GEOLOGICA CARPATHICA
, FEBRUARY 2018, 69, 1, 30–50
doi: 10.1515/geoca-2018-0003
www.geologicacarpathica.com
Life in the fluvial hinterland of the late Sarmatian Sea
(middle Miocene): a rare terrestrial fossil site
in the Styrian Basin (Austria)
MONIKA DOUBRAWA
1
, MARTIN GROSS
2
and MATHIAS HARZHAUSER
3
1
Institute of Earth Sciences, Heidelberg University, Im Neuenheimer Feld 234, D-69120 Heidelberg, Germany; doubrawa@stud.uni-heidelberg.de
2
Universalmuseum Joanneum, Weinzöttlstrasse 16, A-8045 Graz, Austria; martin.gross@museum-joanneum.at
3
Natural History Museum Vienna, Burgring 7, A-1010 Vienna, Austria; mathias.harzhauser@nhm-wien.ac.at
(Manuscript received April 11, 2017; accepted in revised form December 12, 2017)
Abstract: This paper describes the section and fossil content of a former gravel pit in the Eastern Styrian Basin
(SE Austria), which exposes sediments of a fluvial system, ranging from within channel to overbank environments.
A predominately terrestrial gastropod fauna of 15 species so far, was recovered from a palaeosol formed in a moist and
vegetated, floodplain or abandoned channel. Up-section, a shallow freshwater pond/lake developed within the floodplain,
settled by fishes, molluscs and ostracods. By integrating regional geological and biostratigraphical data derived from
the terrestrial gastropod fauna as well as from the other recovered biota, these strata are of late middle Miocene (late
Sarmatian s.str.) age. Hence, this fossil site provides a rare insight into the terrestrial habitats in the hinterland of
the Sarmatian Sea and their biota, which are otherwise barely known in Central Europe.
Keywords: Middle Miocene, Sarmatian s.str, gastropoda, Styria/Austria, terrestrial environments.
Introduction
Outcrops, exposing terrestrial sediments of late Sarmatian
(late Serravallian) age are rare in Austria and the Central
Paratethys as well due to a major hiatus at the middle–late
Miocene transition (e.g., Kováč et al. 2004; Schreilechner
& Sachsenhofer 2007; Harzhauser et al. 2008; Gross et al.
2011a; Stworzewicz et al. 2013; Neubauer et al. 2015a,
2017; Fig. 1). While the marine system of the Sarmatian
Sea is well investigated and a lot of data have been
gathered about palaeobiological and geological changes
(e.g., Papp et al. 1974a; Harzhauser & Piller 2004;
Piller & Harzhauser 2005; Piller et al. 2007), information
on the surrounding continental environments is still
fragmentary.
An abandoned gravel pit (Badenbrunn), situated in the
Eastern Styrian Basin (46°53’51.1” N, 15°46’21.3’’ E; 32 km
SE Graz, 5 km NW Gnas; 320 m a.s.l., Fig. 2A, B), is treated
here. It exposes a palaeosol, rich in terrestrial gastropods.
Additional fossil findings, which include plants, ostracods,
fish remains and mammals, are quite diverse. Hence, this
locality promises a further insight into the so far poorly studied
hinterland of the Paratethyan aquatic system during late
middle Miocene times. The present study focuses on the
taxonomic evaluation of the found gastropods and their
palaeo
environmental and stratigraphic indication, supple-
mented by sedimentological and additional palaeontological
information.
The geological context
The Badenbrunn gravel pit is located in the Eastern Styrian
Basin, more precisely in the Gnas Subbasin (Kröll et al. 1988;
Gross et al. 2007; Fig. 2A). According to detailed geological
mappings by Kollmann (1965) the outcrop exposes upper
Sarmatian sediments (Gleisdorf Formation; Fig. 2B). Close by
exploration drillings of the crude oil mining company RAG
(= Rohöl-Aufsuchungs AG; e.g., F 883 and F 884; ~ 1000 m
SE, ~ 298 m a.s.l.; Fig. 2B) penetrated at a depth of about
9 m clays and marls with abundant “Pirenella picta”
[= Granulolabium bicinctum (Brocchi, 1814)] shells, which is
a marine batillariid gastropod (unpublished report RAG,
1954). In a regional context, these gastropods and the nearby
(around the town of Gnas) occurring oolitic limestones are
typical for the marine upper Sarmatian deposits of the Central
Paratethys (Kollmann 1965; Piller et al. 2007). About 1400 m
to the west (~ 370 m a.s.l.; “Kratzlwirt”, see Fig. 2B), Mytilopsis
ornithopsis (Brusina, 1892) findings are mentioned by Winkler
(1927) and Kollmann (1965). They are indicative for the early
Pannonian (“Zone B” sensu Papp 1951). Drilling F 868
reached at the same sea level, about 500 m towards the W,
beds with abundant dreissenid bivalve fragments (“Congeria
sp.”), which most probable refer to the early Pannonian (early
Tortonian) as well. Accordingly, the section of Badenbrunn is
bracketed by sediments of the late Sarmatian Sea in the foot-
wall and by deposits of brackish Lake Pannon (e.g., Magyar et
al. 1999; Gross et al. 2011b) in the hanging wall.
31
LIFE IN THE FLUVIAL HINTERLAND OF THE LATE SARMATIAN SEA IN THE STYRIAN BASIN (AUSTRIA)
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Langhian
Tortonian
Series
Standard
stages
Messinian
Serra-
vallian
Miocene
Age (Ma)
15
10
9
8
7
11
12
14
13
Middle
Late
7.25
11.62
13.82
Central
Paratethys
Badenian
Pannonian
Chrons
Molluscs
Upper
Zone
Ervilia
Lower
Z.
Ervilia
Mohren-
sternia
Zone
Sarmati-
mactra
vitaliana
Zone
Mytilopsis
ornithopsis
Zone
Mytilopsis
hoernesi
Zone
C5r.2n
C5r.2r-1n
C5n.2n
C5An.1n
C5An.2n
C5Ar.1r
C5An.1r
C5r.3r
C5r.2r
C5r.2r-1r
C5r.1r
C5r.1n
Age
(Ma)
Early Pannonian
Late Sarmatian
Early Sarmatian
11
12
11.5
12.5
Ries Fm.
Gratk.
CG
F
. & P
.
