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
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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
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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
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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.
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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.
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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.
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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,
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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).
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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).
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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.
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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.
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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.
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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
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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.
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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.
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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).
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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.
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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.
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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:
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- 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. The study was performed within the frame of the project 13741-BIO financed by the Austrian
Science Fund and the grant project of Ministry of Education, CR No. J13/98:113100006. Its results shall
also contribute to the efforts of the ESF (European Science Foundation) network EEDEN (Environment and
Ecosystem Dynamics of the Eurasian Neogene).
118
Annalen des Naturhistorischen Museums in Wien 105 A
References
ANDRAE, K.-J. (1861): Ein neuer Beitrag zur Tertiär-Flora Siebenbürgens. – Abh. Naturwiss.
Vereins Sachsen, Thüringen, 2: 23-30. – Dresden.
ANDREÁNSZKY, G. (1955): Új fajok Magyaregregy alsó-helvéti emeletéboől. – Ann. Inst. geol.
publ. hung., 44: 14-16, 152-153. – Budapest.
ASH, A., ELLIS, B., HICKEY, L.J., JOHNSON, K., WILF, P. & WING, S. (1999): Manual of Leaf
Architecture – morphological description and categorization of dicotyledonous and netveined monocotyledonous angiosperms. – 65 p. – Washington (Smithsonian Inst.).
BERGER, W. (1952): Die altpliozäne Flora der Congerienschichten von Brunn-Vösendorf bei
Wien. – Palaeontogr. B, 92: 79-121. – Stuttgart.
––– (1953): Pflanzenreste aus dem miozänen Ton von Weingraben bei Draßmarkt (Mittelburgenland) II. – Sitzungsber. Österr. Akad. Wiss. math.-naturwiss. Kl., Abt. 1, 162/1,2:
17-24. – Wien.
––– (1955): Jungtertiäre Pflanzenreste aus dem unteren Lavanttal in Ostkärnten. – N. Jb. Geol.
Paläont. Abh., 100: 402-430. – Stuttgart.
––– & ZABUSCH, F. (1953): Die obermiozäne (sarmatische) Flora der Türkenschanze in Wien.
– N. Jb. Geol. Paläont. Abh., 98/2: 226-276. – Stuttgart.
BÖHME, M., GREGOR, H.-J. & HEISSIG, K. (2002): The Ries and Steinheim Meteorite Impacts and
their Effect on Environmental Conditions in Time and Space. – In: BUFFETAUT E. &
KOEBERL C. (eds.) Geological and Biological Effects of Impact Events. – 217-235. –
Berlin, Heidelberg, New York (Springer-Verlag).
C. (1971): Tertiary flora from the northern part of the Petipsy area (North-Bohemian
Basin). – Rozpr. Ústr. Úst. Geol., 36: 1-119. – Praha.
BŮZEK,
––– , HOLY, F. & KVACEK, Z. (1996): Early Miocene flora of the Cypris Shale (Western
Bohemia). – Acta Mus. Nation. Pragae, Ser. B., Hist. Natur., 52: 1-72. – Praha.
COLLINSON, M., KVACEK, Z. & ZASTAWNIAK, E. (2001): The aquatic plants Salvinia
(Salviniaceae) and Limnobiophyllum (Arales) from the Late Miocene flora of Sośnica
(Poland). – Acta Palaeobot., 41: 253-282. – Kraków.
DENK, T. (2002): Revision of Fagus (oak family) from the Tertiary of Europe and south-western
Asia. – Abstracts, 6th European Paleobotany-Palynology Conference, Athens. – 75-76. –
Athens.
––– & MELLER, B. (2001): Systematic significance of the cupule/nut complex in living and
fossil Fagus. – Int. J. Plant Sci., 162: 869-897. – Chicago.
ERDEI, B. (in press): A vegetation and climate reconstruction of Sarmatian (Middle Miocene)
sites from NE and W Hungary. – Acta Univ. Carol. Geol. – Praha.
ETTINGSHAUSEN, C.V. (1851a): Die Tertiaer-Floren der oesterreichischen Monarchie. 1. Fossile
Flora von Wien. – 1-35. – Wien (K.K. Hof- u. Staatsdruckerei).
