron flos-saturni p. 24, pl. 12, fig. 15 LMJ 76590 "Quercus" daphnes (counterpart) Sideroxylon hepios p. 24, pl. VIII, fig. 4 LMJ 76587 Dicotylophyllum sp. Spiraea zephyri p. 60, pl. XVIII, figs. 22, 23 cf. Rosa sp. Styrax boreale p. 33, pl. XI, fig. 11 ? Mahonia (?) aspera pl. XI, figs. 12, 13 ? Leguminosae cf. Rosa sp. Symplocos parschlugiana p. 33, pl. XI, fig. 10 Vaccinium chamaedrys p. 36, pl. XII, fig. 1 a Dicotylophyllum sp. pl. 12, fig. 11 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 117 Tab. 11: continued taxon reference collection/ no. Vaccinium empetrites p. 37, pl. XII, fig. 2 a LMJ 76588 Dicotylophyllum sp. pl. XII, fig. 2 c revision new illustration Dicotylophyllum sp. Vaccinium icmadophyllum p. 37, pl. XII, fig. 5 a, b Vaccinium myrsinefolia p. 38, pl. XII, fig. 6 Vaccinium vitis-japeti p. 36, pl. XII, figs. 3 a-c Dicotylophyllum sp. LMJ 76589 Dicotylophyllum sp. Dicotylophyllum sp. Age of the flora of Parschlug The age of the coal-bearing deposits has long been the subject of discussion. A late Karpatian age (late Early Miocene) was supposed based on mammal remains described by MOTTL (1970). However, these remains are not appropriate for such a precise dating (personal communication 2002 G. DAXNER-HÖCK). The newly characterized aspects of the assemblage from Parschlug offer some additional information: Engelhardia is more abundant in the Early Miocene and still occurs in the Badenian, e.g. at Weingraben, Wieliczka, but is a relict in the Early Pannonian (Rudabanya), and Early Pliocene (Gérce) in Central Europe. Cedrelospermum offers a similar picture - it survived only to the Sarmatian and we lack certain evidence from the Pannonian onwards. Unequivocal records of Acer vindobonensis, Acer aegopodifolium, and Acer subcampestre GÖPPERT are documented in Central Europe starting from the Sarmatian (rarely from the late Badenian). Also, roburoid oaks immigrated from the east and are documented largely starting from the Sarmatian. None of them occurs in the Parschlug flora. Of younger elements, only Platanus leucophylla is well documented at Parschlug. The first definite record is not older than the Badenian (Platanus leucophylla in Ukraine, Poland - KOVAR-EDER et al. 1994, 1996). Concluding from these considerations, we suspect a Karpatian/Early Badenian age (late Early – early Middle Miocene) for the flora of Parschlug. Acknowledgements Scientific discussions with M. BÖHME, K. HEISSIG, G. RÖSSNER (Munich), H. WALTHER (Dresden), L. HABLY (Budapest), R. SACHSENHOFER (Leoben), E. ZASTAWNIAK, J. WÓJCICKI (Kraków) and M. THIÉBAUT (Lyon) offered us important information. Collection studies were most essential at the Landesmuseum Joanneum and the Institut für Botanik, Karl-Franzens-Universität in Graz, at the Geologische Bundesanstalt and the Naturhistorisches Museum in Wien, and at the Institut für Geowissenschaften, Montanuniversität in Leoben. We thank A. DRESCHER, R. NIEDERL (Graz), and F. STOJASPAL (Wien) for providing us with help in these collections. Comparative investigations were carried out at the Bayerische Staatssammlung Munich and we thank H. MAYR for help there. M. STACHOWITSCH (Wien) kindly provided linguistic correction. A. SCHUMACHER (Wien) undertook a considerable part of the photo-documentation, A. PROCHÁSZKA (Budapest) drew the vegetation reconstruction, and J. SAKALA (Praha) and E. GREWAL (Wien) provided technical assistance. The critical reviews of E. ZASTAWNIAK and H. WALTHER contributed to the final improvements. 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KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis 125 Index Acacia .......................................................................... 73 Acacia parschlugiana ............. 58, 73, 75, 95, 97, 98, 99, 100, 104, 105, 111, 114, 144 Acer aegopodifolium .................................................. 117 Acer decipiens ........................................ 77, 78, 80, 101 Acer integrilobum .... 78, 95, 98, 100, 101, 102, 104, 146 Acer jurenakyi ............................................................ 101 Acer mescekense ...................................................... 101 Acer parschlugianum .......................................... 58, 102 Acer productum .................... 58, 77, 102, 104, 116, 146 Acer pseudocampestre ................. 77, 79, 102, 104, 116 Acer pseudomonspessulanum ... 77, 78, 79, 93, 95, 100, 101, 102, 104, 116, 146 Acer pyrenaicum ......................................................... 77 Acer sp. .... 79, 92, 93, 95, 100, 101, 102, 104, 116, 146 Acer subcampestre ........................................... 101, 117 Acer tricuspidatum .......... 77, 93, 95, 100, 102, 104, 146 Acer trilobatum ............................................ 77, 102, 104 Acer vindobonensis ................................................... 117 Achras lycobroma ............................... 85, 104, 116, 150 Adiantites renatus ............................................... 52, 104 Adiantum renatum ................ 52, 98, 100, 102, 104, 128 Ailanthus ................................... 82, 93, 96, 98, 101, 119 Ailanthus confucii ................ 81, 82, 92, 95, 98, 100, 148 Ailanthus mescekensis ................................................ 81 Ailanthus pythii ....................... 45, 81, 95, 100, 105, 106, 108, 110, 112, 113, 154 Alnus adscendens ....................................................... 59 Alnus formosana ......................................................... 60 Alnus gaudinii ........................................ 60, 95, 100, 132 Alnus julianiformis ........................... 59, 93, 95, 100, 132 Alnus trabeculosa ........................................................ 60 Amorpha stiriaca ......................................... 90, 104, 114 Amygdalus pereger ..................................... 63, 104, 110 Amygdalus quercula .................................................. 104 Andromeda glauca ...................................... 90, 104, 116 Antholites stiriacus ..................... 45, 86, 92, 95, 98, 100, 101, 105, 107, 115, 156 Antholitus sp. ......................................................... 86, 98 Apocynophyllum lanceolatum ...................................... 90 Apocynospermum ....................................................... 