Gleisdorf Fm.
Gleisdorf Fm.
Rollsdorf
SH
Rollsdorf Fm.
Fm./Styrian Basin
proximal - distal
major
hiatus
major
hiatus
further Fms.
not displayed
G
M
gravel
sand
pelite
siliciclastic-oolitic
Badenbrunn
Gratkorn
Mataschen
G
M
SH (Stallhofen Fm.)
Gratk. (Gratkorn Fm.)
CG (Carinthian Gravel)
F. & P. (Feldbach Fm. & Paldau Fm.)
Sar.
Material and sample preparation
The outcrop was vertically logged by visual inspection of the
lithofacies (thickness, colour, grain size, bedding planes, sedi-
mentary structures, macrofossil content); lithofacies coding
follows the scheme of Miall (1996). Samples were taken from
bed 4 and 5 for more detailed palaeontological analyses (Fig. 3).
About 10 kg from layer 4b were repeatedly soaked in water
and dried over 4 weeks and washed through standard sieves
(250/500 μm). Both fractions were picked out completely.
About 400 g of layers 5a and 5b were disintegrated by using
diluted hydrogen peroxide (H
2
O
2
: H
2
O = 1 : 4) and sieved (stan-
dard sieves: 63/125/250/500 μm). Sieve resi duals were picked
out completely. Inspections of microfossils were done through
an optical microscope (Leica M205 C; camera: Leica DFC290;
software: Leica Application Suite) and/or using a scanning
electron microscope (JEOL JSM- 6610LV).
The Badenbrunn section
The lowermost 1.50 m (bed 1) of the outcrop is formed by
orange to yellow, trough cross-bedded, clast-supported, sandy
coarse–medium gravel (Gt) and dm-thick, cross-bedded sand
interlayers (St; Fig. 3). A general fining upward trend is visi-
ble. The uppermost 0.20 m consists of partly carbonatic
cemented fine gravel including irregular,
lateritic encrustations. Above (bed 2),
0.70 m thick grey, trough cross-bedded
medium sand (St) with partly ripple bed-
ded fine sand layers (Sr) is developed.
Scattered pebbles and clay intraclasts are
embedded. From section metres 2.20 to
2.90 (bed 3), the sediments consist of
grey and orange, parallel bedded, badly
sorted sandy silt and silty clay alterna-
tions (cm-scale; Fl) with cm- thick con-
cretionary layers; the bedding planes are
inclined (250/30). Up-section (layer 4a),
1.10 m yellow, indistinctly parallel bed-
ded silt and silty fine sand alternations
(Fl; partly convolute bedded) follow; the
bedding planes are inclined (30–0°) with
about the same dip as bed 3. In the upper-
most 0.10 m rare gastropods are present.
The transition upwards to layer 4b is
gradual. Layer 4b is 0.15 m thick and
comprises orange-yellow to olive-grey
mottled, massive silt (Fr). It contains
abundant gastropods and oxidized root
traces (reaching down into layer 4a) as
well as a few characean gyrogonites and
rare vertebrate remains. This stratum is
conformably overlain by bed 5 (Fm),
which is divided into 0.12 m brown grey,
discontinuously laminated fine sandy silt,
containing abundant leaves and other plant fragments as well
as rare bivalves and ostracods (layer 5a), and 0.13 m grey or
light yellow, laminated clay (layer 5b) with bivalves (union-
ids), plant fragments (i.e., Glyptostrobus cones and twigs;
gyro gonites), fish remains, ostracods and rare gastropods (i.e.,
Ferrissia). A cm-thick layer of light grey, thin laminated clay
forms the boundary with bed 6. 1.00 m of light grey to orange-
brown, horizontal bedded silt and sand alternations follow
above (bed 6; Fl). In the lower 0.15 m abundant plant frag-
ments (i.e., Trapa nuts) are incorporated. About 0.50 m from
the base, a grey, cm-thick, massive clay layer is intercalated.
Close to the upper boundary a ripple-bedded fine sand lens
(Sr), up to 40 cm thick, is developed. The upper boundary is
formed by 50 mm thick, dark–light grey, laminated silty clay
(Fm), containing fragments of plants, mostly leaves. Above
(bed 7), 0.55 m of reddish brown, laminated alterations of silty
clay without plant fragments and silt with plant fragments
(leaves, wood) are recorded (Fl). A thin layer of white, calcic
nodules marks the beginning of bed (8): 0.25 m greyish brown,
horizontal bedded fine to medium sand (Sh). Bed 9 is inter-
nally differentiated by layers of white calcic nodules and its
sedimentary structures: 9a consists of 0.20 m of grey–orange
ripple-bedded sand (Sr). Above (layer 9b), 0.80 m of grey–
orange, low-angle cross-bedded medium sand (Sl) is deve-
loped. A 10 mm thick layer of white, calcic nodules separates
it from the topmost, >2.00 m thick part of the section (layer 9c),
Fig. 1. Stratigraphic position of the Badenbrunn outcrop (modified after Gross et al. 2014;
international and regional stratigraphic scheme based on Neubauer et al. 2015a).
32
DOUBRAWA, GROSS and HARZHAUSER
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
which consists of dm-thick sets of grey–orange, ripple- and
horizontal bedded medium sand (Sr, Sh) with cm-thick layers
of calcic nodules.
Systematic palaeontology
The fossil material is stored in the collection of the
Universal museum Joanneum, Department for Geology &
Palaeontology, Graz (UMJGP). Chronostratigraphic correla-
tions are based on Neubauer et al. (2015b).
Molluscs
The systematic arrangement follows Bouchet et al. (2005),
Jörger et al. (2010) and Welter-Schultes (2012). Names in the
synonymy lists are reproduced strictly as originally provided
by the authors.
Fig. 2. Location of the Badenbrunn pit. A — Geological
sketch of the Styrian Basin (modified after Gross et al.
2007); B — Simplified geological map of the Badenbrunn
area (modified after Kollmann 1965).
33
LIFE IN THE FLUVIAL HINTERLAND OF THE LATE SARMATIAN SEA IN THE STYRIAN BASIN (AUSTRIA)
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Class: Gastropoda Cuvier, 1795
Subclass: Caenogastropoda Cox, 1960
Order: Littorinimorpha Golikov & Starobogatov, 1975
Superfamily: Truncatelloidea Gray, 1840
Family: Bithyniidae Troschel, 1857
Genus: Bithynia Leach, 1818
Bithynia sp.
