ABHANDLUNGEN DER GEOLOGISCHEN BUNDESANSTALT
Abh. Geol. B.-A. ISSN 0378-0864 ISBN 978-3-85316-058-9 Band 65 S. 181–210
Wien, 10. 11. 2010
Fifty Years of Geological Cooperation between Austria, the Czech Republic and the Slovak Republic
An Early Eocene Fauna and Flora from “Rote Kirche” in Gschliefgraben
near Gmunden, Upper Austria
alFréd dulai1, lenka hradecká2, MaGda konzalová3, GyörGy less4, lilian Švábenická2 & harald lobitzer5
4 Text-Figures, 7 Plates, 4 Tables
Österreichische Karte 1:50.000
Blatt 66 Gmunden
Ultrahelveticum
Salzkammergut
Gschliefgraben
Palynomorphs
Nannofossils
Foraminifera
Brachiopods
Rote Kirche
Ypresian
Contents
Zusammenfassung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Location and Geological Setting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Previous Work . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Studied Material . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Brachiopods (A. Dulai) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Systematic Notes on Brachiopods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Taxonomic Composition of the Brachiopod Fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Palaeoecology, Palaeoenvironment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Small Foraminifera (L. Hradecká). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Larger Foraminifera (Gy. Less) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Systematic Part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Calcareous Nannofossils (L. Švábenická) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Microflora – Preliminary Results (M. Konzalová) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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Beiträge zur früheozänen Fauna und Flora der Lokalität Rote Kirche im Gschliefgraben bei Gmunden, Oberösterreich
Zusammenfassung
Erstmals wird eine Brachiopoden-Vergesellschaftung aus dem Eozän Österreichs beschrieben. Sie umfasst 6 Taxa (Gryphus kickxii, Meznericsia hantkeni,
Terebratulina tenuistriata, Orthothyris pectinoides, Megathiris detruncata, Argyrotheca sabandensis?) und stammt aus mergeligen Kalken bzw. sandigen
Mergeln des Ultrahelvetikums der Lokalität Rote Kirche im Gschliefgraben bei Gmunden. Die Dominanz der Genera Gryphus und Terebratulina spricht für
einen relativ tieferen Ablagerungsraum, wahrscheinlich im äußeren Schelfbereich. Groß- und Kleinforaminiferen, kalkige Nannofossilien und Palynomorphen / Dinoflagellaten ermöglichen eine Einstufung der hangenden Ablagerungen des Aufschlusses Rote Kirche als frühes Eozän (spätes Ypresium). Eine
neue Großforaminiferen-Chronosubpecies, Orbitoclypeus multiplicatus gmundenensis, die für die Zone SBZ 10 charakteristisch ist, wird beschrieben. Die
Palynomorphen-Assoziation wird von marinen Dinoflagellaten dominiert. Es konnten aber auch Süß- bzw. Brackwasser-Algenzysten von Zygnemataceae
(Ovoidites elongatus) nachgewiesen werden, die einen terrestrischen Einfluss bezeugen. Im Gegensatz zu den Pollen-Floren des Danium und des Thanetium Eurasiens stellen die ausgestorbenen Pollen-Leitformen der Normapolles im untersuchten (etwas jüngeren) Material lediglich einen geringen Anteil
der Assoziation dar.
1 aLFrÉd dULai: Hungarian Natural History Museum, Department of Palaeontology and Geology, H 1431 Budapest POB 137, Hungary. DULAI@nhmus.hu
2 Lenka hradeCká and LiLian ŠvábeniCká: Czech Geological Survey, Klárov 131/3, CZ 118 21, Praha 1, Czech Republic. lenka.hradecka@geology.cz,
lilian.svabenicka@geology.cz
3 MaGda konzaLová: Institute of Geology v.v.i., Academy of Sciences of the Czech Republic, Rozvojová 135, CZ 165 00 Praha 6, Czech Republic.
konzalova@gli.cas.cz
4 GYÖrGY LeSS: Department of Geology and Mineral Resources, University of Miskolc, H 3515 Miskolc-Egyetemváros, Hungary. foldlgy@uni-miskolc.hu
5 haraLd Lobitzer: Lindaustraße 3, A 4820 Bad Ischl, Austria. harald.lobitzer@aon.at
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Abstract
An integrated study of brachiopods, small and larger foraminifera (orthophragmines and nummulitids), calcareous nannofossils and palynomorphs / dinocysts was carried out from the marly limestones respectively sandy marls of the Ultrahelvetic zone at the locality Rote Kirche in the Gschliefgraben near
to Gmunden in Upper Austria. Microfossils (including larger foraminifera) indicate the Early Eocene, more precisely the early-middle part of the late Ypresian (the NP 11 and NP 13 calcareous nannofossils, the P 7–8 planktonic foraminiferal and the SBZ 10 shallow benthic zones). Eocene brachiopods are
described for the first time from Austria. Six species (Gryphus kickxii, Meznericsia hantkeni, Terebratulina tenuistriata, Orthothyris pectinoides, Megathiris
detruncata, Argyrotheca sabandensis?) were identified, the taxonomic composition of which (based on the dominance of Gryphus and Terebratulina) refers
to deeper water, probably outer shelf environment. These palaeoecological conditions are also confirmed by the composition of larger foraminifera. A new
orthophragminid chronosubspecies, Orbitoclypeus multiplicatus gmundenensis, characteristic for the SBZ 10 Zone, is introduced. The palyno-association is
dominated by marine dinoflagellates but freshwater-brackish algal cysts of Zygnemataceae (Ovoidites elongatus) are also present, testifying terrestrial
input. In the Cretaceous and the Palaeocene (Danian and Thanetian) pollen floras of the Eurasian Normapolles Province Normapolles pollen are a characteristic element. In the investigated association, however, Normapolles are present only in low quantity.
Location and Geological Setting
Previous Work
The Gschliefgraben area comprises a large land slide (e.g.
Koch, 1898; Baumgartner & Mostler,1978; Millahn et
al., 2008; Weidinger, 2009; Weidinger & Niesner, 2009)
SE of the town of Gmunden in Upper Austria, exposing
rocks of Jurassic, Cretaceous and Palaeogene age, which
are attributed to the Ultrahelvetic thrust unit. Due to the
mass movement and an intense tectonic overprint by a
major strike slip system (Egger et al., 2009), extended undisturbed sections do not exist. In the south, the Ultrahelvetic rocks are bordered by middle Triassic limestones of
the northern rim of the Northern Calcareous Alps. In the
north, Upper Cretaceous turbidites of the Rhenodanubian
Flyschzone show a tectonic contact to the Ultrahelvetic
unit (Text-Fig. 1).
In the early geological literature of the Salzkammergut re
gion the Gschliefgraben is mentioned repeatedly. Among
the earliest records are the papers by Joseph August
Schultes (1809) and Paul von Partsch (1826). Carl Lill
von Lilienbach (1830) was particularly surprised to find
there nummulite-bearing sediments containing green mineral grains (glauconite). Ami Boué (1832) was the first
who published a cross section through the Gschliefgraben. Finally Franz von Hauer (1858) described the complex lithologic sequence. He was also the first, who described in detail the Eocene sediments of the “Rote Kirche”
location.
The slope, on which the Gschliefgraben is situated, extends from the eastern shore of Lake Traunsee (423 m)
up to the small rock-cliff of the “Rote Kirche” (840 m),
a famous site for the occurrence of Eocene fossils. The
cliff consists mostly of yellow-orange coloured marly sandstones respectively sandy marls. On the top of
the cliff limestones with nummulites, brachiopods, bivalves and echinoderms with intercalations of grey sandy marls, respectively brittle sandstones are cropping
out (Text-Fig. 2). Glauconite is almost omnipresent, rarely also thin layers of “Bohnerz”, i.e. finely distributed
limonitic ooides.
For a long period the sequence of the Gschliefgraben was
considered being part of the Flysch zone or of the Upper
Cretaceous / Palaeogene Gosau Group of the Northern
Calcareous Alps (e.g. Fugger, 1903). However, Karl Götzinger in 1937 expressed the opinion, that from the palaeogeographic point of view this sequence is part of the
Helvetic zone. From the tectonic point of view Ernst Kraus
(1944) considered the Gschliefgraben as transgressively
overlying the Flysch zone, while for Max Richter & Gotthold Müller-Deile (1940) it represents a tectonic window
of the Helvetic zone underlying the Flysch unit.
Since 1951 the latter opinion was shared by Siegmund
Prey. Prey’s papers, published between 1949 and 1983,
improved the biostratigraphic record of the lithologically
Text-Fig. 1.
Location, regional geology and tectonic position of “Rote Kirche” in Gschliefgraben.
Sketches courtesy Hans Weidinger, Kammerhofmuseum Gmunden.
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posited on the shelf of the European Platform during the
Ypresian transgression within Zone NP 12.
Studied Material
Text-Fig. 2.
Uppermost stratigraphic sequence of the Rote Kirche section. Nummulite-limestone partly rich in ����������������������������������������������������������
“���������������������������������������������������������
Bohnerz” (limonite ooides) alternating with grey, glauconitic sandy marls.
similar, but stratigraphically diverse rocks of the (Ultra)helvetic zone. In 1953 he argued that the Eocene sandy-marly-glauconitic sediments of “Rote Kirche” show Thanetian
(Late Palaeocene) and probably also Ypresian (Early Eo
cene) age, while the top of this section is represented by a
few meters of iron oolithic nummulitic limestones of Lutetian (Middle Eocene) age, which he considered as an equivalent of the “Roterz” beds in Bavaria (Prey, 1953). This
opinion was supported later by an oral communication of
Franz Traub (in Prey, 1975).
In his last paper Prey (1983) subdivided his Ultrahelveticum into two zones, namely the Northern Ultrahelvetic and
the Southern Ultrahelvetic Klippen Zone. According to him
“Rote Kirche” is part of his Northern Ultrahelvetic, which
comprises a complex sequence of light to dark grey, partly variegated Albian to Maastrichtian marls, which are topped by Palaeocene to early Eocene glauconitic, more or
less sandy marls and nummulitic limestones. Middle Eo
cene nummulitic limestones in Adelholzen facies and Clavulinoides szaboi Beds also occur regionally.
Rasser & Piller (2001) deal in detail with facies patterns,
subsidence and sea-level changes in ferruginous and
glauconitic environments of the Palaeogene Helvetic shelf.