87 Aristolochia aesculapi .................................................. 84 Aristolochia parschlugiana .......................................... 84 Asterocalyx stiriacus .................................................... 86 Astrocalyx .................................................................... 86 Azalea hyperborea ...................................... 90, 104, 116 Bauhinia parschlugiana ......................... 74, 76, 104, 114 Berberis (?) ambigua .... 45, 56, 100, 101, 102, 105, 130 Berberis ................................................................. 93, 96 Berberis berberidifolia ................................................. 56 Berberis centiflora ....................................................... 56 Berberis kymeana ..................................................... 101 Berberis pruinosa ........................................................ 56 Berberis teutonica ........... 45, 56, 95, 100, 105, 116, 130 Berchemia multinervis ................................. 95, 100, 148 Betula cf. dryadum .............................. 59, 100, 102, 132 Betula dryadum ........................................... 59, 102, 111 Betula sp. .................................................................. 111 Betula vel Alnus sp. ...... 59, 95, 100, 102, 105, 111, 132 Blechnum dentatum .................................................... 52 Buxus cf. egeriana ................ 71, 95, 100, 108, 112, 142 Buxus egeriana ........................................................... 71 Buxus pliocenica ......................................................... 71 Calocedrus suleticensis ............................................. 101 Capparis ogygia .................................................. 90, 104 Carpinus microptera .................................................. 111 Carpinus oblonga ........................................ 63, 104, 111 Carpinus producta ....................................... 65, 110, 138 Cassia ambigua .................................... 74, 90, 104, 114 Cassia hyperborea .................................................... 104 Cassia memnonia ....................................... 90, 104, 114 Cassia petiolata ................................................. 104, 110 Castanea atavia .................................................... 60, 95 Catalpa europaea ........................................................ 83 ? Cathaya sp. ........................................ 54, 95, 100, 128 Ceanothus europaeus ................................. 76, 102, 104 Ceanothus subrotundus .............................. 55, 102, 105 Cedrela acuminata ...................................................... 75 Cedrelospermum .............................. 66, 67, 68, 69, 92, 99, 101, 117, 119 Cedrelospermum flichei ............................................... 68 Cedrelospermum hablyae ........................................... 69 Cedrelospermum stiriacum ................... 68, 95, 100, 142 Cedrelospermum ulmifolium .............. 95, 100, 105, 107, 108, 109, 110, 112, 113, 116, 142 Celastrus cassinefolius ............................... 90, 105, 114 Celastrus cuneifolius ........................................... 81, 105 Celastrus elaenoides ................................................. 114 Celastrus elaenus ................................. 86, 90, 105, 114 126 Annalen des Naturhistorischen Museums in Wien 105 A Celastrus europaea .............. 84, 95, 100, 105, 114, 150 Celastrus europaeus ... 66, 70, 71, 84, 85, 105, 114, 150 Celastrus noaticus ................................ 88, 90, 105, 114 Celastrus sp. ................................................... 56, 85, 86 Celtis ............................................................. 89, 98, 101 Celtis begonioides ....................................................... 70 Celtis japeti ........................... 70, 95, 100, 105, 111, 142 Celtis trachytica ........................................................... 70 Ceratopetalum parschlugianum .................................. 63 Cercidiphyllum crenatum ...................... 57, 95, 100, 130 Chaetoptelea ......................................................... 67, 68 ? Chaneya sp. .................. 87, 88, 92, 95, 100, 124, 150 Chrysophyllum parschlugianum .................................. 75 Cladrastis .................................................................... 74 Clethra teutonica ................................................. 56, 105 Comptonia laciniata ...................... 64, 75, 105, 110, 111 Comptonia oeningensis ................ 65, 99, 105, 110, 140 Comptonia peregrina ................................................... 65 Comptonia ulmifolia ............................ 68, 105, 110, 142 Comptonia vindobonensis ........................................... 65 Cornus ferox ......... 84, 95, 100, 105, 110, 114, 116, 150 Cotinus (?) aizoon .... 45, 80, 95, 100, 103, 108, 115, 148 Cotinus cogyggria ........................................................ 81 Cotoneaster andromedae ........................... 90, 105, 116 Cotoneaster pusillus ............................................ 90, 116 Craigia bronnii ................... 66, 92, 95, 98, 100, 121, 138 Crataegus oreonis ............................................... 63, 116 Crataegus orionis ................................................ 63, 105 Crataegus teutonica .................................... 56, 105, 116 ? Cupressus sp. .......... 55, 95, 96, 98, 99, 100, 101, 128 Cyperites tertiarius ...................................... 84, 105, 111 Cypselites sp. ........................... 86, 87, 92, 95, 100, 156 Cytisus dionysi .................................... 74, 105, 114, 144 Dalbergia ..................................................................... 74 Daphne sp. .................................................................. 63 Daphnogene polymorpha .... 55, 93, 95, 100, 102, 105, 130 Davidia ........................................................................ 89 Decodon .............................................................. 83, 121 Dicotylophyllum deichmuelleri ..................................... 84 Dicotylophyllum sp. ....... 102, 103, 104, 105, 106, 107, 108, 109, 110, 112, 113, 114, 115, 116, 117 Dicotylophyllum sp. 1 .................................. 