Fig. 4 A–B
Material: layer 4b: 3 opercula (UMJGP 211.480).
Dimensions: largest specimen: height: 2.6 mm, diameter:
1.8 mm.
Remarks: The drop-shaped outline is characteristic for
Bithynia. Identification on the species level, however, is diffi-
cult without the shells.
Palaeoecology: Recent Bithynia species live in vegetated
slow moving or standing freshwater (Welter-Schultes
2012).
gravel
medium sand
fine sand
silt
clay
calcic nodules
clay intraclasts
massive, blocky
parallel bedding
ripple bedding
low-angle cross bedding
cross bedding
convolute bedding
gastropod sample
ostracod sample
gyrogonites
leaves / plant fragments
fructifications
roots
bivalves
gastropods
ostracods
fish scales
fish teeth
mammal remains
2
3
4a
4b
5a
5b
6
7
8
9a
9b
9c
1
cla
y
sand
silt fine
medium coar
se
gra
ve
l
bed number
6
5a
1
4b
9b
= 0.50 m
Gt
St
St
Sr
Fl
Fl
Fr
Fm
Fl
Fl
Fl
Sr
Sh
Sr
Sl
Sr, Sh
2 cm
2 cm
0.5 cm
10 cm
10 cm
Fig. 3. Lithological section of the Badenbrunn
outcrop.
34
DOUBRAWA, GROSS and HARZHAUSER
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Subclass: Heterobranchia Burmeister, 1837
Clade: Panpulmonata Jörger, Stöger, Kano, Fukuda,
Knebelsberger & Schrödl, 2010
Order: Hygrophila Férussac, 1822
Superfamily: Planorboidea Rafinesque, 1815
Family: Planorbidae Rafinesque, 1815
Subfamily: Ancylinae Rafinesque, 1815
Genus: Ferrissia Walker, 1903
Ferrissia cf. wittmanni (Schlickum, 1964)
Fig. 4 C
cf. 1964 Ancylus wittmanni n. sp. – Schlickum, p. 15, pl. 2, figs.
36–38.
cf. 2014 Ferrissia wittmanni (Schlickum, 1964) – Harzhauser et al.,
p. 18, pl. 5, figs. 3–4, 13 [cum syn.].
Material: layer 5b: 1 specimen (UMJGP 211.481).
Dimensions: length: 0.60 mm.
Remarks: A single fragment of a Ferrissia is available
showing a blunt apex with smooth initial part and broad collar
of prominent radial ribs as described for F. wittmanni by
Kowalke & Reichenbacher (2005) and Harzhauser et al.
(2014). This sculpture allows a separation from the wide-
spread middle Miocene Ferrissia deperdita (Desmarest,
1814), which has faint radial threads (see Harzhauser et al.
2014; Salvador & Rasser 2014; Salvador et al. 2016; Neubauer
et al. 2017).
Distribution: F. wittmanni was originally described from
the Ottnangian (early Miocene) of Bavaria (Schlickum 1964)
but was subsequently also documented from middle Miocene
strata of Hungary and Austria (Kókay 2006; Harzhauser et al.
2014).
Palaeoecology: Ferrissia species are freshwater inhabitants
(Welter-Schulte 2012).
Superorder: Eupulmonata Haszprunar & Huber, 1990
Infraorder: Acteophila Dall, 1885
Superfamily: Ellobioidea Adams & Adams, 1855
Family: Carychiidae Jeffreys, 1830
Genus: Carychium Müller, 1773
Carychium sandbergeri Handmann, 1887
Fig. 5 A–F
* 1887 Carychium Sandbergeri Handm. – Handmann, p. 46.
1923 Carychium sandbergeri Handmann – Wenz, p. 1199.
1942 Carychium sandbergeri Handmann – Wenz & Edlauer, p. 84,
pl. 4, fig. 4.
1977 Carychium (Saraphia) sandbergeri Handmann – Strauch,
p. 167, pl. 16, figs. 36, 38.
1974b Carychium sandbergeri Handmann – Papp, p. 381, pl. 17, fig. 2.
1978 Carychium (Saraphia) sandbergeri Handmann – Schlickum,
p. 248, fig. 1.
1981 Carychium (Saraphia) pachychilus Sandberger – Lueger, p. 14,
pl. 1, figs. 5–8, 9–10.
2004 Carychium sandbergeri Handmann – Harzhauser & Binder,
p. 14, pl. 6, figs. 1–2, 11, 14.
Material: layer 4b: 5 specimens and fragments (UMJGP
211.482).
Dimensions: height: 1.80 mm, diameter: 0.75 mm.
Description: Fusiform, slender shells with 4 convex spire
whorls. Bulbous protoconch of 0.3 mm diameter with inflated
initial part and faintly pitted surface. Early teleoconch with
faint spiral threads close to the lower and upper sutures.
Later teleoconch whorls with prominent, prosocline growth
lines. Last whorl high, subcylindric. Aperture with strongly
thicke
ned peristome, well-rounded basal lip and broad,
reflected inner lip. Three prominent lamellae comprising
a slender and large parietal lamella, a slightly weaker and
lower columellar lamella and a knob-like palatal lamella.
Parietal lip well demarcated from base with straight margin,
forming a distinct angulation with the weakly rounded
inner lip.
Remarks: As only a few complete specimens were
avai lable, and they turned out to be very fragile, the columellar
area was not opened for closer inspection, although it
is suggested in Strauch (1977) and Stworzewicz (1999a)
to be essential for identification. The Badenbrunn
specimens agree with shells of C. sandbergeri from
the Pannonian of the Vienna Basin (e.g., Harzhauser &
Binder 2004) in sculpture and apertural features. Extremely
slender morphotypes as described by Strauch (1977)
and Stworzewicz (1999a), however, are
missing at
Badenbrunn.
Distribution: Middle to late Miocene in the Carpathian
Foredeep, Pannonian Basin and Vienna Basin (late Badenian:
Oppeln (Poland); Sarmatian: Rákosd, Várpalota (Hungary);
middle–late Pannonian: Richardhof, Eichkogel (Austria), Öcs
(Hungary); Wenz & Edlauer 1942; Schütt 1967; Harzhauser &
Binder 2004; Kókay 2006).