According to these authors the Rote Kirche exposures belong to the Southern Helvetic facies of the Austro-Bavarian Helvetic Zone, which is part of the Helvetic Shelf and
as such part of the Alpine Foreland. During the Palaeogene a peculiar shallow water carbonate sedimentation
took place on a wide carbonate platform. The sediments
are characterized by the most intensive ferruginisation and
glauconitisation known from Cenozoic shallow water carbonates of the Eastern Alps (Rasser & Piller, 2001).
According to Egger (2007), the Rote Kirche outcrop is the
easternmost exposure of the South Helvetic zone. There
the nummulitic limestone of the Frauengrube Member and
in particular the underlying marly sandstone (Prey, 1983)
yielded calcareous nannoplankton of zone NP 12. This
nannoflora is considered to indicate, that after a sea-level
rise the nummulitic marlstones and limestones were de-
Two of the authors (Harald Lobitzer and Alfréd Dulai) have
visited the locality on 29.04.2010, with the guidance of
two local private collectors, namely Ferdinand Estermann and Karl Bösendorfer from Pinsdorf. Several macromorphic brachiopod specimens were found in the field,
and four samples were collected for washing and checking micromorphic brachiopods (sample 1: upper nummulitic limestone; samples 2–3: middle glauconitic sandstone; sample 4: lower Assilina sandstone). György Less
(Miskolc) has also studied two of these washed residues
for larger foraminifers (samples 1 and 4). During the field
work two additional samples were collected from the upper part of the section (more or less identical with sample
1), for palynological (Magda Konzalová, Prague), nannofossils (Lilian Švábenická, Prague) and small foraminifera
(Lenka Hradecká, Prague) studies. Karl Bösendorfer, one
of the private collectors made it possible to use and study
his brachiopod material from Rote Kirche locality. Collection of the Kammerhofmuseum in Gmunden also contains
about a dozen brachiopod specimens from the same locality.
The newly collected brachiopods and the photographed
specimens are deposited in the collection of the Hungarian
Natural History Museum, Budapest (inventory numbers of
illustrated specimens: M 2010.477.1. – M 2010.509.1). Figured larger foraminifera ������������������������������
specimens marked by E. are deposited in the Eocene collection of the Geological Institute
of Hungary, Budapest. Samples for study of small foraminifers and calcareous nannofossils were prepared in the
Laboratory and deposited in the Collections and Material
Documentation Department of the Czech Geological Survey, Prague. The palynological preparations were made in
the Institute of Geology v.v.i., Academy of Sciences of the
Czech Republic in Prague and are also deposited there.
The samples for the palynological investigation were prepared in the Laboratory of the Czech Geological Survey,
the preparations are deposited in the Institute of Geology
v.v.i., Academy of Sciences, Prague.
Brachiopods
Brachiopods are generally rare in Eocene benthic assemblages, but they were published from several localities and
numerous papers demonstrate their wide geographical
distribution within the Western Tethys. Eocene brachio
pods are known from England to Ukraine and from Belgium to Egypt (see details of their distribution in Bitner &
Boukhary, 2009, Bitner et al., in press, Dulai, submitted).
However, until now Eocene brachiopods were unknown
from Austria. In some cases brachiopods were mentioned
in faunal lists, but no description of Eocene brachiopods
was published from Austrian localities.
Recently Dulai (submitted) studied the Late Eocene (Priabonian) micromorphic brachiopods of two boreholes of
the Upper Austrian Molasse zone (Helmberg-1 and Perwang-1). These samples, due to the solving method in
acetic acid by Kamil Zágoršek (Prague) (Zágoršek & Vávra, 2000), yielded about 2000 very small, micromorphic
brachiopods, representing 10 species of 7 genera, inclu183
ding three new species. The paper describing this fauna is submitted, but the date of appearance of the proceedings volume is uncertain (6th International Brachiopod
Congress, Melbourne, 1.–5. February, 2010).
Eocene deposits around Gmunden and their fossil contents are poorly known. Prey (1983) has listed fossils
of different groups, including also two brachiopods from
this area: Terebratula aequivalvis Schafhäutl and T. hilario
nis Meneghini. Altogether 114 macromorphic brachiopods were collected during our fieldwork representing
two species of large, smooth terebratulides: Gryphus kick
xii (Galeotti, 1837) (108 specimen) and Meznericsia hantkeni
(Meznerics, 1944) (6). Karl Bösendorfer’s private collection also contains large-sized brachiopods of the same
two species (70 G. kickxii and 5 M. hantkeni). The collection of the Kammerhofmuseum in Gmunden contains a
dozen Gryphus specimens. All of the four washed samples yielded more or less small-sized, so-called micromorphic brachiopod specimens. The richest and most
diverse fauna is from the uppermost sample, collected
from the weathered part of nummulitic limestone (sample 1), where the macromorphic brachiopods were also
collected: Terebratulina tenuistriata (Leymerie, 1846) (20), Ar
gyrotheca sabandensis? (Pajaud & Plaziat, 1972) (16), Gry
phus kickxii juv. (3), Orthothyris pectinoides (Koenen, 1894) (1)
and Megathiris detruncata (Gmelin, 1791) (1). Two samples
(sample 2 and 3) of the second outcrop (upper and lower part of a grey glauconitic sandstone) contain very
fragmentary brachiopods. Sample 2 with Terebratulina te
nuistriata (15) and Gryphus kickxii (5) and sample 3 with Te
rebratulina tenuistriata (28), Gryphus kickxii (15) and Argyrotheca
sabandensis? (2). The lowest sample from yellow Assilina
sandstone (sample 4) yielded only 2 fragments of Tere
bratulina tenuistriata.
All of the washed samples contain some other fauna elements, which are only partly studied in detail in this paper
(larger foraminifers by Gy. Less).
Sample 1: small and larger foraminifers (several), worm
tubes (several coiled and some straight), echinoderms
(several echinoid needles and crinoid stalk fragments),
bryozoans (several), decapods (some fragments).
Sample 2: small and larger foraminifers (several), echinoderms (several echinoid needles, some crinoid stalk fragments), fish teeth (few).
Sample 3: small and larger foraminifers (several), echinoderms (several echinoid needles, some crinoid stalk fragments), molluscs (few ostreid fragments), worm tubes
(few), bryozoans (few), fish and shark teeth (few).
Sample 4: small and larger foraminifers (several), echinoderms (some echinoid needles and crinoid stalk fragments), molluscs (few ostreid and pectinid fragments), corals (few fragments), bryozoans (few), and decapods (few).
Systematic Notes on Brachiopods
Phylum Brachiopoda Duméril, 1806
Subphylum Rhynchonelliformea
Brunton, Holmer & Popov, 1996
Williams,
Carlson,
Class Rhynchonellata Williams, Carlson, Brunton, Holmer & Popov, 1996
Order Terebratulida Waagen, 1883
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Superfamily Terebratuloidea Gray, 1840
Family Terebratulidae Gray, 1840
Subfamily Gryphinae Sahni, 1929
Genus Gryphus Megerle von Mühlfeld, 1811
Gryphus kickxii (Galeotti, 1837)
(Pl. 1, Figs. 1–11)
1843 Terebratula Kickxii Galeotti – Nyst, p. 335, Pl. 19, Fig. 4.
in press Gryphus kickxii (Galeotti) – Bitner et al. (p. X), Figs.
3D–I, 4, 5A, B (cum syn.).
? 2010 Carneithyris subregularis (Quenstedt) – Sulser et al. p.
261–264, Text-Figs. 3, 4, 5.
Material: 213 specimens.
Notes: G. kickxii is a medium-sized, smooth brachiopod with
rectimarginate anterior commissure and short incurved
beak. The outline is very variable: elongate oval to subpentagonal or subcircular, as demonstrated on the figures of
Pl. 1. This is the most common brachiopod of the Rote Kirche locality and it was very widespread in the whole of Europe during the Eocene (Bitner et al., in press). About 70
percent of the studied Austrian specimens belong to this
species, which has a very complex taxonomic history and
was described under different names. The two species
names mentioned by Prey (1983) from Rote Kirche (T. ae
quivalvis, T. hilarionis) are probably also synonyms of G. kickxii.
Critical revision of this species was given just recently by
Bitner et al. (in press) on the basis of an extensive Middle Eocene material from the Szőc Limestone of the Bakony
Mts., Hungary. Very similar forms were mentioned from the
Swiss and Austrian Alpine area in different names: T. kick
xii by Ooster (1863) and Moesch (1878), T. aequivalvis and T.
picta by Schafhäutl (1863) and T. subregularis by Quenstedt
(1868–1871). All of these records may also refer to G. kick
xii, but until now detailed study or revision of these faunas
/ localities is missing.
The online version of the Sulser et al. (2010) paper appeared just during the preparation of this manuscript. They
have studied Lutetian (Middle Eocene) brachiopods from
NE Switzerland. Beside some undetermined Terebratuli
na specimens, they have identified their common smooth
terebratulides as Carneithyris subregularis (Quenstedt). They
regarded T. aequivalvis Schafhäutl and T. hilarionis Davidson
as separate species, and also assigned them to the genus
Carneithyris. However, the outer morphology of the Swiss
specimens is similarly variable, than in case of Lutetian
fauna of the Bakony Mts. (Bitner et al., in press) as in
the case of the studied Rote Kirche fauna. The three assemblages seem to be overlapping in outer morphological characters and in variability. The same is true for sub
regularis / aequivalvis length / width comparisons (see Fig. 8
in Sulser et al., 2010). Sulser et al. (2010) attribute their
material to the species subregularis, because Gryphus kickxii is
“ill-defined” and its thorough revision is missing (although
they also recognized the close relationship between kickxii
and subregularis). However, a paper parallel to Sulser et al.
(2010), a recent critical revision on G. kickxii is just prepared
on a rich material from the Hungarian Middle Eocene by
Bitner et al. (in press). As this latter paper justifies the
Meznericsia hantkeni (Meznerics, 1944)
(Pl. 2, Figs. 1–2)
1944 Magellania (s.l.) Hantkeni n. sp. – Meznerics, p. 46, Pl. 3,
Figs. 13–16; Pl. 5, Figs. 21–23.