88, 100, 156 Dicotylophyllum sp. 2 ............................ 88, 95, 100, 156 Dicotylophyllum sp. 3 .................................. 88, 100, 156 Dicotylophyllum sp. 4 .................................. 89, 100, 156 Dicotylophyllum sp. 5 .................................. 89, 100, 156 Dicotylophyllum sp. 6 .................................. 89, 100, 156 Diospyros rugosa ...................................................... 101 Diospyros sp. ............................................................... 63 Diversiphyllum ............................................................. 84 Dryandroides lignitum ......................................... 64, 140 Embothrites borealis .................................................... 99 Embothrium affine ....................................................... 87 Embothrium giganteum ............................................... 69 Embothrium megalopterum ......................................... 69 Embothrium parschlugianum ....................................... 87 Embothrium postsotzkianum ....................................... 87 Embothrium salicinum ................................................. 87 Embothrium sotzkianum .............................................. 87 Embothrium stiriacum ........................................... 68, 69 Embothrium subboreale .............................................. 87 Encephalartos goerceixianus .................................... 101 Engelhardia ............................. 93, 96, 98, 101, 117, 120 Engelhardia macroptera ................. 65, 92, 95, 100, 104, 110, 111, 138 Engelhardia orsbergensis ..................... 65, 95, 100, 138 Evonymus latoniae ......... 84, 85, 95, 100, 105, 114, 150 Fagus feroniae ............................................................ 60 Fagus herthae ............................................................. 80 Fagus kraeuselii .......................................................... 60 Fagus menzelii ............................................................ 60 Fagus pashanica ......................................................... 61 Fagus saxonica ........................................................... 60 Fagus silesiaca ..................................................... 60, 61 Fagus sp. ................. 60, 95, 96,100, 101, 105, 111, 132 Fagus vel Alnus sp. ..................................... 61, 100, 122 Ficus tenuinervis ......................................................... 75 Ficus troglodytarum ..................................................... 63 Fraxinus intermedius ................................................... 63 Fraxinus pachyptera .................................................... 82 Fraxinus praeexcelsior ................................................ 82 Fraxinus primigenia ..... 81, 82, 92, 95, 100, 105, 113, 148 Ginkgo ....................................................................... 101 Glycyrrhiza blandusiae ................................ 90, 105, 114 Glyptostrobus ............................................................ 101 Glyptostrobus europaeus ............. 54, 55, 92, 93, 95, 96, 97, 99, 100, 102, 128 Glyptostrobus oeningensis .................................. 55, 111 Gordonia axillaris ......................................................... 63 cf. ? Gordonia oberdorfensis ................ 63, 93, 100, 136 Hakea parschlugiana ............................................ 65, 87 Hardenbergia orbis veteris .................................. 84, 114 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Hemiptelea .................................................................. 67 Hysterium parschlugianum .......................................... 78 Ilex ambigua ................................ 56, 102, 105, 114, 130 Ilex cyclophylla .................................... 57, 105, 114, 152 Ilex neogena ........................................................ 57, 114 Ilex parschlugiana ................................. 90, 91, 102, 105 Ilex phenophylla ........................................................ 105 Ilex simularis ....................................................... 90, 114 Ilex sphenophylla ........................ 57, 102, 105, 114, 152 Ilex stenophylla ................................... 91, 102, 105, 114 Isoetites brauni .................................................. 106, 111 Juglans bilinica ............................................................ 82 Juglans elaenoides ..................................... 81, 106, 110 Juglans hydrophila ................ 61, 62, 106, 110, 134, 136 Juglans melaena ......................................... 80, 106, 113 Juglans parschlugiana .... 75, 80, 95, 100, 106, 113, 144 Juniperites baccifera ................................... 55, 102, 128 Koelreuteria macroptera .............................................. 98 Laurus palaeo-benzoin ................................................ 85 Ledum limnophilum ..................................... 90, 106, 116 Leguminocarpon ............................................. 73, 74, 99 Leguminocarpos .......................................................... 73 Leguminosites ............................................................. 73 Leguminosites dionysi .......... 45, 74, 100, 105, 114, 144 Leguminosites hesperidum ............ 45, 73, 95, 100, 104, 108, 114, 115, 144 Leguminosites mescekense ........................................ 74 Leguminosites palaeogaeus .................. 45, 74, 100, 95, 106, 115, 144 Leguminosites parschlugiana ...................................... 74 Leguminosites parschlugianus ... . 45, 74, 95, 100, 104, 114, 144 Liquidambar acerifolium .............................. 58, 106, 111 Liquidambar europaea ....... 58, 92, 93, 95, 96, 100, 102, 106, 110, 111, 130 Liquidambar europaeum ..................... 58, 102, 106, 110 Liquidambar magniloculata ......................................... 58 Liquidambar orientalis ................................................. 58 Liquidambar protensum .............................. 58, 106, 111 Liquidambar sp. .......................... 98, 100, 101, 102, 130 Liquidambar styraciflua ............................................... 58 Liquidambar wutzleri ................................................... 58 Liriodendron .............................................................. 