Palaeoecology: Most probably riparian areas and very
moist habitats, as well as woodlands (Schütt 1967; Harzhauser
& Binder 2004).
Fig. 4. A, B: Bithynia sp.; C: Ferrissia cf. wittmanni (Schlickum,
1964).
35
LIFE IN THE FLUVIAL HINTERLAND OF THE LATE SARMATIAN SEA IN THE STYRIAN BASIN (AUSTRIA)
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Carychium nouleti Bourguignat, 1857
Fig. 5 G–I
*1857 Carychium nouleti Bourguignat, p. 226.
1923 Carychium nouleti nouleti Bourguignat – Wenz, p. 1195.
1977 Carychium (Saraphia) nouleti Bourguignat – Strauch, p. 162,
pl. 15, figs. 24–27, pl. 18, fig. 61, pl. 20, fig. 83.
1999a Carychium nouleti Bourguignat – Stworzewicz, p. 269, figs.
19–22.
2004 Carychium nouleti Bourguignat – Binder, p. 194, pl. 3, fig. 2.
2016 Carychium (Saraphia) nouleti Bourguignat – Salvador et al.,
p. 136, fig. 2U.
2017 Carychium nouleti Bourguignat, 1857 s.l. – Neubauer et al.,
p. 740, fig. 4 A–E, J, O.
Material: layer 4b: 2 specimens (UMJGP 211.483).
Dimensions: height: 1.65 mm, diameter: 0.75 mm.
Description: Stout shell of 4.5 whorls with conical spire
and deep sutures. Spire whorls strongly convex; last whorl
very high. Bulbous protoconch 2.5 mm in diameter with
weakly convex initial part and faintly pitted surface, persis-
ting on the first teleoconch whorl, which bears faint spiral
threads. Sculpture of teleoconch consisting of moderately
prominent, densely spaced, prosocline growth lines.
Ovoid aperture with moderately thickened peristome and
weakly reflected inner lip; columellar lamella broad and low,
parietal lamella large and slender, palatal lamella broad,
knob-like.
Remarks: The identification remains tentative without
information on the internal columellar apparatus. At first
glance, the shells might be identified as stout specimens of
C. sandbergeri from which they differ in the much weaker
sculpture, broader aperture and much higher last whorl. In
addition, the parietal lamella is weaker and has a more adapi-
cal position compared to C. sandbergeri. In overall outline, the
shells are reminiscent of the late Miocene Carychiopsis
berthae (Halaváts, 1903), as described by Strauch (1977),
Harzhauser & Binder (2004) and Katona et al. (2015) from
Austria and Hungary. A clear difference, however, is the much
weaker axial sculpture and the absence of the second parietal
lamella.
Distribution: C. nouleti appears during the early Miocene
in France, Austria and Poland and is widespread during the
middle Miocene with records from Austria, Germany and
France (Wenz 1923; Stworzewicz 1999a; Binder 2004;
Salvador et al. 2016). The youngest reliable records are men-
tioned from the late Miocene mammal zone MN9 by
Stworzewicz (1999a) from Poland.
Palaeoecology: Most probably riparian areas and very
moist habitats, as well as woodlands (Schütt 1967).
Infraorder: Stylommatophora Schmidt, 1855
Non-achatinoid clade: sensu Wade et al., 2006
Unassigned “subclade”: Orthuretha Pilsbry, 1900
Superfamily: Pupilloidea Turton, 1831
Family: Vertiginidae Fitzinger, 1833
Subfamily: Gastrocoptinae Pilsbry, 1916
Genus: Gastrocopta Wollaston, 1878
Gastrocopta fissidens (Sandberger, 1863)
Fig. 5 J–O
* 1863 Pupa fissidens sp. nov. – Sandberger, p. 57, pl. 5, figs. 16a–c.
1914 Leucochilus fissidens (Sandberger) – Fischer & Wenz, p. 97,
pl. 6, fig. 19.
1999b Gastrocopta cf. ferdinandi (Andreae 1902) – Stworzewicz,
p. 165, fig. 63.
2013 Gastrocopta fissidens (Sandberger, 1863) – Stworzewicz et al.,
p. 188, fig. 4D.
Material: layer 4b: 13 specimens, many fragments (UMJGP
211.484).
Dimensions: largest specimen: height: 1.80 mm, diameter:
0.91 mm.
Description: Small, ovate-turreted shell with 3.5 strongly
convex teleoconch whorls separated by deeply incised sutures.
Protoconch consisting of 1.2 convex whorls of 0.4 mm dia-
meter with wrinkled microsculpture. Shell surface covered by
prominent, prosocline growth lines. Last whorl moderately
high, convex, terminating in subcircular aperture. Peristome
continuous, broad, weakly reflexed, well detached from base.
Umbilicus narrow, largely covered by inner lip. Parieto-
angular tooth strongly divided into two bent teeth. A weak
infraparietal tooth may be largely reduced in some specimens;
columellar lamella large, subhorizontal; knob-shaped basal
lamella appears deeper in aperture. Three long parietal
lamellae, becoming successively smaller adapically.
Remarks: The species is characterized by its strongly
convex whorls and the bifid parieto-angular tooth. Both
features allow a separation from the similar middle and late
Miocene Gastrocopta ferdinandi (Andreae, 1902) and
G. serotina sensu Stworzewicz et al. (2013). Wenz (1923,
p. 930) doubted, that the Sarmatian specimens from Poland are
conspecific with the Oligocene and early Miocene G. fissidens
and suggested a relation with G. obstructa (Sandberger, 1875),
which however is higher and more cylindrical.
Distribution: The oldest records of G. fissidens derive from
the Oligocene of Germany, where it is also frequently recorded
from lower Miocene sites (Wenz 1923). In Poland, this species
is recorded from the latest early to middle Miocene (mammal
zones MN5 to MN7/8) by Stworzewicz (1999b) and
Stworzewicz et al. (2013). A smaller morphotype was described
as Gastrocopta fissidens infrapontica by Wenz (1927) from the
late Miocene (early Pannonian) of the Vienna Basin.
Palaeoecology: Recent Gastrocopta are found in meso-
phytic and swamp forests (Stworzewicz et al. 2013).