––– (1851b): Die Proteaceen der Vorwelt. – Sitzungsber. kaiserl. Akad.Wiss., math.-naturwiss. Cl., 7: 732-745. – Wien.
––– (1869): Die fossile Flora des Tertiär-Beckens von Bilin. III. – Denkschr. kaiserl. Akad.
Wiss. math.-naturwiss. Cl., 28: 1-110. – Wien.
––– (1878a): Beiträge zur Erforschung der Phylogenie der Pflanzenarten. I-II. – Denkschr.
kaiserl. Akad. Wiss. math.-naturwiss. Cl., 38: 65-80. – Wien.
KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis
119
––– (1878b): Beiträge zur Kenntniss der fossilen Flora von Parschlug in Steiermark. –
Denkschr. kaiserl. Akad. Wiss. math.-naturwiss. Cl., 38: 81-92. – Wien.
––– (1882): Beiträge zur Erforschung der Phylogenie der Pflanzenarten. III-VII. – Denkschr.
kaiserl. Akad. Wiss. math.-naturwiss. Cl., 43: 93-102. – Wien.
––– (1888): Die fossile Flora von Leoben in Steiermark. 1., 2. Theil. – Denkschr. kaiserl.
Akad. Wiss. math.-naturwiss. Cl., 54: 261-318; 319-384. – Wien.
––– (1890): Die fossile Flora von Schönegg bei Wies in Steiermark. 1. Theil. – Denkschr. kaiserl. Akad. Wiss. math.-naturwiss. Cl., 57: 61-112. – Wien.
––– & STANDFEST, F. (1888): Über Myrica lignitum UNGER und ihre Beziehungen zu den
lebenden Myrica-Arten. – Denkschr. kaiserl. Akad. Wiss. math.-naturwiss. Cl., 54: 255260. – Wien.
GIVULESCU, R. (1962): Die fossile Flora von Valea Neagra, Bezirk Crisana, Rumänien. –
Palaeontogr. B, 110: 128-187. – Stuttgart.
––– (1998): Mahonia sp. (n. sp. ?) une nouvelle Mahonia du Pannonien inférieur de Valea
Cri‚ului (Bihor Roumanie). – Stud. Univ. Babe‚-Bolyai, Geol., 43/2: 3-5. – Cluj – Napoca.
GOEPPERT, H. (1855): Die tertiäre Flora von Schossnitz in Schlesien. – 52 p. – Görlitz (Heyn'sche
Buchhandl.).
GREGOR, H.-J. (1982): Die jungtertiäre Floren Süddeutschlands – Paläokarpologie, Phytostratigraphie, Paläoökologie, Paläoklimatologie. – 278 p. – Stuttgart (F. Enke Verl.).
––– (1986): Zur Flora des Randecker Maares (Miozän, Baden-Württemberg). – Stuttgarter
Beitr. Naturkunde, B, 122: 1-29. – Stuttgart.
HABLY, L. (1985): Catalogue of the Hungarian Cenozoic leaf-floras. – Studia Botanica Hungarica,
18: 5-58. – Budapest.
––– (1992): Distribution of legumes in the Tertiary of Hungary. – In: HERENDEEN, P.S. &
DILCHER, D.L., Advances in legume systematics, part 4, the fossil record. – 169-187. –
Kew (Royal Bot. Gardens).
––– (2001): Fruits and leaves of Ailanthus DESF. from the Tertiary of Hungary. – Acta
Palaeobot., 41: 207-219. – Kraków.
––– (2002): The Middle Miocene flora of Magyaregregy – as shown by a study of recent
collections. Abstracts, 6th European Paleobotany-Palynology Conference, Athens. – 9192. – Athens.
––– , ERDEI, B. & KVACEK, Z. (2001): 19th century´s palaeobotanical types and originals of the
Hungarian Natural History Museum. – 235 p. – Budapest (Hungarian Natural History
Museum).
––– , KVACEK, Z. & MANCHESTER, S.R. (2000): Shared taxa of land plants in the Oligocene of
Europe and North America in context of Holarctic phytogeography. – Acta Univ. Carol.
Geol., 44 (1): 59-74. – Praha.