101 Macreightia germanica .......................... 70, 71, 105, 116 Madhuca ...................................................................... 86 Mahonia (?) aspera .... 45, 57, 95, 98, 100, 101, 102, 103, 127 104, 105, 107, 109, 114, 116, 152 Mahonia (?) cyclophylla ............................................ 101 Mahonia nervosa ......................................................... 57 Mahonia sp. ......................................................... 57, 119 Mimosa palaeogaea .................................... 74, 106, 115 Mimosites palaeogaea ........................................ 74, 106 Monocotyledoneae gen.et sp. indet. ........ 100, 105, 108, 111, 112 Monotes ....................................................................... 87 Muscites fontinaloides ............................................... 111 Muscites schimperi ............................................ 106, 111 Myrica deperdita .......................................... 90, 106, 111 Myrica lignitum ....... 61, 64, 65, 89, 90, 91, 92, 100, 10 103, 105, 107, 110, 112, 115, 119 Myrica oehningensis ............... 65, 95, 98, 100, 101, 140 Myrica sp. ...................................................... 65, 99, 100 Myrica ungeri ............................................................... 65 Myrica vindobonensis .................................................. 99 Myrsine doryphora ........................................ 85, 90, 116 Myrsine formosana ...................................................... 71 Myrtus miocenica ........................................ 90, 106, 116 Nemopanthes angustifolius ......................... 90, 106, 115 Nerium bilinicum .......................................................... 83 Nerium oleander .......................................................... 83 Nerium sp. ....................................... 82, 83, 95, 100, 148 Olea mediterranea ..................................................... 106 Osmunda parschlugiana ....... 52, 95, 100, 102, 107, 128 Osmunda regalis ......................................................... 52 Palaeocarya orsbergensis ........................................... 65 Paliurus favonii ............... 76, 92, 95, 100, 102, 106, 148 Paliurus tiliaefolius ....................................................... 76 Paliurus tiliifolius .......................... 76, 95, 100, 102, 104, 106, 109, 115, 148 Parthenocissus ...................................................... 69, 98 Phaseolites orbicularis ........................................ 90, 106 Phaseolites physolobium .................................... 90, 106 Phaseolites securidacus ....... 75, 95, 100, 106, 115, 144 Phaseolites serratus ............................................ 90, 106 Phyllerium lignitum ...................................................... 64 Physiolobum kennedyaefolium ................... 90, 104, 115 Physolobium antiquum ................................ 90, 106, 115 Physolobium orbiculare ............................... 90, 106, 115 Picrasma ..................................................................... 88 Pinites balsamodes ..................................... 53, 106, 111 Pinites centrotos ................................. 53, 106, 111, 128 Pinites furcatus ............................................ 53, 106, 111 128 Annalen des Naturhistorischen Museums in Wien 105 A Pinites goethanus ............................... 53, 106, 111, 128 Pinites hepios ...................................... 53, 106, 111, 128 Pinites leuce .......................................... 53, 54, 107, 111 Pinites oceanines ........................................ 54, 107, 111 Pinites taedaeformis ............................................ 54, 112 Pinus ciliata ................................................................. 54 Pinus hepios ................................................................ 54 Pinus laricio ................................................................. 54 Pinus palaeo-strobus ................................................... 54 Pinus post-taedaeformis .............................................. 54 Pinus prae-cembra ...................................................... 54 Pinus prae-pumilio ....................................................... 54 Pinus prae-silvestris .................................................... 54 Pinus prae-taedaeformis ............................................. 54 Pinus rigios .................................................................. 54 Pinus sp. .... 53, 92, 93, 99, 100, 106, 107, 111, 112, 128 Pistacia lentiscoides .................................... 90, 107, 113 Pistacia lentiscus ......................................................... 81 Pittosporum cuneifolium .............................. 81, 105, 115 Plagiospermum ........................................................... 72 Planera ungeri ......................................... 67, 68, 69, 110 Platanus hispanica ...................................................... 59 Platanus leucophylla .. 58, 89, 93, 95, 97, 98, 99, 100, 101, 104, 107, 112, 116, 117, 132, 134 Platanus neptuni .......................................................... 98 Platanus orientalis ....................................................... 59 Podocarpium ........................................... 73, 96, 98, 101 Podocarpium podocarpum ................. 74, 93, 95, 97, 99, 100, 110, 120, 144 Polyspora axillaris ....................................................... 63 Populus aeoli ...................................... 70, 107, 112, 142 Populus balsamoides ............................................ 97, 99 Populus gigas .............................................. 58, 107, 112 Populus latior ................................ 70, 71, 107, 112, 154 Populus mutabilis ........................................................ 99 Populus populina ........... 70, 71, 93, 95, 97, 98, 99, 100, 101, 107, 112, 142, 154 Populus sp. ............. 70, 95, 96, 100, 101, 105, 116, 142 Populus zaddachii ....................................................... 71 Porana ................................................................... 87, 88 Potamogeton castaliae .............................................. 112 Prinos hyperboreus ............................................... 