1975 Gryphus inkermanicus Zelinskaja sp. nov. – Zelinskaya,
p. 94, Pl. 8, Fig. 1.
in press Meznericsia hantkeni (Meznerics, 1944) – Bitner et
al., p. X, Figs. 5C, D, 6–8.
Material: 11 specimens.
Text-Fig. 3.
Polished surface of nummulitic limestone from Rote Kirche upper locality. The
small sample contains several macromorphic brachiopods (both double valves
and separated valves; probably Gryphus kickxii). The thin sediment infillings in
some specimens indicate the original position of the rock sample. The mostly
sparitic infilling refer to relatively quick sedimentation. Scale bar: 1 cm.
validity of the G. kickxii species, and it has priority over sub
regularis as well as over aequivalvis and hilarionis, in my opinion the Swiss Lutetian material probably also represents a
new record of G. kickxii. Concerning the generic assignment,
on the basis of the internal morphological characters and
the shell ultrastructure, the Hungarian specimens clearly
belong to the short-looped Gryphus (Bitner et al., in press).
The internal characters of the Swiss specimens seem to
be poorly preserved (at least on the basis of Fig. 5a–b in
Sulser et al., 2010). Therefore their generic assignment to
the fundamentally Cretaceous Carneithyris on the basis of
some selected sections seems to be uncertain. Supposedly, the internal morphological characters of these terebratulides are variable similar to the external ones. For
a more certain species and generic assignment of Alpian
Eocene short-looped terebratulides, we need more studies in the future, including statistical comparisons of outer
morphological characters, and serial sections of well-preserved specimens.
The very limited time to prepare this paper inhibits the
investigation of the internal morphology of the brachiopods by serial sections of the specimens at Rote Kirche.
Later on it would be useful to check the intraspecific internal variability of specimens with different outer morpho
logy. However, on the basis of the polished surface of the
nummulitic limestone (Text-Fig. 3), most of the brachiopod
specimens are infilled with sparitic calcite, therefore unfortunately the serial sectioning seems to be a little hopeless.
Notes: M. hantkeni is a large-sized, strongly biconvex,
smooth terebratulide with a massive, strongly incurved
beak and paraplicate anterior commissure. The species
was described by Meznerics (1944) as Magellania (s.l.) Hant
keni. However, on the basis of the distinctive external and
internal morphological characters, Bitner et al. (in press)
recently erected a new genus, Meznericsia for this species.
Zelinskaya (1975) described the same morphology as Gry
phus inkermanicus from the Ukraine and its smaller size prob
ably refers to a juvenile specimen. The specimens from the
Rote Kirche locality have widened the known palaeogeographical distribution of this rare species.
Distribution: Eocene of Hungary, Ukraine and Austria (see
Bitner et al., in press).
Superfamily Cancellothyridoidea Thomson, 1926
Family Cancellothyrididae Thomson, 1926
Subfamily Cancellothyridinae Thomson, 1926
Genus Terebratulina d’Orbigny, 1847
Terebratulina tenuistriata (Leymerie, 1846)
(Pl. 3, Figs. 1–11)
2000 Terebratulina tenuistriata (Leymerie) – Bitner, p. 118–120,
Figs. 2, 3, 4A–F, 5B–G (cum syn.).
in press Terebratulina tenuistriata (Leymerie, 1846) – Bitner et
al., p. X, Fig. 3A–C (cum syn.).
Material: 65 specimens.
Notes: T. tenuistriata is relatively frequent at the Rote Kirche
locality, mainly in the washed residues. This is the commonest species in the Eocene brachiopod assemblages
of the Western Tethys. Bitner (2000) gave detailed analysis of this species and its great variability during the ontogeny. Different sized Rote Kirche specimens confirm this
variability (see Pl. 3, Figs. 1–11). Adults of this species are
characterized by numerous fine ribs and an elongated oval
outline, while juveniles have only 10–12 granular ribs which
increase rapidly in number with the age of brachiopods.
Distribution: Europe: Belgium, Italy, Switzerland, Austria,
Hungary, Poland, Romania, Bulgaria, Ukraine and Turkey;
Asia: Caucasus and Kazakhstan (see details in Bitner et
al., in press).
Distribution: Europe: Great Britain, Belgium, France,
Spain, Italy, Poland, Hungary, Romania, Bulgaria, and Ukraine (see Bitner et al., in press; Dulai, submitted); Africa:
Egypt (see Bitner & Boukhary, 2009).
Family Gibbithyrididae Muir-Wood, 1965
Family Chlidonophoridae Muir-Wood, 1959
Subfamily Gibbithyridinae Muir-Wood, 1965
Subfamily Orthothyridinae Muir-Wood, 1965
Genus Meznericsia Bitner, Dulai & Galácz, 2010
Genus Orthothyris Cooper, 1955
185
Orthothyris pectinoides (Koenen, 1894)
(Pl. 2, Fig. 3)
1894 Terebratulina pectinoides Koenen – Koenen, p. 1354–
1355, Pl. 99, Figs. 8–9.
2008 Orthothyris pectinoides (Koenen) – Bitner & Dulai, p. 35,
Fig. 4.9–16 (cum syn.).
Material: 1 specimen.
Notes: This species seems to be very rare at the Rote
Kirche locality, but it is a dominant faunal element in the
recently studied nearby Helmberg and Perwang samples
(Upper Austrian Molasse Zone, Late Eocene) (Dulai, submitted). The small, subcircular specimen agrees well with
those hitherto described, however it is more similar to the
Hungarian specimens (Bitner & Dulai, 2008, Figs. 4.10,
4.14) than to the more juvenile Austrian ones. Until recently, this species was attributed to the genus Terebratulina, but
Bitner & Dieni (2005) and later Bitner & Dulai (2008) and
Dulai (submitted) attributed it to the genus Orthothyris created by Cooper (1955) for Late Cretaceous brachiopods.
On the basis of the Helmberg and Perwang materials, Dulai (submitted) recognized that Orthothyris and the very similar Terebratulina alternate with each other along the Upper
Eocene layers and probably were competitors of the same
ecological niches.
Distribution: Eocene of Germany (Koenen, 1894), Ukraine
(Zelinskaya, 1975), Italy (Bitner & Dieni, 2005), Hungary
(Bitner & Dulai, 2008) and Austria (Dulai, submitted; and
this paper).
Superfamily Megathyridoidea Dall, 1870
Family Megathyrididae Dall, 1870
Genus Megathiris d’Orbigny, 1847
Megathiris detruncata (Gmelin, 1791)
(Pl. 2, Fig. 4)
2007 Megathiris detruncata (Gmelin) – Dulai, p. 2–3, Figs. 2,
1–2 (cum syn.).
2008 Megathiris detruncata (Gmelin) – Bitner & Dulai, p. 35–
36, Figs. 5.1–4 (cum syn.).
Material: 1 specimen.
Notes: M. detruncata has very wide distribution both stratigraphically and geographically. It is one of the most common species in Palaeogene, Neogene and Recent shallow
water assemblages. However, it is rare in deeper water environments, as it is also confirmed by the Helmberg and
Perwang samples (Dulai, submitted), as well as the Rote
Kirche locality (1 known juvenile specimen only).
Distribution: Eocene: Italy, Hungary, Austria (see details in
Bitner & Dulai, 2008; Dulai, submitted); Oligocene: Hungary (Dulai, 2010); Miocene: Central Paratethys (see details in Bitner & Dulai, 2004 and Dulai, 2007); Recent:
Mediterranean, Eastern Atlantic and Caribbean Sea (Logan, 1979; Brunton & Curry, 1979; Cooper, 1977).
Genus Argyrotheca Dall, 1900
Argyrotheca sabandensis? (Pajaud & Plaziat, 1972)
(Pl. 2, Figs. 5–11)
1972 Cistellarcula sabandensis nov. sp. – Pajaud & Plaziat, p.
450–451, Text-Figs. 2–3, Pl. 1, Fig. 2.
Material: 18 specimens.
186
Notes: This small sized micromorphic species is relatively
frequent in the washed residues of the
����������������������
Rote
������������������
Kirche locality. Although all of the studied specimens are small and not
very well preserved, they show remarkable similarity with
A. sabandensis described by Pajaud & Plaziat (1972) from
the Late Palaeocene of Spain. The outline of the specimens, the shape of the beak area, the length of the hinge,
the number and character of the ribs seem to be consistent with A. sabandensis. However, some uncertainties are
caused by the very poor illustration of this species in the
original description. Based on external and internal morphological characters, genus Cistellarcula was synonymised
with Argyrotheca by Calzada & Urquiola (1994). If the identification of these specimens is correct, the known stratigraphical distribution of this species is widened by this
record from Late Palaeocene to Early Eocene.
Distribution: Late Palaeocene (Thanetian) of Spain, and
Early Eocene (Ypresian) of Austria (Gmunden).
Taxonomic Composition of the
Brachiopod Fauna
Linguliformea and Craniiformea brachiopods are missing;
all studied specimens belong to the Rhynchonelliformea
subphylum. Within Rhynchonelliformea, all specimens belong to the
���������������������������������������������������
order
�����������������������������������������������
Terebratulida (rhynchonellides and thecideides are missing). Within terebratulides the short-looped
superfamily Terebratuloidea is dominant (Gryphus 69 %,
Meznericsia 3.6 %) but Cancellothyridoidea is also significant (Terebratulina 21 %, Orthothyris 0.3 %). Two genera belonging to the superfamily Megathyridoidea are much less
numerous (Argyrotheca 5.8 %, Megathiris 0.3 %).
The above mentioned taxonomic composition is based
on all studied specimens and therefore supposedly little
biased against the micromorphic species: both the private collection and the material from ����������������
the Kammerhofmu������������
seum contain only macromorphic brachiopods (they did
not examine washed materials). However, if we check only
the new material of the upper nummulitic limestone (from
where both macromorphic and micromorphic brachiopod specimens were intensively collected), the taxonomic
composition does not change significantly: Gryphus 71 %,
Meznericsia 3.9 %, Terebratulina 13.2 %, Orthothyris 0.7 %, Argy
rotheca 10.5 % and Megathiris 0.7 %). The only significant difference is that Argyrotheca is more frequent, while Terebratulina
is less common.