––– & THIÉBAUT, M. (2002): Revision of Cedrelospermum (Ulmaceae) fruits and leaves from
the Tertiary of Hungary and France. – Palaeontogr. B, 262: 71-90. – Stuttgart.
HANTKE, R. (1954): Die fossile Flora der obermiozänen Oehninger-Fundstelle Schrotzburg (Schienerberg, Süd-Baden). – Denkschr. Schweiz. Naturforsch. Ges., 80 (2): 27-118. – Zürich.
HEER, O. (1859): Flora tertiaria Helvetiae. III. – 378 p. – Winterthur (Wurster & Comp.).
120
Annalen des Naturhistorischen Museums in Wien 105 A
HEIZMANN, E.P.J. (1983): Die Gattung Cainotherium (Cainotheriidae) im Orleanium und im
Astaracium Süddeutschlands. – Eclog. Geol. Helvetiae, 76/ 3: 781-825. – Basel.
HERENDEEN, P.S. (1992a): A re-evaluation of the fossil genus Podogonium HEER. In:
HERENDEEN, P.S. & DILCHER, D.L., Advances in legume systematics, part 4, the fossil
record. – 3-18. – Kew (Royal Bot.Gardens).
––– (1992b): Podocarpium podocarpum comb. nov., the correct name for Podogonium knorrii
HEER, nom. illegit. (fossil Fabaceae). – Taxon, 41: 731-736. – Utrecht.
––– (1992c): The fossil history of the Leguminosae from the Eocene of southeastern North
America. In: HERENDEEN, P.S. & DILCHER, D.L., Advances in legume systematics, part 4,
the fossil record. – 85-160. – Kew (Royal Bot. Gardens).
HICKEY, L.J. (1977): Stratigraphy and Paleobotany of the Golden Valley Formation (Early
Tertiary) of Western North Dakota. – Mem. Geol. Soc. Amer., 150: 1-181. – Boulder.
ILJINSKAYA, I.A. (1962): Tortonskaja flora Svoshovice i pliocenovye flory Zakarpatja. – Paleont.
Zhur., 1962/3: 102-110. – Moskva.
––– (1964): The Tortonian flora of Swoszowice. – Trudy Bot. Inst. AN SSSR, ser. 8, Paleobotanica, 5: 113-144. – Leningrad.
––– & SHTEFYRTSA, A.G. (1971): Iskopaemye Smilacaceae i Dioscoreaceae iz Miotsena
Moldavii. – Bot Zhur., 56/2: 175-183. – Leningrad.
JÄHNICHEN, H., MAI, H.D. & WALTHER, H. (1977): Blätter und Früchte von Engelhardia LESCH. ex
BL. (Juglandaceae) aus dem europäischen Tertiär. – Fedd. Repert., 88: 323-363. – Berlin.
KNOBLOCH, E. (1968): Bemerkungen zur Nomenklatur tertiärer Pflanzenreste. – Acta Mus.
Nation. Pragae, B, 24/3: 121-152. – Praha.
––– & KVACEK, Z. (1976): Miozäne Blätterfloren vom Westrand der Böhmischen Masse. –
Rozpr. Ústr. Úst. Geol., 42: 1-131. – Praha.
––– & KVACEK, Z. (1982): Miozäne Pflanzenreste aus der Umgebung von Tamsweg (Niedere
Tauern). – Acta Univ. Carolinae, Geol., 1981/2: 95-120.
––– & KVACEK, Z. (1996): Miozäne Floren der südböhmischen Becken. – Sbor. Geol. Ved,
Paleont., 33: 39-77. – Praha.
KOVAR, J. (1982): Eine Blätter-Flora des Egerien (Ober-Oligozän) aus marinen Sedimenten der
Zentralen Paratethys im Linzer Raum (Österreich). – Beitr. Paläont. Österr., 9: 1-209. – Wien.
KOVAR-EDER, J. (1988): Obermiozäne (pannone) Floren aus der Molassezone Österreichs. –
Beitr. Paläont. Österr., 14: 19-121. – Wien.
––– & KRAINER, B. (1991): Flora und Sedimentologie der Fundstelle Reith bei Unterstorcha,
Bezirk Feldbach in der Steiermark (Kirchberger Schotter, Pannonium C, Miozän). – Jb.
Geol. Bundesanst., 134/4: 737-771. – Wien.