64, 86 Prinsepia serra ..................... 45, 56, 62, 72, 95, 98, 100, 101, 103, 109,112,152 Prinsepia sinensis ....................................................... 72 ? Prinsepia sp. ............................................ 73, 100, 142 Pronephrium stiriacum ................................ 52, 100, 128 Prunus atlantica .......................................... 91, 107, 116 Prunus euri .................................................. 68, 107, 116 Prunus paradisiaca ............................... 66, 91, 107, 116 Prunus theodisca ........................................ 62, 107, 116 Pteleaecarpum europaeum ......................................... 98 Pteris parschlugiana ........................... 52, 102, 107, 128 Pterocarya castaneifolia ........................................ 98, 99 Pterospermum ferox ............................................ 84, 110 Pungiphyllum ............................................................. 101 Pyrus euphemes ......................................... 91, 107, 110 Pyrus mini ..................................................... 71, 72, 116 Pyrus minor ................................................. 91, 107, 110 Pyrus theobroma ......................................... 91, 107, 110 Quercus aspera ............................ 57, 76, 102, 107, 152 Quercus chlorophylla .................................. 85, 103, 107 Quercus coccifera ....................................................... 62 Quercus commutata .................................... 64, 112, 140 Quercus cyclophylla .................................... 62, 107, 112 Quercus daphnes .............. 72, 83, 85, 93, 95, 100, 103, 104, 107, 108, 116, 150 Quercus drymeja ..... 61, 62, 72, 93, 95, 99, 100, 101, 103 106, 107, 108, 110, 112, 113, 134 Quercus elaena ........................................... 85, 103, 107 Quercus gmelini .................................................. 91, 112 Quercus hamadryadum ............................... 91, 103, 107 Quercus ilex ................................................................ 61 Quercus kubinyii .................................................... 62, 95 Quercus lignitum ......................... 64, 103, 107, 112, 140 Quercus lonchitis ......................................................... 68 Quercus mediterranea ..... 56, 60, 61, 62, 93, 95, 98, 99, 100, 101, 103, 107, 112, 116, 134 Quercus myricaefolia .......................................... 91, 112 Quercus myrsinaefolia ................................................. 56 Quercus myrtilloides ..................... 71, 91, 108, 112, 142 Quercus pseudocastanea ......................................... 101 Quercus pseudorobur ................................................ 101 Quercus serra ................. 63, 72, 88, 103, 108, 112, 152 Quercus sosnowskyi ............................................. 62, 63 Quercus urophylla ....................................... 61, 108, 112 Quercus zoroastri ................ 61, 62, 63, 81, 93, 95, 100, 106, 108, 110, 112, 136 Rhamnus aizoides ....................................... 91, 108, 115 Rhamnus aizoon ................... 80, 81, 103, 108, 115, 148 Rhamnus degener ....................................... 91, 108, 115 Rhamnus pygmaeus ................................... 91, 108, 115 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Rhododendron flos-saturni .................. 85, 108, 116, 150 Rhus cotinus ................................................................ 81 Rhus cuneolata ........................................... 91, 108, 113 Rhus elaeodendroides .................... 81, 85, 91, 108, 113 Rhus herthae ...................................... 80, 108, 113, 144 Rhus napearum ........................................... 91, 108, 113 Rhus palaeocotinus ..................................................... 81 Rhus retine .................................................. 91, 108, 113 Rhus triphylla .............................................. 68, 108, 113 Rhus zanthoxyloides ................................... 91, 108, 113 Rhytisma aceris ..................................................... 77, 78 Robinia ........................................................................ 73 Robinia hesperidum ................ 73, 87, 91, 108, 115, 144 cf. Rosa sp. ........................... 71, 95, 100, 109, 116, 142 Salix sp. ................................................................. 98, 99 Salvinia ................................................................ 93, 118 Salvinia cerebrata ........................................................ 53 Salvinia cf. mildeana ................................... 53, 100, 128 Salvinia intermedia ...................................................... 53 Salvinia microphylla ..................................................... 53 Salvinia reussii ............................................................ 53 Sapindus pythii .............................. 62, 81, 108, 113, 154 Saportaspermum ................................................. 92, 101 Saportaspermum occidentale ...................................... 87 Saportaspermum sp. .... 87, 95, 100, 102, 108, 115, 156 Sapotacites longepetiolatus ....................................... 85 Sideroxylon hepios ...................................... 92, 108, 116 Sideroxylon salicites .................................................... 98 Smilacites sagittata ..................................... 83, 103, 108 Smilax ....................................................... 83, 86, 93, 99 Smilax grandifolia .................................................. 83, 86 Smilax sagittata ........................................................... 83 Smilax sagittifera ............ 83, 93, 95, 100, 103, 108, 148 Sorbus ......................................................................... 88 Spargianum acheronticum .......................... 84, 108, 112 Sphaeria mediterranea ................................................ 62 Sphaeria palaeo-sapindi ..................................... 81, 154 Spiraea zephyri ..................................... 71, 72, 109, 116 Styrax boreale ............................................. 57, 109, 116 Symplocos parschlugianus ........................................ 116 Taxodites dubius ........................................ 