Palaeoecology, Palaeoenvironment
According to Logan (1979) and Logan et al. (2004) Argy
rotheca, Megathiris and Lacazella dominate in shallow water environments (ranging down to about 200 m) of the Recent
Mediterranean, while Gryphus, Terebratulina, Platidia and Meg
erlia characterize the eurybathic species, which are more
typical of the bathyal zone. The absence of thecideids,
the limited rate of Megathyridoidea (Argyrotheca, Megathiris)
and the dominance of Gryphus and Terebratulina clearly refer
to deeper water environment at the Rote Kirche locality,
maybe in outer shelf environments, as suggested also by
larger foraminifera (see later). The distributional pattern of
the ��������������������������
Mediterranean upper bathyRecent Gryphus vitreus along ������������������������������
al continental slope was intensively studied by Emig & Arnaud (1988) and Emig (1989a, b).
Ecologically, the described brachiopods belong to three
categories. Most of the species are attached by a strong
and short pedicle to hard substrates: Gryphus, Argyrotheca,
Megathiris, Orthothyris. However, Terebratulina is attached directly to the loose sediment by a root-like, divided pedicle.
Meznericsia is an extinct genus without recent representatives, but the very convex valves, and extremely incurved
small beak may refer to non functional pedicle, at least in
the adult phase. It should mean that after the “normal”,
attached juvenile stage, the large-sized, nearly globular
adult specimens secondarily became free-living, probably
on soft substrate.
Larger brachiopods can offer hard substrate for epifaunal encrusters, but the amount of epibionts is very variable both on fossil and Recent forms. Only two out of
213 studied specimens (0.9 %) of Gryphus kickxii show
remains of worm tubes (some similar worms were also
seen on large nummulitids). Both coiled worm tubes are
situated on the ventral valve, very near to the terminal
part of the anterior margin (Pl. 1, Figs. 10–11). Taking
into consideration the life position of Gryphus, the ventral
valve and mainly the terminal part of the ventral valve
is situated at the highest point. These brachiopods are
sometimes densely packed, and only these parts of the
shells are available as solid substrate for the settlement of larvae. It suggests that they encrusted the ventral valves of Gryphus during the life of the brachiopods.
As they attached very near to the anterior margin, the
worms probably benefited from the feeding currents of
the brachiopod. Similar situations were reported e.g.
on the Palaeozoic Mucrospirifer (Schumann, 1967), on
the Devonian Anathyris (Alvarez & Taylor, 1987), on the
Eocene Paraplicirhynchia (Bitner, 1996), on the Miocene
Argyrotheca (Dulai, 2007) and on the Cenozoic and Recent Tegulorhynchia (Lee, 1980).
Small Foraminifera
Material and method
One sample from the locality Rote Kirche was collected
for foraminiferal analysis. The sample was washed in the
Laboratory of the
�����������������������������������������
Czech
�������������������������������������
Geological Survey in Prague using the standard washing method. The size of the sieve of
0.063 mm and coarse fraction was kept. The
���������������
f����������
oraminiferal assemblage was studied by a Nikon binocular microscope.
Results
The studied sample contains a relatively rich foraminiferal assemblage but the preservation of foraminiferal tests
is mostly bad. Bryozoa and Echinodermata remains were
also found, as well as some ostracods and fish teeth. In
the anorganic
�������������������������������������������������������
part of �������������������������������������
the material
���������������������������������
grains of glauconite appear.
In the foraminiferal assemblage benthic species prevail,
especially Heterolepa eocaena (Guembel). Among other benthonic foraminifers Spiroplectammina pectinata (Reuss), Globoro
talites sp., Planulina costata (Hantken), Pararotalia lithothamnica
(Uhlig) and Textularia sp. are present.
Planktonic species are less abundant. Specimens of
Truncorotalia aequa (Cushman & Renz), Subbotina triloculinoides
(Plummer), Turborotalia primitiva (Finlay) and Globorotalia aragon
ensis Nuttall were found. Some of the recognized species
were mentioned in previous papers from the Palaeogene
of the Austrian Helvetic Zone (Gohrbandt, 1963, 1967;
Wille-Janoschek, 1966).
Stratigraphic interpretation
The presence of G. aragonensis in the studied sample allows
to attribute this assemblage to the planktonic Zone P7
(Globorotalia formosa) to P8 (Globorotalia aragonensis) of the Early
Eocene according to Blow (1969) and Berggren (1971).
Palaeoecological interpretation
Abundance of benthic foraminifers and a smaller amount
of planktonic ones characterize shallow-water conditions
at certain times.
Larger Foraminifera
Introduction
Larger foraminifera occur in great quantity in two samples.
These are the uppermost nummulitic limestone (sample
1) and the lower outcrop with Assilina (sample 4). They are
represented by nummulitids (genus Nummulites and Assilina)
and orthophragmines, which is an informal collective term
for Eocene orbitoidal forms uniting two systematically independent families, such as Discocyclinidae (consisting
of genus Discocyclina and Nemkovella) and Orbitoclypeidae
(with genus Orbitoclypeus and Asterocyclina). The preservation
of fossils is average in both samples, megalospheric (A)
forms are in great majority.
Methods
The inner morphology of larger foraminifera could be
studied by opening them by the splitting method with
pliers and painting with violet ink (described in detail
in Less, 1981). In the determination of larger foraminifera the morphometric method (described in detail by
Drooger, 1993) was followed, i.e. in each sample specimens were grouped into populations, the members of
which are clearly distinguishable from the specimens of
the other populations of the same sample. Taxonomic
determinations are based on these populations (as a
whole) and not on their separate individuals. These taxa
are in most cases the members of a long-lasting and
continuous evolutionary chain called lineage or phylum.
In the case of orthophragmines lineages correspond to
species while for the genus Nummulites and Assilina they
form a series of chronospecies.
In the determination of orthophragmines we focused on
the internal features found in the equatorial section, thus
we adopted principles and nomenclature used by Less
(1987) as illustrated in Text-Fig. 4 and explained in the
header of Table 1. Numerous orthophragminid lineages
(their validity is proven biometrically by Less & Ó. Kovács,
2009) are used for biostratigraphic purposes after being
artificially segmented into chronosubspecies separated
from each other by arbitrary biometric limits of the mean
deuteroconchal size, the most rapidly evolving parameter.
A synopsis of subspecies identification based on the outer
cross-diameter of the deuteroconch (parameter d) is given
in Özcan et al. (2010). A revised stratigraphy of late Ypresian to middle Lutetian orthophragmines is presented in
Özcan et al. (2007b).
187
Representatives of nine orthophragminid lineages could
be found in the Gmunden samples. They are figured
in Pl. 4, biometric data are summarized in Table 1.
Because of the limited space, a complete statistical
evaluation with the number of specimens (№), arithmetical mean and standard error (s.e.) is given only
for parameter d, the crucial parameter in subspecific
determination. If the population consists of only a
single specimen, no subspecies is determined, in the
case of only two or three specimens, the subspecies
is determined as “cf.”. Since most orthophragmines
found in the Gmunden samples are recently discussed
in Özcan et al. (2007a, 2007b, 2010) and Less et al.
(2007), we do not repeat here their description with
the exception of Orbitoclypeus multiplicatus gmundenensis n.
ssp. (see in the systematical part), which represents
the most advanced developmental stage of the lineage
known so far.
Nummulitids appeared to be less diverse in the Gmunden
samples. Four lineages could be identified, some small
Nummulites have not been determined on the specific level. The segmentation of lineages into chronospecies by
Schaub (1981) is typological and based mainly on microspheric (B) forms, however we also could use the mean
proloculus (the first chamber) diameter of the megalospheric (A) forms in the Schaub collection measured by
Less (1998b). Nummulitids are figured in Pl. 5, biometric
data of the inner cross-diameter of the proloculus (parameter P) are summarized in Table 2. The specific determination within lineages is briefly discussed at particular
samples.
Results
Sample 1 consists of a relatively rich assemblage of larger foraminifera dominated by both orthophragmines and
nummulitids. The specific composition is as follows:
Outer cross-diameter of the embryon
Parameters
deuteroconch
Discocyclina archiaci cf. archiaci (Schlumberger) – Pl.
4, Fig. 6.
D. fortisi fortisi (d’Archiac) – Pl. 4, Figs. 1–3.
D. pulcra cf. landesica Less – Pl. 4, Fig. 5.
D. dispansa taurica Less – Pl. 4, Figs. 4, 7.
Nemkovella evae evae Less – Pl. 4, Figs. 8, 9.
N. strophiolata cf. fermonti Less – Pl. 4, Fig. 10.
Orbitoclypeidae
Orbitoclypeus schopeni crimensis Less – Pl. 4, Figs. 12–
14.
O. multiplicatus gmundenensis n. ssp. Less – Pl. 4,
Figs. 15–19.
Asterocyclina alticostata (Nuttall) indet. ssp. – Pl. 4,
Fig. 11.
Nummulitidae
Nummulites nemkovi Schaub – Pl. 5, Figs. 1–4.
N. irregularis Deshayes – Pl. 5, Figs. 6–8.
N. indet. sp. (radiate forms)
Assilina plana Schaub – Pl. 5, Figs. 9, 10.
Comments on nummulitids: According to Schaub (1981)
Nummulites nemkovi, N. irregularis and Assilina plana are members of the N. distans, N. irregularis and A. spira lineage, respectively. Specific identification within lineages is based
on the measurements by Less (1998b). Concerning the N.
distans lineage, the mean proloculus diameter (Pmean) given
in Table 2 best fits to N. nemkovi. It is considerably larger
than the characteristic values of N. haymanensis, the ancestor of N. nemkovi, and significantly smaller than those of N.
distans, the offspring. In the case of the N. irregularis lineage,
the dimension of the proloculus fits well N. irregularis and is
considerably smaller than that of N. maior, the successor.
Finally, the proloculus diameter of Assilina with an open spi-
Adauxiliary chamberlets
protoconch
d (µm)
Orthophragmines:
Discocyclinidae
width
height
Equatorial chamberlets
annuli/
0.5 mm
width
height
Subspecific
determination
N
W (µm)
H (µm)
n
w (µm)
h (µm)
Species
Sample
N°.
range
mean±s.e.
range
mean
range
range
range
range
range
range
Discocyclina
archiaci
Gmunden 1
3
415–510
462
260–295
278
25−30
40−45
65−75
8−9
35−40
70−90
cf. archiaci
Gmunden 1
18
550–910
719±26
260–440
352
38−52
40−55
50−70
8−10
35−40
65−80
fortisi
indet. ssp.