––– & MELLER, B. (2001): Plant assemblages from the hanging wall sequence of the opencast
mine Oberdorf N Voitsberg, Styria (Austria, Early Miocene, Ottnangian). – Palaeontogr.
B, 259: 65-112. – Stuttgart.
––– , GIVULESCU, R., HABLY, L., KVACEK, Z., MIHAJLOVIC, D., TESLENKO, Y., WALTHER, H.,
ZASTAWNIAK, E. (1994): Floristic changes in the areas surrounding the Paratethys during
Neogene time. – In: BOULTER, M.C. & FISHER, H.C. (eds.): Cenozoic Plants and Climates
of the Arctic. – NATO-ASI Ser. I: Global Environmental Change, 27: 347-369. –
Heidelberg (Springer).
KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis
121
––– , KVACEK, Z., ZASTAWNIAK, E., GIVULESCU, R., HABLY, L., MIHAJLOVIC, D., TESLENKO, Y.
& WALTHER, H. (1996): Floristic trends in the vegetation of the Paratethys surrounding
areas during Neogene time. – In: BERNOR, R., FAHLBUSCH, V. & MITTMANN, H.-W. (eds.):
The Evolution of Western Eurasian Later Neogene Faunas. – 399-409. – New York
(Columbia University Press).
KRISHTOFOVICH, A.N. & BAYKOVSKAYA, T.N. (1965): Sarmatskaya flora Krynki. – 136 p. –
Moskva-Leningrad (Nauka).
KVACEK, Z., BŮZEK, C. & HOLY, F. (1982): Review of Buxus fossils and a new large-leaved species from the Miocene of Central Europe. – Rev. Palaeobot. Palyn., 37: 361-394. –
Amsterdam.
––– , BŮZEK, C. & MANCHESTER, S. R. (1991): Fossil fruits of Pteleaecarpum WEYLAND - tiliaceous, not sapindaceous. – Bot. Gaz., 153: 522-523. – Chicago.
––– & HURNÍK, S. (2000): Revision of Early Miocene plants preserved in baked rocks in the
North Bohemian Tertiary. – Acta Mus. Nation. Pragae, B, Hist. Nat. Ser., 56: 1-48. – Praha.
––– , MANCHESTER, S.R., ZETTER, R. & PINGEN, M. (2002 a): Fruits and seeds of Craigia bronnii
(Malvaceae – Tilioideae) and associated flower buds from the late Miocene Inden Formation,
Lower Rhine Basin, Germany. – Rev. Palaeobot. Palyn., 119: 311-324. – Amsterdam.
––– & SAKALA, J. (1999): Twig with attached leaves, fruits and seeds of Decodon (Lythraceae)
from the Lower Miocene of northern Bohemia, and implications for the identification of
detached leaves and seeds. – Rev. Palaeobot. Palynol., 107: 201-222. – Amsterdam.
––– , VELITZELOS, D., & VELITZELOS, E. (2002 b): Late Miocene flora of Vegora, Macedonia,
N. Greece. – 173 p. – Athens (Athens University).
––– & WALTHER, H. (1984 a): Nachweis tertiärer Theaceen Mitteleuropas nach blatt-epidermalen Untersuchungen I. Teil – Epidermale Merkmalkomplexe rezenter Theaceen. –
Fedd. Repert., 95: 209-227. – Berlin.
––– & WALTHER, H. (1984 b): Nachweis tertiärer Theaceen Mitteleuropas nach blatt-epidermalen Untersuchungen II. Teil – Bestimmung fossiler Theaceen-Sippen. – Fedd. Repert.,
95: 331-346. – Berlin.
––– & WALTHER, H. (1989): Paleobotanical studies in Fagaceae of the Tertiary. – Pl. Syst.
Evol., 162: 213-229. – Wien.
––– & WALTHER, H. (1991): Revision der mitteleuropäischen tertiären Fagaceen nach blattepidermalen Charakteristiken IV. Teil Fagus LINNÉ. – Fedd. Repert., 102: 471-534. – Berlin.
––– & WALTHER, H. (1998): The Oligocene volcanic flora of Kundratice near Litomerice,
Ceské stredohorí Volcanic Complex (Czech Republic) – a review. – Acta Mus. Nation.