Family: Vertiginidae Fitzinger, 1833
Subfamily: Nesopupinae Steenberg, 1925
Genus: Negulopsis Nordsieck, 2014
Negulopsis suturalis (Sandberger, 1858)
Fig. 5 P–R
* 1858 Pupa suturalis A. Braun – Sandberger, p. 54, Pl. 6, figs.
2–2a.
36
DOUBRAWA, GROSS and HARZHAUSER
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
1919 Negulus suturalis gracilis n. var. – Gottschick & Wenz, p. 9,
pl. 1, figs. 11–13.
1981 Negulus suturalis gracilis Gottschick u. Wenz – Lueger, p. 18,
pl. 2, figs. 2a–b.
2004 Negulus gracilis Gottschick & Wenz – Harzhauser & Binder,
p. 18, pl. 6, figs. 9–10.
2008 Negulus gracilis Gottschick & Wenz – Harzhauser et al., p. 50,
fig. 5.4.
2013 Negulus suturalis (Sandberger, 1858) – Stworzewicz et al.,
p. 191, fig. 4L.
2014 Negulus suturalis (Sandberger, 1858) – Harzhauser et al.,
p. 854, figs. 8K–N.
2015 Negulopsis suturalis (Sandberger, 1858) – Harzhauser et al.,
p.29, pl. 5, figs. 5–6.
Material: layer 4b: 1 specimen, 1 aperture fragment
(UMJGP 211.485).
Dimensions: height: 1.70 mm, diameter: 0.80 mm.
Remarks: This widespread species was described and
discussed in detail recently by Stworzewicz et al. (2013) and
Harzhauser et al. (2014, 2015). We follow these authors and
especially Stworzewicz (1999b) who treat the late Miocene
Negulopsis gracilis (Gottschick & Wenz, 1919) as a subjec-
tive junior synonym of N. suturalis. The alleged morphologi-
cal features characterizing N. gracilis (smaller size and slender
shape) are also observed in various early and middle Miocene
populations of N. suturalis.
Distribution: N. suturalis is known from the Oligocene of
Germany, the early Miocene of Germany, Czech Republic,
Poland, Austria, Hungary and the middle Miocene of Poland,
Hungary and Austria (Stworzewicz 1999b; Kókay 2006;
Harzhauser et al. 2014). During the late Miocene, it is
widespread in Austria, Hungary and Slovakia (Harzhauser
& Binder 2004). The last occurrence is documented from
the Messinian of the Po Basin in Italy (Harzhauser et al. 2015).
Palaeoecology: N. suturalis seems to have preferred moist
habitats in riparian areas.
Unassigned “subclade”
Superfamily: Clausilioidea Gray, 1855
Family: Filholiidae Wenz 1923
Genus: Triptychia Sandberger, 1875
Fig. 5. A–F: Carychium sandbergeri Handmann, 1887; G–I: Carychium nouleti Bourguignat, 1857; J–O: Gastrocopta fissidens (Sandberger,
1863); P–R: Negulopsis suturalis (Sandberger, 1868).
37
LIFE IN THE FLUVIAL HINTERLAND OF THE LATE SARMATIAN SEA IN THE STYRIAN BASIN (AUSTRIA)
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Triptychia sarmatica Kókay, 2006
Fig. 6 D–G
2006 Triptychia leobersdorfensis sarmatica nov. ssp. – Kókay ,
p. 82, pl. 31, figs. 5–7, pl. 41, figs. 6–7.
2006 Triptychia (Triptychia) leobersdorfensis sarmatica Kókay,
2006 – Schnabel, p. 153, pl. 4, figs. 53–54.
Material: layer 4b: 11 specimens (UMJGP 211.354,
211.355, 211.357, 211.361, 211.367, 211.370, 211.372,
211.382, 211.404, 211.405a, 211.405b).
Dimensions: largest specimen (without initial spire whorls):
height: 42 mm, diameter: 11 mm.
Description: Moderately large, slender shell consisting of
>12 teleoconch whorls (due to sediment compaction most
Fig. 6. A–C: Palaeotachea sylvestrina (Schlotheim, 1820); D–G: Triptychia sarmatica Kókay, 2006; H–I: Archaeozonites costatus Sandberger,
1875; J: Klikia giengensis Klein, 1846; K–N: Pseudochloritis gigas (Pfeffer, 1930); O–P: Pseudoleacina sp.; Q: Anodonta cf. carinthiaca
Modell, 1957.
38
DOUBRAWA, GROSS and HARZHAUSER
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
specimens appear very broad); earliest spire whorls turreted,
weakly convex; later spire whorls widening, nearly flat. Early
spire whorls with very prominent, regular and densely spaced
axial ribs, which become weaker, thinner and slightly wavy on
later spire whorls and continue on last whorl. Weak, narrow
spiral incisions may appear in the upper half of the last whorl.
Adapical tips of the axial ribs produce a weakly crenulate,
slightly wavy suture. Aperture not wide, drop-shaped; two
columellar lamellae with a slightly more prominent upper
lamella; parietal lamella larger; none of the lamellae reach the
peristome. Sinulus long, narrow, acute. Columellar lip and
parietal lip well detached from base.
Remarks: This species is frequently found at Badenbrunn.
Despite the moderate preservation, its size, characteristic axial
sculpture and the long, narrow and acute sinulus correspond
well with T. sarmatica from the Sarmatian of Várpalota in
Hungary. The late Miocene Triptychia leobersdorfensis (Troll,
1907) has a shorter and wider sinulus and its axial sculpture is
less prominent.
Palaeoecology: Triptychia species are considered as indica-
tors for moist and warm forests (Binder 2002, 2008; Schnabel
2007).
Unassigned “subclade”: Limacoidea
Superfamily: Limacoidea Lamarck, 1801
Family: Limacidae Lamarck, 1801
Limacidae gen. et sp. indet.
Fig. 7 A
Material: layer 4b: 4 specimens and fragments (UMJGP
211.486).
Dimensions: largest specimen: width: 2 mm, length: 3 mm.
Remarks: The elongate ovoid outline of the calcitic plates
and their projecting, asymmetrical nucleus support their place-
ment in the Limacidae. For the same reasons as for the
Milacidae, the authors refrain from identification on species or
genus level.
Family: Agriolimacidae Wagner, 1935
Agriolimacidae gen. et sp. indet.