D. fortisi
p (µm)
number
Gmunden 4
1
−
−
45
60
9−14
40
70
Gmunden 1
11
165–260
214±9
110–160
129
13−21
30−35
45−60
11−15
25−30
50−70
taurica
Gmunden 4
4
160–260
205±18
90–150
122
13−20
30−35
45−55
12−15
25
45−60
taurica
D. pulcra
Gmunden 1
2
570–665
618
260
48
40−50
80−110
6−7
25
100−120
cf. landesica
Nemkovella
evae
Gmunden 1
10
205–290
246±9
105–180
153
11−15
50–60
45–60
11–13
30–40
40–60
evae
N. strophiolata
Gmunden 1
2
115–145
130
60–90
75
6−7
40
25−30
18
25−30
30−35
cf. fermonti
Orbitoclypeus
schopeni
Gmunden 1
17
295–550
418±13
150–335
235
28−40
40−50
50−60
8–10
30−40
60−100
crimensis
O. multiplica
tus
Gmunden 1
11
455–790
613±32
250–430
318
32–48
45–80
50–70
6.5–8
40–45
75–100
gmundenensis
n. ssp.
Asterocyclina
alticostata
Gmunden 1
1
185
8
60–120
60
13
30–35
35–45
indet. ssp.
D. dispansa
800
255
Table 1.
Statistical data of orthophragminid populations. No: number of specimens, s.e.: standard error.
188
ral (the basic feature of their arrangement into the A. spira
lineage) in sample Rote Kirche 1 falls between A. adrianen
sis (the ancestor) and A. laxispira (the offspring) and corresponds well to A. plana.
Age: This assemblage clearly determines the SBZ 10 Zone
by Serra-Kiel et al. (1998) and the OZ 6 Zone by Less
(1998a), indicating the early part of the late Ypresian (=
Cuisian). Moreover, the OZ 6 Zone suggests the higher
part of the SBZ 10 Zone. The correlation of orthophragminid (OZ) zones with shallow benthic (SBZ) and planktic
zonations is given in Özcan et al. (2010). Discocyclina fortisi
fortisi, Nummulites nemkovi and Assilina plana are zonal markers,
whereas the range of all the other taxa includes this zone.
Discocyclina archiaci archiaci and Orbitoclypeus multiplicatus are not
known from younger strata, moreover this latter species
in older layers is represented by O. m. kastamouensis, a more
primitive developmental stage than the newly described O.
m. gmundenensis. In the meantime Orbitoclypeus schopeni crimen
sis, Discocyclina dispansa taurica, D. pulcra, Nemkovella strophiolata,
Asterocyclina alticostata and Nummulites irregularis are unknown
from older horizons.
Facies: The richness of orthophragmines and the presence
of nummulitids with an open spiral in combination with the
lack of Nummulites with granules and porcellaneous forms
(alveolinids and genus Orbitolites) indicate the deeper part of
the photic shelf, very probably the outer ramp.
Sample 4 contains a considerably less���������������
�������������������
diverse assemblage, in which the genus Assilina dominates. Orthophragmines and the representatives of the genus Nummulites are
subordinate. The specific composition is as follows:
Orthophragmines:
Discocyclinidae
Discocyclina dispansa taurica Less
D. fortisi indet. ssp.
Nemkovella indet. sp. (only a B-form was found)
Nummulitidae:
Assilina aff. placentula (Deshayes) – Pl. 5, Figs. 5, 11,
12.
Nummulites indet. sp. (small radiate forms).
Comments on Assilina: The representatives of this genus
in sample 4 have a considerably tighter spiral than that in
sample 1. Therefore, they are ranged into the A. exponens lineage. Based on the measurements by Less (1998b), the
proloculus diameter in the given sample (see Table 2) is intermediate between A. placentula (characteristic for the Low
er Cuisian, see Serra-Kiel et al., 1998) and A. cuvillieri (oc-
Text-Fig. 4.
The measurement system of megalospheric orthophragmines in equatorial
section. See the header of Table 1 for explanation.
curring in the Upper Cuisian). Such forms are determined
by Schaub (1981) as A. aff. placentula, mainly from the Mid
dle Cuisian.
Age: Although the presence of Assilina aff. placentula suggests Middle Cuisian (SBZ 11) as discussed above, this
rather narrow time-span cannot be confirmed by other
larger foraminifera. The range of Discocyclina dispansa taurica is
SBZ 10–12 (Özcan et al., 2007b, updated by Zakrevskaya
et al., in review), i.e. the whole late Ypresian (SBZ 10–12),
which is a more cautious age-estimate for sample 4.
Facies: This sample indicates a slightly less��������������
������������������
deep environment than that of sample 1, since orthophragmines are
subordinate and Assilina aff. placentula with a tighter spiral
replaces the representatives of the A. spira lineage with a
more open spiral. Meanwhile forms, characteristic for the
middle ramp (Nummulites with granules) or for the inner ramp
(porcellaneous forms like alveolinids and the genus Orbito
lites) are still missing. To sum up: the shallower part of the
outer ramp seems to be the most realistic assumption.
Systematic Part
Order Foraminiferida Eichwald, 1830
Family Orbitoclypeidae Brönnimann, 1946
Taxon
Sample
N°
Proloculus diameter (P) in mm
Range
Mean ± s.e.
Nummulites nemkovi
Gmunden 1 15
260 – 620
482,3 ± 17,6
N. irregularis
Gmunden 1
8
150 – 350
241,9 ± 22,6
Assilina plana
Gmunden 1 17
185 – 390
321,5 ± 18,8
A. aff. placentula
Gmunden 4 15
270 – 560
350,3 ± 21,5
Table 2.
Statistical data of the inner cross-diameter of the proloculus of nummulitid
populations (in µm).
No: number of specimens, s.e.: standard error.
Genus Orbitoclypeus Silvestri, 1907
Orbitoclypeus multiplicatus (Gümbel, 1870)
Emended diagnosis: Average-sized, inflated, unribbed
forms with “marthae” type rosette. The medium-sized to
moderately large embryon is excentrilepidine, rarely eulepidine. The numerous, “varians” type adauxiliary chamberlets are rather wide and medium high as well as the
equatorial chamberlets. The annuli are usually moderately
undulated; the growth pattern is of the “varians” type. O.
multiplicatus is subdivided into four successive subspecies
as defined below:
189
O.
O.
O.
O.
m.
m.
m.
m.
haymanaensis dmean < 310 µm
multiplicatus dmean = 310–420 µm
kastamonuensis dmean = 420–550 µm
gmundenensis dmean > 550 µm.
Orbitoclypeus multiplicatus gmundenensis n. ssp. Less
Pl. 4, Figs. 15–19.
Etymology: Named after the city of Gmunden.
Holotype: Specimen E.10.31 (Pl. 4, Figs. 18, 19.).
Depository: Geological Institute of Hungary, Budapest.
Paratypes: All the other specimens from Gmunden, sample 1, illustrated in Pl. 4, Figs. 15–17.
Type locality: Gmunden (Austria), sample Rote Kirche 1.
Type level: Lower Upper Ypresian, the OZ 6 orthophragminid and the SBZ 10 shallow benthic zone.
Diagnosis: Orbitoclypeus multiplicatus populations with dmean
exceeding 550 µm.
Description (see also Table 1): Moderately large (3–5 mm),
inflated, unribbed forms with “marthae” type rosette. The
embryon is rather large, mostly excentrilepidine, some
times eulepidine. The numerous “varians” type adauxiliary
chamberlets are rather wide and relatively high. The equatorial chamberlets are also fairly wide and moderately high.
The annuli can be slightly undulated; their growth pattern
is of the “varians” type.
Remarks: Representatives of the Orbitoclypeus multiplicatus lineage are mostly known from the Thanetian and early Ypresian (Ilerdian), in the SBZ 3 to 8 and OZ 1b to 4 Zones.
Özcan et al. (2007b) reported one single specimen with similar characteristics as in Gmunden from the SBZ 10/11 or
OZ 6/7 Zones corresponding to the lower part of the Upper
Ypresian (Cuisian) of Kiriklar (N Turkey). Our material from
Gmunden consisting of eleven specimens allows us to introduce the most advanced developmental stage of the lineage as a new chronosubspecies.
Orbitoclypeus multiplicatus gmundenensis is hardly distinguishable
from O. schopeni schopeni and O. zitteli with similar embryonic size and type. Its equatorial chamberlets, however, is
slightly wider than those of the other two taxa, which have
a different stratigraphical position.
Range: Early part of the late Ypresian (Cuisian), the SBZ 10
and OZ 6 Zones. It may include the SBZ 9 and 11 as well
as the OZ 5 and 7 Zones.
Gmunden (Austria) and very probably Kiriklar (Turkey).
Calcareous Nannofossils
Method
Nannofossils were investigated in the fraction of 2–30
µm, separated by decantation following the methodology described in Svobodová et al. (2004). Simple smearslide was mounted by Canada Balsam and inspected at
a 1000× magnification, using an oil-immersion objective
on a Nikon Microphot-FXA transmitting light microscope.
Biostratigraphic data were interpreted applying the zonations of Martini (1971) and Varol (1998).
190
Results
The s���������������������
tudied f�������������
raction 2–30 �������������������������
µm (samples A and B) contained predominantly anorganic material. The nannofossil
abundance in sample A was generally 10–20 specimens
per 1 field of view of the microscope, whereas sample
B was extremely poor, only 1–3 specimens per 1 field of
view of the microscope. Calcareous nannofossils were
poorly preserved in both samples. Discoasterids and large
placoliths were mostly fragmented and discoasterids and
the central fields of placoliths partly etched, partly overgrown with calcite. Some specimens cannot be identified
due to the poor preservation especially in sample Rote
Kirche B.
Sample A
The nannofossil assemblage is characterized by a higher
number of discoasterids exclusively of rosette shape, and
by the rare presence of specimens of the genera Reticulofe
nestra, Helicosphaera and Lophodolithus (Pl. 6).