55, 109, 112 Taxodites oeningensis ........................................ 55, 109 Taxodium distichum miocenicum ................................ 55 Ternstroemites pereger ..... 45, 63, 72, 93, 95, 100, 104, 105, 110, 111, 116, 138 Tetraclinis salicornioides ..................................... 98, 101 129 Tilia ........................................................ 93, 98, 101, 122 Tilia lignitum ........................................................ 66, 138 Tilia longebracteata ......................... 66, 92, 95, 100, 138 Tilia milleri ................................................................... 66 Toxicodendron herthae ..... 80, 95, 100, 106, 108, 113, 144 Tremophyllum tenerrimum .................................... 68, 99 Trigonobalanopsis ....................................................... 98 Ulmus .............................................................. 68, 93, 98 Ulmus americana ........................................................ 68 Ulmus braunii .............................................................. 67 Ulmus bronnii ...................................................... 66, 103 Ulmus carpinoides ....................................................... 67 Ulmus parschlugiana 45, 67, 68, 95, 100, 103, 109, 138 Ulmus parvifolia ................................................................ ............................................................. 68, 109, 112, 142 Ulmus plurinervia ....... 66, 67, 68, 72, 93, 95, 96, 98, 9 100, 103, 109, 110, 112, 138, 142 Ulmus praelonga ................................. 69, 109, 112, 142 Ulmus pyramidalis ................................................. 67, 99 Ulmus quercifolia .......................... 72, 92, 103, 109, 112 Ulmus zelkovaefolia ...................... 67, 69, 103, 109, 142 Vaccinium chamaedrys ............................... 92, 109, 116 Vaccinium empetrites .................................. 92, 109, 117 Vaccinium icmadophilum ............................ 92, 109, 117 Vaccinium myrsinefolium ............................ 92, 109, 117 Vaccinium vitis-japeti ................................... 92, 109, 117 Widdringtonia baccifera ............................................... 55 Widdringtonia ungeri ................................................... 55 Widdringtonites ungeri ........................................ 55, 109 Xylomites aceris decipientis ........................................ 78 Xylomites aristolochiae ................................................ 84 Xylomites daphnes ...................................................... 85 Xylomites drymejae ............................................. 61, 134 Xylomites liquidambaris ............................................... 58 Xylomites quercus serrae ............................................ 72 Xylomites rhamni aizoonis .......................................... 81 Zelkova ................................. 66, 67, 69, 93, 98, 99, 101 Zelkova praelonga ................................................. 69, 98 Zelkova ungeri ........................................ 69, 98, 99, 112 Zelkova zelkovifolia ....... 68, 69, 93, 95, 96, 98, 99, 10 103, 109, 110, 112, 142 Ziziphus paradisiaca .................................................. 101 Ziziphus protolotus ...................................... 76, 109, 115 Ziziphus renata ............................................ 76, 104, 115 Ziziphus tremula .......................................... 76, 109, 115 130 Annalen des Naturhistorischen Museums in Wien 105 A Plate 1 Osmunda parschlugiana (UNGER) ANDREÁNSZKY 1 - LMJ 76520, holotype of Pteris parschlugiana UNGER (1847: pl. 36, fig. 6), 2 x 2 - NHMW 1878/6/6795, a - 1 x, b - 3 x Pronephrium stiriacum (UNGER) KNOBLOCH & Z. KVACCEK 3 - IBUG Ett. coll. 111, 2 x Salvinia cf. mildeana GOEPPERT 4 - IBUG Ett. coll. 113, 2 x Adiantum renatum UNGER 5 - IBUG Ett. coll. 344, 5 x Pinus sp. div. 6 - GBA 2002/01/26, 3-short-needled, 1x 7 - NHMW 1878/6/9706, 2-needled, 1 x 8 - NHMW 1878/6/9780, 3-long-needled, 1 x 9 - NHMW 1878/6/2479, male catkin, 4 x 10 - LMJ 76500, male catkin, syntype of Pinites centrotos UNGER (1852: pl. 37, fig. 4), 3 x 11 - LMJ 76491, seed, syntype of Pinites goethanus UNGER (1852: pl. 35, fig. 18), 2 x 12 - LMJ 76501, seed, syntype of Pinites hepios UNGER (1852: pl. 35, fig. 9), 2 x 13 - IBUG Ett. coll.195, cone scale, 1 x Glyptostrobus europaeus (BRONGNIART) UNGER 14 - NHMW 2001B0017/0001, branched twig, syntype of Juniperites baccifera UNGER (1845: pl. 21, fig.1), 1 x 15 - IBUG Ett. coll.190 a, twig taxodioid, 1 x 16 - IBUG Ett. coll.162, seed cone, 2 x ? Cupressus sp., twigs, all 2 x 17 - GBA 2002/01/24 18 - GBA 2002/01/23 19 - NHMW 1845/0039/0003 ? Cathaya sp. 20 - needle, IBUG Ett. coll. 343, 1 x 21 - 23 cone scales, all 2 x, 21 - IBUG Ett. coll. 318 22 - IBUG Ett. coll. 335 23 - IBUG Ett. coll. 317 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 1 132 Annalen des Naturhistorischen Museums in Wien 105 A Plate 2 Liquidambar europaea A. BRAUN 1 - LMJ 76492, holotype of Liquidambar acerifolia UNGER (1852: pl. 43, fig. 28), 1 x 2 - NHMW 1878/6/2406, a three-lobed leaf, 1 x 3 - NHMW 1878/6/9542, 1 x 4 - LMJ 76492, holotype of Liquidambar protensa UNGER (1852: pl. 43, fig. 27), 1 x 5 - NHMW 1878/6/9052, a five-lobed leaf, 1 x Liquidambar sp. - fructus 6 - NHMW 1878/6/9538, 1 x Cercidiphyllum crenatum (UNGER) R. BROWN 7 - NHMW 1878/6/6510, 1 x. Daphnogene polymorpha (A. BRAUN) ETTINGSHAUSEN 8 - NHMW 2001B0017/0002, 1 x Berberis teutonica (UNGER) KOVAR-EDER & Z. KVACCEK comb.nov. 9 - NHMW 1878/6/2153, neotype, a - 1 x, b - 2 x 10 - NHMW 1878/6/2442, 1 x Berberis (?) ambigua (UNGER) KOVAR-EDER & Z. KVACCEK comb.nov. 11 - LMJ 76519, holotype of Ilex ambigua UNGER (1847: pl. 50, fig. 14), 1.5 x KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 2 134 Annalen des Naturhistorischen Museums in Wien 105 A Plate 3 Betula cf. dryadum BRONGNIART 1 - IBUG Ett. coll. 725, fruitlet, 5 x Fagus vel Alnus sp. 2 - NHMW 1878/6/9137, 1 x Betula vel Alnus sp. 3 - LMJ 76489, Fagus deucalionis UNGER (1852, pl. 18, fig. 24), a - 1 x, b - detail of leaf margin, 2 x 4 - NHMW 2001B0017/0004, 1 x Alnus gaudinii (HEER) KNOBLOCH & Z. KVACCEK 5 - NHMW 1878/6/9412, 1 x Y Alnus julianiformis (STERNBERG) Z. KVACCEK & HOLY 6 - IBUG Ett. coll. 284, 1 x Fagus sp., leaf, all 1 x 7 - IBUG Ett. coll. 989 8 - NHMW 1878/6/2491 9 - IBUG Ett. coll. 986 Platanus leucophylla (UNGER) KNOBLOCH 10 - IBUG Ett. coll. 1140, 1 x KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 3 136 Annalen des Naturhistorischen Museums in Wien 105 A Plate 4 Quercus drymeja UNGER, all 1 x 1 - LMJ 76524 A, lectotype (UNGER 1847: pl. 32, fig. 1 right) 2 - LMJ 76549, figured as Juglans hydrophila in UNGER (1850b: pl. 53, fig. 7) 3 - NHMW 1878/6/6557, figured by ETTINGSHAUSEN (1878: pl. 3, fig. 10 - bearing the holotype of Xylomites drymejae ETTINGSHAUSEN). 