Pragae, B, 54/1-2: 1-42. – Praha.
LIU, YUSHENG, ZETTER, R., MOHR, B.A.R. & FERGUSON, D.K. (2001): The flowers of an extinct
legume from the Miocene of southern Germany. – Palaeontogr. B, 256: 159-174. – Stuttgart.
MAI, H.D. (1983): Studien an Endocarpien europäischer und westasiatischer Arten der Gattung
Acer L. (Aceraceae). – Gleditschia, 10: 37-57. – Berlin.
––– (1984a): Die Endocarpien bei der Gattung Acer L. (Aceraceae) – eine biosystematische
Studie. – Gleditschia, 11: 17-46. – Berlin.
––– (1984b): Karpologische Untersuchungen der Steinkerne fossiler und rezenter
Amygdalaceae (Rosales). – Fedd. Repert., 95: 299-329. – Berlin.
122
Annalen des Naturhistorischen Museums in Wien 105 A
––– (1986): Über Typen und Originale tertiärer Arten von Pinus L. (Pinaceae) in mitteleuropäischen Sammlungen – ein Beitrag zur Geschichte der Gattung in Europa. – Fedd.
Repert., 97: 571-605. – Berlin.
––– (1994): Fossile Koniferenreste in der meridionalen Zone Europas. – Fedd. Repert., 105:
207-227. – Berlin.
––– (1995): Tertiäre Vegetationsgeschichte Europas. – 691 p. – Jena, Stuttgart, New York
(Gustav Fischer Verlag).
––– & WALTHER, H. (1988): Die pliozänen Floren von Thüringen, Deutsche Demokratische
Republik. – Quartärpaläontologie, 7: 55-297. – Berlin.
MANCHESTER, S.R. (1989): Systematics and fossil history of the Ulmaceae. – Syst. Assoc. Spec.
Vol., 40B: 221-251. – Kew.
––– (1994): Inflorescence bracts of fossil and extant Tilia in North America, Europe, and
Asia: patterns of morphological divergence and biogeographic history. – Amer. J. Bot.,
81: 1176-1185. – New York.
MELLER, B., KOVAR-EDER, J. & ZETTER, R. (1999): Lower Miocene diaspore-, leaf-, and palynomorph – assemblages from the base of the lignite-bearing sequence in the opencast
mine Oberdorf N Voitsberg (Styria, Austria) as indication of a "Younger Mastixioid"
vegetation. – Palaeontogr. B, 252: 123-179. – Stuttgart.
MEYER, W. & MANCHESTER, S.R. (1997): The Oligocene Bridge Creek flora of the John Day
Formation, Oregon. – Univ. California Publ. Geol. Sci., 141: 1-195. – Berkeley.
MOTTL, M. (1970): Die jungtertiären Säugetierfaunen der Steiermark, Südost-Österreichs. –
Mitteil. Mus. Bergbau, Geol. u. Tech. Landesmus. Joanneum, 31. – Graz.
NEUBAUER, F., FRITZ, H., GENSER, J., KURZ, W., NEMES, F., WALLBRECHER, E., WANG, X. &
WILLINGSHOFER, E. (2000): Structural evolution within an extruding wedge: model and
application to the Alpine-Pannonian system. – In: LEHNER, F.K. & URAI, J.L. (eds.):
Aspects of tectonic faulting. – 141-153. – Berlin (Springer).
OBERHAUSER, R. (ed.) (1980): Der geologische Aufbau Österreichs. – 695 p. – Wien, New York
(Springer).
PALAMAREV, E.H. & PETKOVA, A.S. (1987): La macroflore du Sarmatien. – In: TZANKOV, V.
(ed.): Les fossiles de Bulgarie VIII. 1. – 273 p. – Sofia (Acad. Bulg. Sci.).
PENNINGTON, T.D. (1991): The genera of Sapotaceae. – 300 p. – Kew, New York (R. Bot.
Gardens – N. Y. Bot. Gard.)
PETRASCHEK, W.E. (1922-1929): Kohlengeologie der österreichischen Teilstaaten I-II. – 484 p.
Katowice (Kattowitzer Buchdruck. u. Verlags. Sp. Akc.).