The following species have been found: Coccolithus pela
gicus, C. eopelagicus, Sphenolithus radians, S. moriformis, S. edi
tus, Campylosphaera dela, C. eodela, Helicosphaera seminulum, H. lo
phota, Neococcolithes protenus, N. protenus-dubius, Cyclococcolithus
(Ericsonia) formosus, Zygrhablithus bijugatus, Calcidiscus protoannulus,
Micrantholithus flos, Pontosphaera pulcheroides, P. pulchra, Thoracos
phaera sp., Discoaster barbadiensis, D. lodoensis (7 rays, mostly
in fragments), D. kuepperi, D. sp., Toweius rotundus, T. crassus,
Girgisia gammation, Clausicoccus fenestratus, Chiasmolitus solitus, C.
eograndis (fragments), C. consuetus, C. sp., Lophodolithus mochlo
porus, L. nascens, Braarudosphaera turbinea (probably reworked
from the older sediments of the lowermost Palaeocene,
Danian age).
Sample B
Poor nannofossils are characterized by a higher number
of specimens of the genus Toweius. The assemblage consists of species Coccolithus pelagicus, C. eopelagicus, rare Ellipsoli
thus macellus, Chiasmolithus solitus, C. bidens, C. eograndis, Discoaster
binodosus, D. barbadiensis, D. kuepperi, D. multiradiatus, Zygrhablithus
bijugatus, Neochiastozygus junctus, Lophodolithus nascens, Sphenolithus
moriformis, Campylosphaera eodela, Pontosphaera pulchra, Coronocyc
lus sp., rare pentaliths of Braarudosphaera bigelowii bigelowii, B.
bigelowii parvula and Micrantholithus sp., Clausicoccus fenestratus,
Toweius crassus, T. rotundus, T. pertusus.
The assemblage also contained reworked species from older sediments of the lower and middle Palaeocene age,
such as Fasciculithus cf. ulii, Cruciplacolithus tenuis, Sullivania danica
and Markalius astroporus (Danian).
Stratigraphic interpretation
Sample A: Upper part of Lower Eocene (Ypresian), zone
NP 13 sensu Martini (1971) according to the presence of
Discoaster lodoensis (7 rays), rare Lophodolithus mochloporus and
Reticulofenestra dictyoda.
Sample B: Lower Eocene (Ypresian), the uppermost part
of zone NP 11 (Martini, 1971), i.e. NNTe1D (sensu Varol,
1998) according to the joint presence of Discoaster kuepperi
and Ellipsolithus macellus, and the relative abundance of To
weius spp.
Palaeoecologic interpretation
The presence of calcareous nannofossils indicates a sea
of average salinity, with an abundance of discoasterids,
relatively warm waters, the presence of the genera Pont
osphaera, Helicosphaera and penthaliths shallow-water conditions; the etching of placoliths and discoasterids may be
the result of carbonate dissolution caused by the release
of carbon dioxide during the oxidation of organic matter
(Švábenická et al., 2010).
Discussion
Calcareous nannofossils of the Rote Kirche outcrop have
already been studied by Egger et al. (2009). They mentioned an assemblage of zone NP 12 with Discoaster lodoen
sis and Tribrachiatus orthostylus (Type B). Sample Rote Kirche
A of the present study provided nannofossils of zone NP
13 with the genus Reticulofenestra. This small difference in results might be caused by taking samples from dissimilar
places of outcrop.
Varol (1998) mentioned the first occurrence of the genus
Reticulofenestra within zone NNTe5 and correlated it with the
uppermost part of the standard nannoplankton zone NP
12, i.e. with the upper part of the Lower Eocene. The first
occurrence of Lophodolithus mochloporus is stated by PerchNielsen (1985) within NP 13.
Joint occurrence of Discoaster kuepperi and Ellipsolithus macel
lus in sample B delimits the short stratigraphic range within zone NNTe1D (Varol, 1998). This is supported also by
the occurrence of Discoaster multiradiatus, its last occurrence
known from NP 11 (Perch-Nielsen, 1985).
The nannofossil content and stratigraphic interpretation of
samples published by Egger et al. (2009) different from
this study (samples A and B) may indicate a deposition in a
longer period of time, spanning an interval from NP 11 (upper part) up to NP 13.
Microflora – Preliminary Results
Taxonomically varied microfossils of dinocysts, spores,
pollen, remains of foram linings, tiny cuticles and xylitic
splinters were obtained from the aleuritic sample derived
from the section at Rote Kirche. Marine dinocysts, lack of
typical Mesozoic cheirolepidaceous conifers and rare Normapolles characterize the assemblage. Reworked specimens from the Upper Cretaceous, composition of pollen
taxa (Icacinaceae, cf. Sapotaceae) and comparable DinoZones point to the early Palaeogene. Observable organic
matter originated rather in a near-shore than a far offshore
environment.
Characteristic of the assemblage
Residues obtained by solution and maceration of the sample (Laboratory of the Geological Survey, Prague; geology and location of the sample site, Egger, 1996, 2007)
contained no rich assemblage of palynomorphs. They are
mainly composed of dinocyst microplankton (Table 3), with
co-occurrence of foraminiferal linings and accessories of
terrestrial plants, spores, pollen and other organic debris
(Table 4).
F e r n spores belong to the Osmundaceae, Schizaeaceae,
Lygodiaceae, Gleicheniaceae and document presence of
the terrestrial flora of the nearby coastland area.
C o n i f e r s are represented by at least two groups. The
first is documented by inaperturate pollen, resembling taxodiaceous pollen, the second comprises pollen provided
with a bisaccate apparatus (bladders), grouped within Pinaceae. Both are commonly known from the Cretaceous
and Tertiary pollen assemblages.
Characteristic feature of the present microflora is a small
number of coniferous pollen. Cheirolepidaceae pollen
grains, common in the Cretaceous deposits, were not recorded. This could be in good accordance with their disappearing in the Palaeogene.
F l o w e r i n g - p l a n t pollen genera (Normapolles and
other angiospermous pollen) were represented by solitary
species and single records (Table 4), in contrast to the non
marine environments (e.g. Menat, Borna, Geiseltal a.o.).
D i n o c y s t s dominated in the assemblage, pointing together with other organic remains to the ample nutrient
supply. Some of dinocysts show poor preservation (broken
cysts or only partly preserved specimens). These features
may be interpreted as the result of reworking and/or transport on the shelf.
Striking is relatively abundant dark organic matter, amorphous or with preserved structure.
According to the residual phyto- and microzoo-remains,
the flourishing associations can be considered in the time
of silty clay deposition.
Conclusions
The pollen and several dinocyst records provided data
for the preliminary evaluation of the relative age of the
investigated assemblage (Chateauneuf, 1980; Kedves,
1969, 1970; Kedves & Russel, 1982; Krutzsch, 2004;
Krutzsch & Vanhoorne, 1977; Davey et al., 1966;
Köthe, 1990; Lentin & Williams, 1993; Stanley, 1965
ex Williams et al., 1998); based on the dinocysts and
several flowering plant taxa, it is obviously the Palaeogene age, mostly the Palaeocene (Zone D 3) and Early
Eocene (Zone D 4, D 5, D 6). The Early Eocene (Ypresian) age has also been supported by nannofossil zones
NP 11 with Discoaster kuepperi and Ellipsolithus macellus and
NP 13 with Discoaster lodoensis and Reticulofenestra dictyoda,
as well as larger benthic foraminifera indicating the SBZ 10
Zone. Dinocysts, preliminarily recorded, show their range often within the Zone D 4, D 5, in comparison with
the palynological investigation of the borehole sections in NW Germany, Lower Saxony area (Köthe, 1990).
The fragmentary remains of some plankton specimens
and taxa predominantly known from Cretaceous deposits (Ilyina et al., 1994; Marheinecke, 1986) are considered as results of reworking and bioturbation processes, possible also at a very short time scale and
within thin layers.
The observation of amorphous organic matter and evidently organic remains allows to consider rather bay or
near shore sedimentation, not a far offshore environment.
Calcareous nannofossils indicate a warmer sea of normal
salinity. The presence of organic remains in the depositional area is supported also by the mode of nannofossil
preservation: carbonate dissolution of coccoliths is usually
caused by the release of carbon dioxide during oxidation
of organic matter.
191
Dinoflagellates
Remarks
cf. Adnatosphaeridium vittatum Williams
& Downie 1966
partly preserved Early Eocene
cf. Achomosphaera aff. triangulata
(Gerlach 1961) Davey & Williams
1969
cf. Apteodinium sp.
Age
D Zones and
Subzones
D 6b
Early Eocene
Late Palaeocene,
Early Eocene,
Late Eocene (LO?)
Köthe (Kö), 1990, NW Germany,
Gartow, Early Eocene (E Eo)
Kö, 1990,
Gartow, E Eo
partly preserved Early Eocene
Cordosphaeridium sp. – compared with
C. fibrospinosum Davey & Williams
1966 and C. trompetum (Cookson &
Eisenack 1982) Lentin & Williams
1985
Pl. 7, Figs. 1, 2 in present paper
References
Kö, 1990, Gartow, E Eo
D 4, D 5b
Kö, 1990, D 4 Late Palaeocene,
Gartow, D 5b E Eo; (D 7 a. D 8
Late Eo, more in Kö, 1990)
D3
Palaeocene
Palaeocene, bore Söhlingen,
ibid.
D 5b
Early Eocene,
whole Palaeocene D 4 D 4, 4na, nb
Kö, 1990; D 5b E Eo, Gartow
D 4 Palaeocene,
bore Penningsehl, Kö, 1990
[now Areoligera (Achomosphaera) danica]
Pl. 7, Fig. 6 in present paper
Upper Cretaceous,
particularly in
L./U. Maastrichtian
Cretaceous, Marheinecke, 1986
Dipsilidinium pastielsii (Davey &
Williams 1966) Bujak, Downie,
Eaton, Williams 1980
Early Eocene
Areoligera senonensis Lejeune-Carpen1938 sensu Köthe 1990
tier
probably
reworked
D 5b
Kö, 1990, Gartow
cf. Odontochitina sp.
partly preserved Palaeocene
D 4a (rare)
Kö, 1990
? Ceratiopsis sp.
partly preserved Palaeocene (e.g.)