Fig. 7 B
Material: layer 4b: 1 specimen (UMJGP 211.487).
Dimensions: width: 1.9 mm, length: 3.2 mm.
Remarks: Based on the outline with the subparallel mar-
gins and the only moderately displaced nucleus, this single
specimen is placed here in the Agriolimacidae as described by
Reuse (1983) and Frank (2006). As for all slugs, a more
detailed identification is impossible.
Superfamily: Zonitoidea Mörch, 1864
Family: Zonitidae Mörch, 1864
Genus: Archaeozonites Sandberger, 1875
Archaeozonites costatus Sandberger, 1875
Fig. 6 H–I
* 1875 Archaeozonites costatus Sandberger – Sandberger, p. 604.
1916 Zonites (Aegopsis) costatus (Sandberger) – Gottschick &
Wenz, p. 21, pl. 1, figs. 1a–c.
1976 Archaeozonites costatus Sandberger – Schlickum, p. 18, pl. 5,
fig. 66.
2002 Miozonites costatus (Sandberger) – Binder, p. 168, pl. 1, figs.
14–16, pl. 3, fig. 8, pl. 7, fig. 3, pl. 8, fig. 4.
2013 Miozonites costatus (Sandberger, 1874) – Stworzewicz et al.,
p. 194, fig. 5L.
2014 Miozonites costatus (Sandberger) – Harzhauser et al., p. 33, pl.
10, figs. 13–19.
2015a Archaeozonites costatus Sandberger – Salvador et al., p. 259,
figs. 3Q–R.
2015b Archaeozonites costatus Sandberger, 1875 – Salvador et al.,
p. 206, figs. 2O–P.
2016 Archaeozonites costatus Sandberger, 1875 – Salvador &
Rasser, p. 49, fig. 2Z.
Material: layer 4b: 2 specimens (UMJGP 211.383,
211.396).
Dimensions: Both specimens are deformed, which makes
precise measurements difficult. 211.383 is vertically com-
pressed: height: 12 mm, diameter: 22 mm. 211.396 shows
horizontal compression: height: 4 mm, diameter: 15 mm.
Description: Medium sized shells consisting of at least 4
teleoconch whorls; spire whorls moderately convex with deep
sutures. Last whorl with weak angulation at mid-whorl, coin-
ciding with the maximum diameter. Sculpture consisting of
densely spaced and slightly irregular prosocline ribs on spire
whorls; ribs become slightly sigmoidal on last whorl and fade
out below angulation. Umbilicus moderately wide and deep.
Aperture destroyed.
Remarks: The fragmented (or subadult) specimens are
rather high spired and have a very strong axial sculpture. In
these features, they correspond well to the shells described by
Binder (2004) from the late early Miocene of the Korneuburg
Basin in Austria. The sculpture is also identical to specimens
from the early middle Miocene of Styria described by
Fig. 7. A: Limacidae gen. et sp. indet.; B: Agriolimacidae gen. et sp.
indet.; C–D: Milacidae gen. sp. indet.
39
LIFE IN THE FLUVIAL HINTERLAND OF THE LATE SARMATIAN SEA IN THE STYRIAN BASIN (AUSTRIA)
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Harzhauser et al. (2014) supporting the identification of the
poorly preserved material.
Distribution: A widespread species, which appears during
the late early Miocene and becomes ubiquitous in the middle
Miocene. Occurrences are documented from Switzerland,
southern Germany, the North Alpine Foreland Basin (Germany,
Austria), the Korneuburg Basin (Austria) and the Rein Basin
(Austria; Wenz 1923; Binder 2002, 2004; Harzhauser et al.
2014; Salvador & Rasser 2014, 2016; Salvador et al. 2015a).
The northernmost distribution was described by Stworzewicz
et al. (2013) from the Sarmatian of Poland. Middle Miocene
specimens from drillings in Hungary, described by Kókay
(2006) as “Aegopsis costatus” might rather represent another
species.
Palaeoecology: Moist woodland (Binder 2002; Harzhauser
et al. 2014).
Superfamily: Parmacelloidea Fischer, 1856
Family: Milacidae Ellis, 1926
Milacidae gen. et sp. indet.
Fig. 7 C–D
Material: layer 4b: 7 specimens and fragments (UMJGP
211.488).
Dimensions: largest specimen: width: 3 mm, length: 5 mm.
Remarks: Based on their symmetrical, calcitic, broad ovoid
plates with nearly straight sides and a peripheral, convex
nucleus, we place these specimens in the Milacidae. The iden-
tification of Milacidae species or genera based solely on plates
is nearly impossible, because diagnostic features of slugs are
mostly related to soft body anatomy (Wiktor & Likharev
1979).
Unassigned “subclade”
Superfamily: Helicoidea Rafinesque, 1815
Family: Elonidae Gittenberger, 1979
Genus: Klikia Pilsbry, 1895
Klikia giengensis (Klein, 1846)
Fig. 6 J
* 1846 Helix giengensis – Klein, p. 69, pl. 1, fig. 9.
1911 Klikia giengensis Krauss – Wenz, p. 85, pl. 4, figs. 6–10.
1916 Klikia (Klikia) giengensis (Klein) – Gottschick & Wenz, p. 56.
1923 Klikia (Klikia) giengensis giengensis (Klein) – Wenz, p. 539.
1976 Klikia (Klikia) giengensis (Klein) – Schlickum, p. 16, pl. 4, fig. 58.
2006 Klikia (Klikia) giengensis (Klein) – Kókay, p. 91, pl. 25, fig. 9.
2013 Klikia giengensis (Klein) – Stworzewicz et al., p. 195, fig. 6A.
2014 Klikia giengensis (Klein) – Harzhauser et al., p. 33, pl. 11, figs.
9–11, 22.
Material: layer 4b: 1 fragmentary specimen (UMJGP
211.348b).
Dimensions: about 10 mm diameter.
Description: A single fragment is available comprising 5
densely coiled spire whorls with flat spire. Microsculpture
consisting of a dense pattern of tiny papillae on entire spire.
Base and aperture are destroyed or covered by sediment.
Remarks: Despite the fragmentary preservation, the flat
spire, the tightly coiled spire whorls and the dense pattern of
papillae are very characteristic for K. giengensis and allow
a separation from all other species of the genus.