PINGEN, M., FERGUSON, D.K. & COLLINSON, M.E. (1994): Homalanthus costatus MAI: a new Miocene
fruit of Cinnamomum SCHAEFFER (Lauraceae). – Palaeontogr. B, 232: 155-174. – Stuttgart.
REID, E.M. & CHANDLER, M.E.J. (1926): The Bembridge flora. – Brit. Mus. Catal. Cainoz. Plants
I. 8: 1-206. – London.
RIEDERLE, R. (1997): Die Sandgrube Ursberg bei Thannhausen - Stratigraphie einer neuen miozänen
Fundstelle aus der Molasse Bayerisch-Schwabens. – Doc. naturae, 110: 103 - 118. – München.
––– & GREGOR, H.-J. (1997): Die Tongrube Kirrberg bei Balzhausen – eine neue Fundstelle
aus der Oberen Süßwassermolasse Bayerisch-Schwabens – Flora, Fauna, Stratigraphie. –
Doc. naturae, 110: 1-53. – München.
KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis
123
RÜFFLE, L. (1963): Die obermiozäne (sarmatische) Flora vom Randecker Maar. – Paläont. Abh.,
1(3): 139-298. – Berlin.
SACHSENHOFER, R. F., KUHLEMANN, J. & REISCHENBACHER, D. (2002): Das Miozän der östlichen Norischen Senke. In: G.W. MANDL (ed.): Arbeitstagung 2001. – 135-145. – Wien (Geol. B.-Anst.).
SAPORTA, G. DE (1865): Études sur la végétation du sud-est de la France a l´époque tertiaire. –
Ann. Sci. Natur. Sér. 5 (Bot.), 4: 5 (149)-264 (408). – Paris.
SCHIMPER, W. PH. (1874): Traité de paléontologie végétale. III. – 896 p. – Paris (J.B. Baillière et fils).
SCHMID, H. (1983): Eine miozäne Blatt- und Fruchtflora von der Fossilfundstelle Sandgrube
Dumerth in Burtenbach. – Günzburger Hefte, 20. – Günzburg.
SCHMID, W. & GREGOR, H.-J. (1983): Gallenbach – eine neue mittelmiozäne Fossilfundstelle in
der westlichen Oberen Süßwassermolasse Bayerns. – Ber. Naturwiss. Ver. Schwaben, 87
(3/4): 51-63. – Augsburg.
SCHMIDT, C. (1976): Obermiozäne Flora von Derching bei Augsburg. – Ber. Naturwiss. Ver.
Schwaben, "Aus der Schwäbischen Heimat", 1976 (3/4): 52-56. – Augsburg.
––– (1980): Ein Profil von pflanzenführenden Schichten der Sandgrube Derching. – Ber. Naturwiss. Ver. Schwaben, 84 (1/2): 13-15. – Augsburg.
SCHMITT, H. & BUTZMANN, R. (1997): Entrischenbrunn – Statistische Untersuchungen an einer
neuen Florenfundstelle aus der Oberen Süßwassermolasse im Landkreis Pfaffenhofen a.
d. Ilm. – Doc. naturae, 110: 55-87. – München.
SCHWEIGERT, G. (1992): Die untermiozäne Flora (Karpatium, MN 5) des Süßwasserkalkes von
Engelswies bei Meßkirch (Baden-Würtemberg). – Stuttgarter Beitr. Naturkunde Ser. B,
188: 1-55. – Stuttgart.
STIZENBERGER, E. (1851): Übersicht der Versteinerungen des Grossherzogthums Baden. – 144 p.
– Freiburg i. Breisgau (Verl. Univ.-Buchhandlung J. Diernfellner).
STRÖBITZER, M. (1999): Die fossilen Blattvergesellschaftungen von Lintsching (Tamsweger
Becken, Salzburg, Miozän). – Beitr. Paläont., 24: 91-153. – Wien.
STRÖBITZER-HERMANN, M. (2003): Systematik, Variabilität, regionale und stratigraphische Verbreitung und Ökologie der Gattung Acer L. in Mitteleuropa vom Oligo- bis ins Pliozän.
PhD thesis, Univ. Wien.