4 - GBA 2002/01/40 5 - NHMW 1878/6/2447 6 - GBA 2002/01/108 7 - NHMW 1878/6/9388 Quercus mediterranea UNGER, all 1 x 8 - LMJ 76524 B, lectotype of Quercus mediterranea UNGER (1847: pl. 32, fig. 1 top left) 9 - NHMW 1845/0034/0004, syntype of Quercus mediterranea UNGER (1847: pl. 32, fig. 9) 10 - LMJ 76507, UNGER (1852: pl. 18, fig. 1) 11 - GBA 1864/01/05 12 - GBA 2002/01/19 13 - IBUG Ett. coll. 944 14 - NHMW 1878/6/7532 15 - NHMW 1878/6/9381 16 - NHMW 1878/6/9374 Platanus leucophylla (UNGER) KNOBLOCH 17 - NHMW 1878/6/7713, 1 x KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 4 138 Annalen des Naturhistorischen Museums in Wien 105 A Plate 5 Quercus zoroastri UNGER, all 1 x 1 - NHMW 1878/6/2401, neotype of Quercus zoroastri UNGER 2 - LMJ 76866, figured as Juglans hydrophila by UNGER (1850b: pl. 53, fig. 6) 3 - NHMW 1878/6/6478 4 - GBA 2002/01/42 cf. ? Gordonia oberdorfensis KOVAR-EDER 5 - NHMW 1878/6/2009, 1 x 6 - NHMW 1878/6/2038, a - 1 x, b - detail of venation, 2 x 7 - NHMW 1878/6/2025, 1 x 8 - NHMW 1878/6/2005, 1 x KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 5 140 Annalen des Naturhistorischen Museums in Wien 105 A Plate 6 Ternstroemites pereger (UNGER) KOVAR-EDER & Z. KVACCEK comb. nov., all 1 x 1 - NHMW 1878/6/8169, neotype, a - complete leaf, b - detail of margin, 3 x 2 - NHMW 1878/6/8171 3 - NHMW 1853/26/473 4 - GBA 2002/01/41 5 - GBA 2002/01/39 6 - LMJ 76562 B 7 - GBA 2002/01/111 Engelhardia macroptera (BRONGNIART) UNGER, all 1 x 8 - NHMW 1878/6/2698 9 - LMJ 76540, syntype of Carpinus producta UNGER (1850b: pl. 32, fig. 6) Engelhardia orsbergensis (WEBER) JÄHNICHEN, MAI & WALTHER 10 - IBUG Ett. coll. 723, a - 1 x, b - 2 x 11 - NHMW 1878/6/8951, 1 x 12 - GBA 2002/01/22, 1 x Tilia longebracteata ANDRAE 13 - IBUG Ett. coll. 1663, 3 x 14 - IBUG Ett. coll. 1541, as Tilia lignitum in ETTINGSHAUSEN (1869: pl. 42, fig. 6), 1 x 15 - GBA 2002/01/31, 1.5 x Craigia bronnii (UNGER) Z. KVACCEK, BŮZZEK & MANCHESTER 16 - IBUG Ett. coll. 2804a, capsule valve, 2 x 17 - GBA 2002/01/35, capsule valve, 1 x Ulmus plurinervia UNGER, all 1 x 18 - NHMW 1878/6/9665 19 - NHMW 1878/6/9667, neotype of Ulmus plurinervia UNGER 20 - NHMW 1878/6/9082 21 - NHMW 1878/6/9155 22 - NHMW 1878/6/7592 Ulmus parschlugiana KOVAR-EDER & Z. KVACCEK sp. nov., all 2 x 23 - IBUG Ett. coll. 1100, group of fruits, holotype 24 - NHMW 1878/6/9658, paratype 25 - NHMW 1878/6/9081 26 - NHMW 1878/6/9651, paratype KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 6 142 Annalen des Naturhistorischen Museums in Wien 105 A Plate 7 Myrica lignitum (UNGER) SAPORTA 1 - LMJ 76510 right, Quercus lignitum UNGER (1852: pl. 17, fig. 6), 1 x 2 - LMJ 76504, Quercus lignitum UNGER (1852: pl. 17, fig. 1), 1 x 3 - NHMW 1878/6/9312, a - 1 x, b - detail of venation, 1.5 x 4 - NHMW 1878/6/9309, a - 1 x, b - detail of venation, 3 x 5 - GBA 1851/04/10, Dryandroides lignitum (UNGER) ETT. in ETTINGSHAUSEN (1851b: pl. 5, fig. 5), 1 x 6 - LMJ 76503, Quercus lignitum UNGER (1852: pl. 17, fig. 4 - lectotype), 1 x 8 - LMJ 76510 left, Quercus commutata UNGER (1852: pl. 17, fig. 10), 1 x 9 - NHMW 1878/6/7382, a - 1 x, b - detail of venation, 1.5 x Myrica oehningensis (A. BRAUN) HEER 7 - LMJ 76546, Comptonia oeningensis UNGER (1850b: pl. 29, fig. 3), 1 x KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 7 144 Annalen des Naturhistorischen Museums in Wien 105 A Plate 8 Cedrelospermum ulmifolium (UNGER) KOVAR-EDER & Z. KVACCEK comb. nov. (foliage) 1 - NHMW 1878/6/9573, 1 x 2 - GBA 2002/01/38, 1 x 3 - IBUG Ett. coll. 135, 1 x 4 - IBUG Ett. coll. 1085, a - 1 x, b - detail of venation, 5 x 5 - LMJ 76536, lectotype of Comptonia ulmifolia UNGER (1850b: pl. 29, fig. 5), 1 x Cedrelospermum stiriacum (ETTINGSHAUSEN) KOVAR-EDER & Z. KVACCEK comb. nov. (fruit) 6 - IBUG Ett. coll. 2899, 2 x Celtis japeti UNGER 7 - NHMW 1878/6/7654, neotype, 1 x Zelkova zelkovifolia (UNGER) BŮZZEK & KOTLABA, all 1 x 8 - GBA 2002/01/18 9 - NHMW 1987/57, lectotype of Ulmus zelkovaefolia UNGER (1845: pl. 26, fig. 7) 10 - NHMW 1878/6/9642 11 - LMJ 76487, holotype of Ulmus praelonga UNGER (1852: pl. 20, fig. 20) Cedrelospermum ulmifolium (UNGER) KOVAR-EDER & Z. KVACCEK comb. nov. vel Ulmus plurinervia UNGER 12 - LMJ 76488, as Ulmus parvifolia A. BRAUN in UNGER (1852: pl. 20, fig. 22), 2 x ? Prinsepia sp. 13 - NHMW 1878/6/9747, twig, 1 x cf. Rosa sp. 14 - IBUG 1059, 1 x Y ? Buxus cf. egeriana Z. KVACCEK, BŮZZEK & HOLY 15 - LMJ 76502, as Quercus myrtilloides (UNGER 1852: pl. 18, fig. 17), 2 x 17 - GBA 2002/01/14, 1.5 x Y Buxus cf. egeriana Z. KVACCEK, BŮZZEK & HOLY 16 - LMJ 76524C (reverse side), 1.5 x Populus populina (BRONGNIART) KNOBLOCH 18 - LMJ 76506, holotype of Populus aeoli UNGER (1852: pl. 21, fig. 2), 1 x Populus sp. - fructus, all 2 x 19 - NHMW 1878/6/2387 20 - IBUG 1666 21 - NHMW 1878/6/9896 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 8 146 Annalen des Naturhistorischen Museums in Wien 105 A Plate 9 Leguminosites palaeogaea (UNGER) KOVAR-EDER & Z. KVACCEK comb. nov. 1 - NHMW 2002B0017/0003, neotype, 1 x Leguminosites hesperidum (UNGER) KOVAR-EDER & Z. KVACCEK comb. nov., all 1 x 2 - NHMW 1878/6/9109 3 - NHMW 1878/6/8783 4 - GBA 1864/01/21, lectotype of Robinia hesperidum UNGER (1864: 21, pl. 4, fig.13) Leguminosites dionysi (UNGER) KOVAR-EDER & Z. KVACCEK comb. nov. 5 - LMJ 76577 holotype of Cytisus dionysi UNGER (1864: pl. 4, fig. 1), 1 x Leguminosites parschlugianus (UNGER) KOVAR-EDER & Z. KVACCEK comb. nov., all 1 x 6 - NHMW 1878/6/8889 7 - NHMW 1878/6/9895, neotype Podocarpium podocarpum (A. BRAUN) HERENDEEN 8 - IBUG 2245, pod, 1 x 9 - GBA 2002/01/27, pod, 1 x 10 - GBA 2002/01/28 leaflet, 1 x 11 - NHMW 1878/6/8884 leaflet, a - 1 x, b - 2 x "Acacia" parschlugiana UNGER 12 - NHMW 1878/6/9117, neotype, 1 x Phaseolites securidacus