Distribution: This species appears already during the early
Miocene (Binder 2004) and occurs around the early/middle
Miocene boundary in France and southern Germany. In the
Styrian Basin, it is recorded from lower Badenian strata
(Harzhauser et al. 2014) and is frequently mentioned from
the middle Miocene Silvana-beds in Germany (Mörsingen,
Hohenmemmingen, Zwiefaltendorf) (Schlickum 1976).
Sarmatian occurrences are documented from Várpalota in
Hungary (Kókay 2006).
Palaeoecology: This species seems to have preferred moist
woodlands (Harzhauser et al. 2014).
Family: Helicidae Rafinesque, 1815
Genus: Palaeotachea Jooss, 1912
Palaeotachea sylvestrina (Schlotheim, 1820)
Fig. 6 A–C
* 1820 Helicites sylvestrinus – Schlotheim, p. 99.
1923 Cepaea sylvestrina sylvestrina (Schlotheim) – Wenz, p. 690.
1974b Cepaea sylvestrina (Schlotheim) – Papp, p. 391, pl. 19, figs.
1–5.
2016 Palaeotachea sylvestrina (Schlotheim, 1820) – Höltke &
Rasser, p. 245, figs. 5.6–5.9, 6.5.
Material: layer 4b: 8 specimens (UMJGP 211.408,
211.409a, 211.409b, 211.410, 211.411a, 211.412, 211.416,
211.418).
Dimensions: largest specimen: height: 15 mm, diameter:
21 mm.
Description: Ovoid-conical shells comprising 3.5 teleoconch
whorls and 1.25 smooth and weakly convex protoconch
whorls. Spire whorls moderately convex with distinct growth
lines on early whorls and deep sutures; last whorl high,
strongly convex. Peristome reflexed, distinctly prosocline in
lateral view. Surface smooth except for growth lines. Colour
pattern in UV-light consisting of three spiral bands on last
whorl; the uppermost spiral band occurs slightly above the
position of the maximum diameter followed by a second one
slightly below and a third one on the base.
Remarks: All specimens are deformed by sediment com-
paction and the apertures are filled by sediment. Therefore, the
identification is tentative. Nevertheless, the high spire and
deep suture allow a separation from the similar Palaeotachea
silvana (Klein, 1853) (see Höltke & Rasser 2016) and the
colour pattern agrees with typical P. sylvestrina from the
Sarmatian of Austria and Hungary (Papp 1974b).
Distribution: Middle Miocene in the North Alpine Foreland
Basin, Southern Germany and the Vienna Basin spanning
mammal zones MN6 to MN7/8 (Papp 1974b; Höltke & Rasser
2016). Early Miocene occurrences, mentioned as Cepaea cf.
40
DOUBRAWA, GROSS and HARZHAUSER
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
sylvestrina by Salvador (2013) need confirmation and are not
listed under this species by Höltke & Rasser (2016). Late
Miocene occurrences might rather represent P. etelkae
(Halaváts, 1923) (Lueger 1981).
Palaeoecology: The ecological requirements of this extant
genus are unknown. Like the recent species of Cepaea, the
Palaeotachea species seem to have settled a broad range of
habitats (see Welter-Schultes 2012 for Cepaea ecology).
Genus: Pseudochloritis Boettger, 1909
Pseudochloritis gigas (Pfeffer, 1930)
Fig. 6 K–N
* 1930 Tropidomphalus (Pseudochloritis) gigas – Pfeffer, p. 76.
1957 Tropidomphalus (Pseudochloritis) gigas Pfeffer – Papp, p. 86.
1967 Tropidomphalus (Pseudochloritis) gigas Pfeffer – Schütt,
p. 219, fig. 23.
1974b Tropidomphalus (Pseudochloritis) gigas Pfeffer – Papp,
p. 389, pl. 18, figs. 2a–c.
1981 Tropidomphalus (Pseudochloritis) gigas Papp – Lueger, p. 58,
pl. 12, fig. 4, pl. 13, fig. 4, pl. 16, fig. 5.
2008 Pseudochloritis gigas (Pfeffer) – Binder, p. 174, pl. 2, figs.
4a–c.
Material: layer 4b: 10 specimens (UMJGP 211.340,
211.346, 211.347, 211.348a, 211.380, 211.381, 211.413,
211.415, 211.417a, 211.417b).
Dimensions: largest specimen: height: 26 mm, diameter:
33 mm.
Description: Large helicoid shells with depressed spire;
protoconch comprising 1.5 feebly convex whorls with delicate
ribs and tiny papillae. Teleoconch consisting of 3.5 whorls
with incised suture and wide, high and regularly convex last
whorl; sculpture consisting of weak prosocline growth lines,
being typically most prominent close to the upper suture.
Terminal last whorl bends down abruptly close to the aperture;
peristome thickened with strongly reflexed lip and slight
constriction right before aperture. Deep, moderately wide
umbilicus partly covered by inner lip. No colour pattern visible
in UV-light.
Remarks: This is the largest gastropod species at
Badenbrunn and can be recognized easily by its depressed
spire, strongly reflexed peristome and only partly covered
umbilicus. The Pannonian Pseudochloritis depressus (Wenz,
1927) is smaller and the spire whorls are less tightly coiled;
P. richarzi (Schlosser, 1907) is much smaller and has a higher
spire (see Lueger 1981).
Distribution: P. gigas is widespread during the Sarmatian
in the North Alpine Foreland Basin, the Vienna Basin, the
Lavanttal Basin and the Bakony Mountains (Lueger 1981;
Kókay 2006; Binder 2008). The youngest records derive from
the early Pannonian of the Vienna Basin (Lueger 1981).
Palaeoecology: Lueger (1981) proposed vegetated areas
near creeks or ponds as the habitat of this species;
Binder (2008) and Höltke & Rasser (2015) discussed moist
habitats.
Achatinoid clade: sensu Wade et al., 2006
Superfamily: Testacelloidea Gray, 1840
Family: Oleacinidae H. & A. Adams, 1855
Genus: Pseudoleacina Wenz, 1914
Pseudoleacina sp.
Fig. 6 O–P
2006 Palaeoglandina gracilis porrecta (Gobanz) 1854 – Kókay, p. 85,
pl. 32, fig. 8 [non Paleoglandina porrecta (Gobanz, 1854)].