D4
Kö, 1990
Isabelidinium sp. (former Chatangiella
Vozzhenikova 1967)
probably
reworked
Upper Cretaceous
(Campanian, Maastrichtian);
Palaeocene,
Early Eocene
(another type, with
broad cingulum)
Isabelidinium cf. cooksoniae (Alberti
1959) Lentin & Williams 1977
reworked
Cretaceous
Coniacian,
Campanian
Late Cretaceous, Coniacian,
Campanian, e.g. Siberia, Ilyina
et al., 1994
cf. Homotryblium aff. tenuispinosum
Davey et al. 1966
Eocene
London Clay
Davey et al., 1966
Thalassiphora cf. pelagica (Eisenack
1954) Eisenack & Gocht 1960,
T. delicata Davey et al. 1966
Cretaceous, Palaeogene, Neogene
London Clay
Kö, 1990, e.g. Early Eocene,
NW Germany, Gartow, Davey et
al., 1966, Eocene
Cretaceous,
Tertiary, Pleistocene
freshwater
and brackish
water
genus commonly known from
the basinal deposits
? Fromea sp.
e.g. Canada, Siberia, Ilyina et
al., 1994;
Chatangiella ?, Palaeocene, South
Dakota, Stanley, 1965
(ex Williams et al., 1998);
Isabelidinium sp., Early Eocene,
NW Germany, Kö, 1990
vermiculate
surface
Chlorophyta – Zygnemataceae,
freshwater green algae
Ovoidites elongatus (Hunger 1952)
Krutzsch 1959
Table 3.
Plankton (selected taxa).
192
Filicinae – ferns
Leiotriletes adriennis (Potonié & Gelletich 1933)
Krutzsch 1959
Leiotriletes microadriennis Krutzsch 1959
Mesozoic, Tertiary
Schizaeaceae, Lygodium type
Appendicisporites cf. auritus Aggassie in Singh 1983
Schizaeaceae, Lygodium type
Palaeocene (e.g. Menat), Eocene
(Geiseltal, Messel) and other sites of
Tertiary deposits
Mesozoic, Palaeogene (predomiGleicheniaceae
nantly)
Mesozoic, reworked
Schizaeaceae
Cicatricosisporites sp.
Mesozoic, Palaeogene
Schizaeaceae
Trilites menatensis Kedves 1982
Palaeocene, Eocene
Lygodiaceae (after Kedves in Kedves &
Russel, 1982)
Rugulatisporites quintus Pflug 1953
Mesozoic, Tertiary
Osmundaceae
?Palaeocene, Eocene, Neogene
Pinaceous conifers
Cretaceous, Tertiary
Pinaceae
Cretaceous, Tertiary
Taxodiaceous pollen – commonly known
from Cretaceous and Tertiary
Cretaceous, Palaeogene
extinct
Upper Cretaceous, Palaeocene, Early
Eocene common
Minorpollis sp.
Cretaceous, Palaeogene
extinct
cf. Complexiopollis vancampoe Diniz et al. 1974,
smaller-sized form
Cretaceous
reworked
Portugal, Upper Cretaceous, L. to M.
Turonian is considered
Gleicheniidites sp.
Conifers
Pityosporites sp. – Pityosporites minutus (Zaklinskaja
1957) Nagy 1985, ?Pityosporites strobipites (Woodehouse 1933) Krutzsch 1971
Pityosporites sp.
Pl. 7, Fig. 3 in present paper
Inaperturopollenites Thomson & Pflug 1953
Inaperturopollenites hiatus (Potonié 1931) Thomson
& Pflug 1953 (as Taxodiaceaepollenites sp.) in
Mesozoic
Angiospermae - Flowering plants
Normapolles
Angiospermae - Flowering plants
cf. Triporopollenites robustus Pflug 1953 subfsp.
minor Kedves 1970
cf. Betulaceae (after Kedves, 1970)
cf. Compositoipollenites sp.
Palaeogene
cf. Icacinaceae
aff. Intratriporopollenites sp.
Palaeogene, Neogene
cf. Malvaceae, Tilioideae
Tricolpites, Tricolpopollenites – Tricolpo(roi)pollenites group
– reticulate morphotypes s.l.
Tricolporopollenites exactus (Potonié 1931) Thomson
& Pflug 1953
Tricolporopollenites cf. gracillimus Krutzsch & Vanhoorne 1977
Early Cretaceous, Tertiary
Hamamelidaceae, Platanaceae, partly
extinct
Fagaceae, Castaneoideae
aff. Tricolporopollenites globus Deák 1960
Eocene (Hungary)
Palaeogene, Neogene
Palaeogene,
Epinois „Bild“ sensu Krutzsch
Pl. 7, Fig. 4 in present paper
Tetracolporopollenites sp.
Pl. 7, Fig. 5 in present paper
Late Landenian, palynozone 11 after
Krutzsch (in Krutzsch & Vanhoorne,
1977), Early Eocene
Incertae sedis; Sapotaceae
(after Kedves, 1969)
Incertae sedis, ?Sapotaceae
Other plant remains
filamentous Algae or Cyanobacteria
Precambrian – Recent,
environmentally controlled
charcoal splinters (rare)
tiny cuticle fragments
Remains of zoo-plankton
different linings of microforaminifera
Animal cuticle / epidermis remains
Table 4.
Vascular plants
193
Plate 1
Fig. 1:
Fig. 2:
Fig. 3:
Fig. 4:
Fig. 5:
Fig. 6:
Fig. 7:
Fig. 8:
Fig. 9:
Fig. 10:
Fig. 11:
194
Gryphus kickxii (Galeotti, 1837).
a – dorsal view, b – lateral view, c – anterior view.
Rote Kirche 1; L: 11.4 mm, W: 10.8 mm, Th: 5.5 mm.
M 2010.477.1., 2×.
Gryphus kickxii (Galeotti, 1837).
Dorsal view.
Rote Kirche 1; L: 12.4 mm, W: 9.7 mm, Th: 6.3 mm.
M 2010.478.1., 2×.
Gryphus kickxii (Galeotti, 1837).
a – dorsal view, b – anterior view.
Rote Kirche 1; L: 18.8 mm, W: 17.3 mm, Th: 10.0 mm.
M 2010.479.1., 2×.
Gryphus kickxii (Galeotti, 1837).
a – dorsal view, b – lateral view, c – anterior view.
Rote Kirche 1; L: 19.5 mm, W: 20.5 mm, Th: 12.1 mm.
M 2010.480.1., 2×.
Gryphus kickxii (Galeotti, 1837).
Dorsal view.
Rote Kirche 1; L: 19.2 mm, W: 16.1 mm, Th: 10.1 mm.
M 2010.481.1., 2×.
Gryphus kickxii (Galeotti, 1837).
a – dorsal view, b – lateral view, c – anterior view.
Rote Kirche 1; L: 24.4 mm, W: 24.5 mm, Th: 12.9 mm.
M 2010.482.1., 2×.
Gryphus kickxii (Galeotti, 1837).
Dorsal view.
Rote Kirche 1; L: 25.8 mm, W: 20.9 mm, Th: 11.8 mm.
M 2010.483.1., 2×.
Gryphus kickxii (Galeotti, 1837).
a – dorsal view, b – lateral view.
Rote Kirche 1; L: 28.5, W: 24.0 mm, Th: 14.6 mm.
M 2010.484.1., 2×.
Gryphus kickxii (Galeotti, 1837).
Dorsal view.
Rote Kirche 1; L: 27.7 mm, W: 31.2 mm, Th: 14.0 mm.
M 2010.485.1., 2×.
Tube worm on Gryphus kickxii (Galeotti, 1837).
Ventral view.
Rote Kirche 1; L: 21.8 mm, W: 23.6 mm, Th: 12.0 mm.
M 2010.486.1., 2×.
Tube worm on Gryphus kickxii (Galeotti, 1837).
Ventral view.
Rote Kirche 1; L: 21.2 mm, W: 19.3 mm, Th: 10.6 mm.
M 2010.487.1., 2×.
195
Plate 2
Fig. 1:
Fig. 2:
Fig. 3:
Fig. 4:
Fig. 5:
Fig. 6:
Fig. 7:
Fig. 8:
Fig. 9:
Fig. 10:
Fig. 11:
196
Meznericsia hantkeni (Meznerics, 1944).
a – dorsal view, b – lateral view, c – anterior view.
Rote Kirche 1; L: 30.8 mm, W: 27.8 mm, Th: 18.2 mm.
M 2010.488.1., 2×.
Meznericsia hantkeni (Meznerics, 1944).
a – dorsal view, b – lateral view, c – posterior view.
Rote Kirche 1; L: 29.1 mm, W: 25.6 mm, Th: 19.0 mm.
M 2010.489.1., 2×.
Orthothyris pectinoides (Koenen, 1894).
Dorsal view.
Rote Kirche 1; L: 2.7 mm, W: 2.6 mm.
M 2010.490.1., 20×.
Megathiris detruncata (Gmelin, 1791).
Dorsal view.
Rote Kirche 1; L: 1.8 mm, W: 2.1 mm.
M 2010.491.1., 20×.
Argyrotheca sabandensis? (Pajaud & Plaziat, 1972).
Dorsal view.
Rote Kirche 1; L: 2.4 mm, W: 2.0 mm.
M 2010.492.1., 20×.
Argyrotheca sabandensis? (Pajaud & Plaziat, 1972).
Dorsal view.
Rote Kirche 1; L: 2.3 mm, W: 2.0 mm.
M 2010.493.1., 20×.
Argyrotheca sabandensis? (Pajaud & Plaziat, 1972).
Dorsal view.
Rote Kirche 1; L: 1.6 mm, W: 1.5 mm.
M 2010.494.1., 20×.
Argyrotheca sabandensis? (Pajaud & Plaziat, 1972).
Lateral view.
Rote Kirche 1; L: 2.0 mm, Th: 1.0 mm.
M 2010.495.1., 20×.
Argyrotheca sabandensis? (Pajaud & Plaziat, 1972).
Oblique lateral view.
Rote Kirche 1; L: 2.6 mm, Th: 1.4 mm.
M 2010.496.1., 20×.
Argyrotheca sabandensis? (Pajaud & Plaziat, 1972).
Ventral view.
Rote Kirche 1; L: 2.4 mm, W: 2.2 mm.
M 2010.497.1., 20×.
Argyrotheca sabandensis? (Pajaud & Plaziat, 1972).
Ventral view.
Rote Kirche 1; L: 2.0 mm, W: 1.8 mm.