TAKHTAJAN, A. (1982): Magnoliophyta fossilia URSS. 2. Ulmaceae - Betulaceae. – 216 p. –
Leningrad (Nauka).
TANAI, T. (1983): Revisions of Tertiary Acer from East Asia. Jour. Fac. Sci., Hokkaido Univ.,
Ser. IV, 20/4: 291-390. – Sapporo.
––– & OZAKI, K. (1977): The genus Acer from the upper Miocene in Tottori prefecture,
western Japan. Jour. Fac. Sci, Hokkaido Univ., Ser. IV, 17/4: 575-606. – Sapporo.
TRYON, A.F. & LUGARDON, D. (1990): Spores of the Pteridophytes. – New York.
UNGER, F. (1841-1847): Chloris protogaea. – CX, 150 p. – Leipzig (W.Engelmann).
––– (1848): Die fossile Flora von Parschlug. – Steiermärk. Z., n. F., 9/1: 3-39. – Graz.
––– (1849): Blätterabdrücke aus dem Schwefelflötze von Schwoszowice in Galicien. – Separatum, from Haidingers Naturwiss. Abh., 3:121-128. – Wien.
––– (1850a): Genera et species plantarum fossilium. – XL, 627 p. – Wien (W. Braumüller).
124
Annalen des Naturhistorischen Museums in Wien 105 A
––– (1850b): Die fossile Flora von Sotzka. – Denkschr. kaiserl. Akad. Wiss. math.-naturwiss.
Cl., 2: 127-197. – Wien.
––– (1852): Iconographia plantarum fossilium. – Denkschr. kaiserl. Akad. Wiss. math.-naturwiss. Cl., 4: 73-118. – Wien.
––– (1860): Sylloge plantarum fossilium. I. – Denkschr. kaiserl. Akad. Wiss. math.-naturwiss.
Cl., 19: 1-48. – Wien.
––– (1864): Sylloge plantarum fossilium. II. – Denkschr. kaiserl. Akad. Wiss. math.-naturwiss. Cl., 22: 1-36. – Wien.
––– (1866): Sylloge plantarum fossilium. III. – Denkschr. kaiserl. Akad. Wiss. math.-naturwiss. Cl., 25: 1-76. – Wien.
––– (1867): Die fossile Flora von Kumi auf der Insel Euboea. – Denkschr. kaiserl. Akad.
Wiss. math.-naturwiss. Cl., 27: 27-87. – Wien.
VELITZELOS, D. ed. (2002): Field guide to the Neogene of the Island of Evia – Early Miocene
flora of Kymi. 6th European Paleobotany-Palynology Conference, Athens. – 61 p. –
Athens (University of Athens).
WALTHER, H. (1972): Studien über tertiäre Acer Mitteleuropas. – Abh. Staatl. Mus. Mineral.
Geol., 19: 1-309. – Dresden.
––– & ZASTAWNIAK, E. (1991): Fagaceae from Sośnica and Malczyce (near Wroclaw,
Poland). A revision of original materials by Goeppert 1852 and 1855 and a study of new
collections. – Acta Palaeobot., 31: 153-199. – Kraków.
WANG, YUFEI & MANCHESTER, S.R. (2000): Chaneya, a new genus of winged fruit from the
Tertiary of North America and eastern Asia. – Int. J. Plant Sci., 161: 167-178. – Chicago.
WEBER, L. & WEISS, A. (1983): Bergbaugeschichte und Geologie der österreichischen Braunkohlenvorkommen. – Archiv Lagerstättenforsch. Geol. Bundesanst., 4: 1-317. – Wien.
WEYLAND, H. (1937): Beiträge zur Kenntnis der rheinischen Tertiärflora. II. Erste Ergänzungen
und Berichtigungen zur Flora der Blätterkohle und des Polierschiefers von Rott im
Siebengebirge. – Palaeontogr. B, 83: 67-122. – Stuttgart.
ZASTAWNIAK, E. & WALTHER, H. (1998): Betulaceae from Sośnica near Wroclaw (Poland) – a
revision of GOEPPERTs original materials and a study of more recent collections. – Acta
Palaeobot., 38: 87-145. – Kraków.
KOVAR-EDER & al: The Miocene Flora of Parschlug – Revision and Synthesis
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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"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
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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
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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)
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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
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