UNGER, both 1 X 13 - NHMW 1878/6/2517 14 - LMJ 76569, lectotype of Phaseolites securidacus UNGER (1864: pl. 5, fig. 9) "Juglans" parschlugiana UNGER, both 1 x 15 - LMJ 76559, lectotype of Juglans parschlugiana UNGER (1860: pl. 19, fig. 2) 16 - NHMW 1878/6/2569 Toxicodendron herthae (UNGER) Z. KVACCEK & WALTHER, all 1 x 17 - LMJ 76562, lectotype of Rhus herthae UNGER (1860: pl. 20, fig. 8) 18 - NHMW 1878/6/2027 19 - NHMW 1878/6/9252 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 9 148 Annalen des Naturhistorischen Museums in Wien 105 A Plate 10 Acer integrilobum WEBER sensu WALTHER, all 1 x forma A 1 - GBA 2002/01/49 2 - LMJ 77894 3 - LMJ 76531, syntype of Acer pseudomonspessulanum UNGER (1847: pl. 43, fig. 1) 4 - IBUG Ett. coll. 84 forma B 5 - NHMW 1878/6/6594 6 - NHMW 1878/6/2544 Acer pseudomonspessulanum UNGER emend. STRÖBITZER-HERMANN, all 1 x 7 - LMJ 77899 8 - LMJ 76522, lectotype of Acer pseudomonspessulanum UNGER (1847: pl. 43, fig. 2) 9 - NHMW 1878/6/9156 Acer tricuspidatum BRONN, all 1 x 10 - LMJ 76526 as Acer productum A. BRAUN in UNGER (1847: pl. 42, fig. 8) 11 - IBUG Ett. coll. 1554 12 - LMJ 77900A Acer sp. – fruit, form-group 3 13 - NHMW 1878/6/9253, 2 x Acer sp. – fruit, form-group 2 14 - NHMW 1878/6/9891, 2 x Acer sp. – fruit, form-group 2 ? 15 - IBUG Ett. coll. 2803, 2 x Acer sp. – fruit, form-group 1 16 - IBUG Ett. coll. 1549, 2 x KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 10 150 Annalen des Naturhistorischen Museums in Wien 105 A Plate 11 Paliurus tiliifolius (UNGER) BŮZZEK 1 - NHMW 1878/6/8584, 1 x Paliurus favonii UNGER 2 - GBA 2002/01/36, 1.5 x 3 - NHMW 1878/6/8583, epitype, 1 x 7 - LMJ 76518, lectotype, 1 x Berchemia multinervis (A. BRAUN) HEER, all 1 x 4 - NHMW 1878/6/9107 5 - NHMW 1878/6/2078 Cotinus (?) aizoon (UNGER) KOVAR-EDER & Z. KVACCEK comb. nov., all 1 x 6 - GBA 2002/01/8 8 - LMJ 76575, lectotype of Rhamnus aizoon UNGER (1864: pl. 3, fig. 44) 9 - LMJ 77607 10 - GBA 2002/01/11 Ailanthus confucii UNGER 11 - NHMW 1878/6/2121, 2 x Fraxinus primigenia UNGER, all 1 x 12 - IBUG Ett. coll. 1387 13 - Neotype, NHMW 1878/6/8155 14 - NHMW 1878/6/9889 15 - IBUG Ett. coll. 1385 Nerium sp. 16 - IBUG Ett. coll. 1405, capsule, 1.5 x 17 - NHMW 1878/6/8173, a - 1 x, b - detail of venation, 4 x 18 - NHMW 1878/6/8175, 1 x Smilax sagittifera HEER emend. HANTKE, all 2 x 19 - GBA 1847/03/20 20 - IBUG Ett. coll. 399 KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 11 152 Annalen des Naturhistorischen Museums in Wien 105 A Plate 12 "Celastrus" europaea UNGER, both 1 x 1 - LMJ 76576, lectotype of Celastrus europaeus UNGER (1864: pl. 2, fig. 10) 2 - LMJ 76563, syntype of Celastrus europaeus UNGER (1864: pl. 2, fig. 12) "Evonymus" latoniae UNGER 3 - LMJ 76574, lectotype of Evonymus latoniae UNGER (1864: pl. 2, fig. 25), 1.5 x 4 - NHMW 1878/6/2063, 1 x 5 - NHMW 1878/6/2742, 1 x "Cornus" ferox UNGER, both 1 x 6 - NHMW 1878/6/8109 part, neotype 7 - NHMW 1878/6/ 6566 counterpart, neotype ? Chaneya sp., both 1 x 8 - NHMW 1878/6/8741 9 - NHMW 1878/6/8742 "Quercus" daphnes UNGER 10 - LMJ 76591 as Achras lycobroma UNGER (1866: pl. 8, fig. 1), 1 x 11 - LMJ 76590 counterpart to the holotype of Rhododendron flos-saturni UNGER (1866: pl. 12, fig. 15), a - 1 x, b - detail of venation, 2 x 12 - NHMW 1878/6/9460, a - 1 x, b - detail of venation, 2 x 13 - NHMW 1878/6/7557, 1 x 14 - NHMW 1878/6/9459, 1 x 15 - LMJ 76525, lectotype of Quercus daphnes UNGER (1847: pl. 31, fig. 3), 1 x KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 12 154 Annalen des Naturhistorischen Museums in Wien 105 A Plate 13 Mahonia (?) aspera (UNGER) KOVAR-EDER & Z. KVACCEK comb. nov. 1 - LMJ 76571, as Ilex sphenophylla UNGER (1864: pl. 3, fig. 3), 2 x 2 - LMJ 76579, as Ilex cyclophylla UNGER (1864: pl. 3, fig. 8), 1 x 3 - LMJ 76529, lectotype of Quercus aspera UNGER (1847: pl. 30, fig. 2), 1 x 4 - LMJ 76532, syntype of Quercus aspera UNGER (1847: pl. 30, fig. 1), 1 x 5 - GBA 2002/01/45, 1 x 6 - NHMW 1878/6/2758, 1 x 7 - GBA 2002/01/46, 1 x 8 - GBA 2002/01/44, 1 x Prinsepia serra (UNGER) KOVAR-EDER & Z. KVACCEK comb. nov. 9 - LMJ 76528, lectotype of Quercus serra UNGER (1847: pl. 30, fig. 5), a - 1 x, b - detail of venation and margin, 1.5 x 10 - LMJ 76495, epitype of Quercus serra UNGER (1852: pl. 18, fig. 16), a - 1 x, b - detail of venation and margin, 2 x 11 - NHMW 1878/6/9509, 1 x 12 - NHMW 1878/6/9671, 1 x 13 - NHMW 1878/6/7538, a - 1 x, b - detail of venation and margin, 1.5 x 14 - NHMW 1878/6/9502, 1 x 15 - NHMW 1878/6/9527, 1 x 16 - GBA 2002/01/16, 1 x 17 - GBA 2002/01/15, 1 x KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 13 156 Annalen des Naturhistorischen Museums in Wien 105 A Plate 14 Populus populina (BRONGNIART) KNOBLOCH 1 - LMJ 76505 as Populus latior A. BRAUN in UNGER (1852: pl. 21, fig. 4), 0.8 x Ailanthus pythii (UNGER) KOVAR-EDER & Z. KVACCEK comb. nov., all 1 X 2 - NHMW 1878/6/2649, a pinnate leaf 3 - NHMW 1878/6/2525, leaflet 4 - LMJ 76557, leaflet, lectotype of Sapindus pythii UNGER (1860: pl. 14, fig. 8) 5 - NHMW 1878/6/6484, leaflet, as Sapindus pythii UNGER in ETTINGSHAUSEN (1878 b: pl. 3, fig. 5 bearing Sphaeria palaeo-sapindi ETTINGSHAUSEN) KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 14 158 Annalen des Naturhistorischen Museums in Wien 105 A Plate 15 Dicotylophyllum sp. 1 1 - NHMW 1878/6/2091, 1 x Dicotylophyllum sp. 2, both 1 x 2 - GBA 2002/01/109 3 - NHMW 1878/6/6555 Dicotylophyllum sp. 6 4 - IBUG Ett. coll. 1083, 1 x 5 - IBUG Ett. coll. 1084, a - 1 x, b - detail of margin and venation, 2 x Saportaspermum sp., all 2 x 6 - NHMW 1878/6/8029 7 - NHMW 1878/6/8028 8 - IBUG Ett. coll. 1346 Dicotylophyllum sp. 3 9 - GBA 2002/01/20, a - 1.5 x, b - 1 x 10 - NHMW 1878/6/8571, 1 x Dicotylophyllum sp. 4 11 - GBA 2002/01/21 Dicotylophyllum sp. 5 12 - NHMW 1878/6/7507, 1 x Antholithes stiriacus KOVAR-EDER & Z. KVACCEK sp. nov. (flowers) 13 - IBUG Ett. coll. 432, ca. 7 x 14 - IBUG Ett. coll. 427, ca. 7 x 15 - NHMW 1878/6/9870, holotype, 5 x Cypselites sp., seed 16 - IBUG Ett. coll. 1374, ca. 6 x KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis Plate 15

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The Miocene flora of Parschlug (Styria, Austria)revision and synthesis

The Miocene flora of Parschlug (Styria, Austria)revision and synthesis

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