M 2010.498.1., 20×.
197
Plate 3
Fig. 1:
Fig. 2:
Fig. 3:
Fig. 4:
Fig. 5:
Fig. 6:
Fig. 7:
Fig. 8:
Fig. 9:
Fig. 10:
Fig. 11:
198
Terebratulina tenuistriata (Leymerie, 1846).
Dorsal view.
Rote Kirche 1; L: 2.1 mm, W: 1.5 mm.
M 2010.499.1., 20×.
Terebratulina tenuistriata (Leymerie, 1846).
Dorsal view.
Rote Kirche 1; L: 3.1 mm, W: 2.6 mm.
M 2010.500.1., 20×.
Terebratulina tenuistriata (Leymerie, 1846).
Dorsal view.
Rote Kirche 1; L: 2.5 mm, W: 1.8 mm.
M 2010.501.1., 20×.
Terebratulina tenuistriata (Leymerie, 1846).
Ventral view.
Rote Kirche 1; L: 2.6 mm, W: 1.9 mm.
M 2010.502.1., 20×.
Terebratulina tenuistriata (Leymerie, 1846).
Ventral view.
Rote Kirche 1; L: 2.5 mm, W: 2.2 mm.
M 2010.503.1., 20×.
Terebratulina tenuistriata (Leymerie, 1846).
Lateral view.
Rote Kirche 1; L: 2.6 mm, Th: 1.1 mm.
M 2010.504.1., 20×.
Terebratulina tenuistriata (Leymerie, 1846).
Oblique lateral view.
Rote Kirche 1; L: 2.7 mm, W: 1.3 mm.
M 2010.505.1., 20×.
Terebratulina tenuistriata (Leymerie, 1846).
Ventral view.
Rote Kirche 1; L: 3.9 mm, W: 2.9 mm.
M 2010.506.1., 15×.
Terebratulina tenuistriata (Leymerie, 1846).
Dorsal view.
Rote Kirche 1; L: 5.5 mm, W: 4.6 mm.
M 2010.507.1., 15×.
Terebratulina tenuistriata (Leymerie, 1846).
Dorsal view.
Rote Kirche 1; L: 5.2 mm, W: 3.9 mm.
M 2010.508.1., 15×.
Terebratulina tenuistriata (Leymerie, 1846).
Dorsal view.
Rote Kirche 3; L: 9.1 mm, W: 7.5 mm.
M 2010.509.1., 15×.
199
Plate 4
Megalospheric orthopragmines (A-forms) from Gmunden, Gschliefgraben, sample Rote Kirche 1.
Figs. 1–3:
Figs. 4, 7:
Fig.
5:
Fig.
6:
Figs.
8, 9:
Fig.
10:
Fig.
11:
Figs. 12–14:
Figs. 15–19:
Figs.
Fig.
200
Discocyclina fortisi fortisi (d’Archiac)
Fig. 1: E.10.16.
Fig. 2: E.10.17.
Fig. 3: E.10.18.
Discocyclina dispansa taurica Less.
Fig. 4: E.10.20.
Fig. 7: E.10.21.
Discocyclina pulcra cf. landesica Less.
E.10.05.
Discocyclina archiaci cf. archiaci (Schlumberger).
E.10.19.
Nemkovella evae evae Less.
Fig. 8: E.10.22.
Fig. 9: E.10.23.
Nemkovella strophiolata cf. fermonti Less.
E.10.32.
Asterocyclina alticostata (Nuttall) indet. ssp.
E.10.24.
Orbitoclypeus schopeni crimensis Less.
Fig. 12: E.10.26.
Fig. 13: E.10.27.
Fig. 14: E.10.25.
Orbitoclypeus multiplicatus gmundenensis n. ssp. Less.
Fig. 15: E.10.28.
Fig. 16: E.10.29.
Fig. 17: E.10.30.
Figs. 18, 19: holotype, E.10.31.
1–18: Equatorial sections, 40×.
19: External view, 25×.
201
Plate 5
Nummulitids from Gmunden, Gschliefgraben.
Figs.
1–4:
Figs. 5, 11, 12:
Figs.
6–8:
Figs.
9, 10:
Nummulites nemkovi Schaub.
sample Rote Kirche 1.
Figs. 1, 2: E.10.06.
Fig. 3: E.10.07.
Fig. 4: E.10.08.
Assilina aff. placentula (Deshayes).
sample Rote Kirche 4.
Fig. 5: E.10.11.
Fig. 11: E.10.12.
Fig. 12: E.10.13.
Nummulites irregularis Deshayes.
sample Rote Kirche 1.
Figs. 6, 7: E.10.09.
Fig. 8: E.10.10.
Assilina plana Schaub.
sample Rote Kirche 1.
Fig. 9: E.10.14.
Fig. 10: E.10.15.
Fig.
Figs.
B-form, 5×, all the others are A-forms, 10×.
External views, all the others are equatorial sections.
202
5:
1, 5, 6:
203
Plate 6
Calcareous nannofossils, samples Rote Kirche A, B.
PPL – plane-polarized light, XPL – cross-polarized light. For magnification see Fig. 1.
Fig.
1:
Fig.
2:
Fig.
3:
Fig.
4:
Fig.
5:
Fig.
6:
Fig.
7:
Fig.
8:
Fig.
9:
Fig.
10:
Fig.
11:
Fig.
12:
Fig.
13:
Fig.
14:
Fig.
15:
Fig.
16:
Figs. 17, 18:
Fig.
19:
Fig.
20:
Fig.
21:
Fig.
22:
Fig.
23:
Fig.
24:
Figs. 25, 26:
Figs. 27, 28:
Fig.
29:
Fig.
30:
204
Braarudosphaera turbinea Stradner.
Sample A, XPL.
Markalius astroporus (Stradner) Hay & Mohler.
Sample B, XPL.
Girgisia gammation Bramlette & Sullivan.
Sample A, XPL.
Toweius crassus (Bramlette & Sullivan) Perch-Nielsen.
Sample B, XPL.
Toweius rotundus Perch-Nielsen.
Sample A, XPL.
Clausicoccus fenestratus (Deflandre & Fert) Prins.
Sample A, XPL.
Ellipsolithus macellus (Bramlette & Sullivan) Sullivan.
Sample B, XPL.
Lophodolithus nascens Bramlette & Sullivan.
Sample A, XPL.
Lophodolithus mochloporus Deflandre.
Sample A, XPL.
Helicosphaera seminulum Bramlette & Sullivan.
Sample A, XPL.
Helicosphaera lophota Bramlette & Sullivan.
Sample A, XPL.
Calcidiscus protoannulus (Gartner) Loeblich & Tappan.
Sample A, XPL.
Discoaster multiradiatus Bramlette & Riedel.
Sample B, PPL.
Discoaster binodosus Martini.
Sample B, PPL.
Discoaster barbadiensis Tan.
Sample A, PPL.
Discoaster sp.
Sample A, PPL.
Discoaster kuepperi Stradner.
Sample B, PPL.
Fig. 17: high focus.
Fig. 18: low focus.
Discoaster lodoensis Bramlette & Riedel.
Sample A, PPL.
Chiasmolithus bidens (Bramlette & Sullivan) Hay & Mohler.
Sample B, XPL.
Chiasmolithus solitus (Bramlette & Sullivan) Locker.
Sample A, XPL.
Chiasmolithus sp.
Sample A, XPL.
Reticulofenestra dictyoda (Deflandre) Stradner.
Sample A, XPL.
Reticulofenestra sp. cf. R. dictyoda (Deflandre) Stradner.
Sample A, XPL.
Sphenolithus moriformis (Brönnimann & Stradner), Bramlette & Wilcoxon.
Sample A, XPL, 25–0o, 26–45o.
Sphenolithus radians Deflandre.
Sample A, XPL, 27–0o, 28–45o.
Rhabdosphaera sp.
Sample A, XPL.
Zygrhablithus bijugatus (Deflandre) Deflandre.
Sample A, XPL.
205
Plate 7
Figs. 1, 2:
Fig.
3:
Fig.
4:
Fig.
5:
Fig.
6:
206
Cordosphaeridium sp.
Dinocyst, one specimen at two optical levels.
Size 110 µm. Light microscope photo.
Pityosporites sp.
Pollen of Pinaceae.
Size 90 µm. Light microscope photo.
aff. Tricolporopollenites globus Deák 1960.
Angiospermous pollen, incertae sedis vel Sapotaceae.
Size 30 µm. Light microscope photo.
Tetracolporopollenites sp.
Angiospermous pollen, incertae sedis vel ?Sapotaceae.
Size 44 µm. Light microscope photo.
Remains of dinocyst.
?Areoligera (Achomosphaera) danica type. Probably reworked.
Size of the remains 75 µm. SEM micrograph.
207
Acknowledgements
We are grateful to private collectors (Ferdinand Estermann and Karl Bösendorfer) for showing us the locality
and make it possible to study their brachiopod collection.
Best thanks to Hans Egger from the Austrian Geological
Survey for critical reading of parts of the manuscript. We
thank Hans Weidinger (Gmunden) for a critical review of
the introduction and Text-Figure 1. We are also indebted
to Hans for the possibility to study the brachiopods from
the Kammerhofmuseum in Gmunden. A. Dulai and Gy.
Less were supported by the Hungarian Scientific Research
Fund (OTKA K 77451 and 60645, respectively). Small Foraminifera and calcareous nannofossil investigations have
been made in the frame of the Research Goal of the Czech
Geological Survey MZP 0002579801. Eszter Hankó (Budapest) took the macroscopic brachiopod photos. The
SEM micrographs were taken in the SEM Laboratory of the
Hungarian Natural History Museum, Budapest (Hitachi S2600N). The polished surface of the nummulitic limestone
was prepared by Péter Gulyás (Budapest). �������������
Magda �������
Konzalová expresses her thanks to Dr. H. Lobitzer (Bad Ischl) and
Dr. M. Svobodová CSc. (Prague) for the kind providing of
the samples and preparations from the locality. The SEM
micrographs were taken in the SEM Laboratory of the IG
AS CR by Dr. Z. Korbelová CSc. The study of plant microfossils was supported by Project No. AVOZ 301305 16 of
the Institute of Geology AS CR, v.v.i. Cz.
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Received: 17. September 2010, Accepted: 12. October 2010
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Lenka Hradecka