2006 Poiretia ex. aff. taurinensis (Sacco) 1886 – Kókay, p. 86, pl.
32, fig. 11 [non Palaeoglandina taurinensis (Sacco, 1886)].
Material: layer 4b: 1 compressed specimen (UMJGP
211.358), 2 fragments (UMJGP 211.369, 211.375).
Dimensions: UMJGP 211.358 height of fragment: 36 mm,
diameter (compressed): 15 mm.
Description: The available specimens show an elongated
ovoid shell, with high, nearly straight-sided spire whorls and
moderately high, subcylindric last whorl, attaining about 1/3
of the total height. Suture narrow but deeply incised. Sculpture
consisting of prominent, densely spaced, slightly irregular,
weakly prosocline axial ribs. The axial ribs are united in
an indistinct narrow adsutural cord along the upper suture.
Peristome thin; aperture drop-shaped; columella slightly
twisted, anteriorly truncated. Inner lip forming a distinct callus
attached to the base.
Remarks: Kókay (2006) documented this species under
two names from Sarmatian deposits of Várpalota in Hungary.
All his specimens are fragmentary and derive from drillings.
One specimen was identified by Kókay (2006) as
Palaeoglandina gracilis porrecta (Gobanz, 1854), which
differs in its inflated last whorl and much shorter spire (see
Harzhauser et al. 2014 for discussion on this species). The
same features allow a clear separation from the Miocene
Palaeoglandina gracilis (von Zieten, 1832) (see Harzhauser et
al. 2014 and Salvador et al. 2015a). Another specimen was
identified by Kókay (2006) as Poiretia ex. aff. taurinensis
(Sacco, 1886) from which it differs clearly in its fusiform out-
line and much stronger sculpture (see Ferrero-Mortara et al.
1984 for the syntype of the early Miocene Italian Glandina
taurinensis Sacco, 1886). Pseudoleacina eburnea (Klein,
1853), from the middle and late Miocene of Germany and
Austria, is much smaller and has much more delicate sculpture
(see Harzhauser & Binder 2004; Salvador et al. 2015a).
Pseudoleacina rakosdensis (Gaál, 1911), from the Sarmatian
of Hungary, differs in its slender outline and the axial ribs,
which fade out below the shoulder. Pseudoleacina kleiniana
(Pilsbry, 1909), from the middle Miocene of southern Germany,
differs in its fusiform last whorl (see Schlickum 1976).
The specimen from Badenbrunn and the coeval specimens
from Hungary seem to represent a new, undescribed species.
Unfortunately, the preservation of the available material is
not suitable to serve as type specimens to formally establish
a new species.
Distribution: This species is known so far only from the
Sarmatian of Várpalota in Hungary and Badenbrunn in Styria.
41
LIFE IN THE FLUVIAL HINTERLAND OF THE LATE SARMATIAN SEA IN THE STYRIAN BASIN (AUSTRIA)
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Class: Bivalvia Linnaeus, 1758
Subclass: Palaeoheterodonta Newell, 1965
Superfamily: Unionoideae Rafinesque, 1820
Family: Unionidae Rafinesque, 1820
Genus: Anodonta Lamarck, 1799
Anodonta cf. carinthiaca Modell, 1957
Fig. 6 Q
cf. 1957 Anodonta splendens carinthiaca n. ssp. – Modell, p. 98,
figs. 3–4.
cf. 2001 Anodonta (Anodonta) splendens carinthiaca Modell, 1957
– Schultz, p. 379, figs. 8a–b.
Material: layer 5b: 1 specimen and several fragments
(UMJGP 211.489).
Dimensions: length: 54.9 mm, height: 32.8 mm.
Description: An articulated specimen with moderately
fragile valves is available; hinge and umbonal area are
destroyed. Dorsal margin straight, passing into evenly rounded
anterior margin. Ventral margin rounded; posterior margin
subangulate with long, nearly straight posterodorsal area.
Maximum height in posterior third.
Remarks: Based on the characteristic outline and the rather
thin shell, the specimen seems to represent an Anodonta carin
thiaca, which was described by Modell (1957) from the
Sarmatian of Slovenia. This species was separated by Modell
(1957) from the mainly early Miocene Anodonta splendens
(Goldfuss, 1837) because of stratigraphic reasons. A syste-
matic revision of this group, however, is beyond the scope of
this paper.
Palaeoecology: Anodonta species prefer calm freshwater in
lakes, and oxbow lakes with muddy or sandy substrate and
rarely appear in running waters (Welter-Schultes 2012).
Ostracods
Suprageneric classification and terminology of valve
characters follow Meisch (2000) and Sames (2011), respec-
tively. Abbreviations: L = left valve, R = right valve; f = female,
j = juvenile, A-1, ... = juvenile stages; l = length, h = height (both
in millimetres).
Class: Ostracoda Latreille, 1802
Order: Podocopida Sars, 1866
Superfamily: Cypridoidea Baird, 1845
Family: Candonidae Kaufmann, 1900
Subfamily: Candoninae Kaufmann, 1900
Genus: Fabaeformiscandona Krstić, 1972
Fabaeformiscandona cf. balatonica (Daday, 1894)
Fig. 8. A–C
Material: layer 5b: 4 valves, 28 fragments (UMJGP
211.490).
Dimensions: Rf: l = 0.98–1.01 mm, h = 0.52–0.54 mm; Lf:
l = 1.02 mm, h = 0.52– 0.63 mm.
Remarks: At Badenbrunn only few, fragmented female
valves have been found. They are very similar to F. balatonica
– at least in a wider sense (e.g., Absolon 1970; Janz 1997;
Meisch 2000; Pipík 2001; Gross 2004; Fuhrmann & Goth
2011; Witt 2011; Fuhrmann 2012). The early Karpatian–middle
Badenian (Central Paratethyan distribution) species Fabae
formiscandona pokornyi (Kheil, 1964), which could be the
ancestor of F. balatonica, differs by its more pointed postero-
ventral end (e.g., Witt 1998, 2000; Gross 2006; Reischenbacher
et al. 2007). Due to the limited material and the still unresolved
variability of this species(-group), an assured determination is
not possible (Janz 1997; Meisch 2000).
Distribution: (of F. balatonica s.l.): middle Miocene
(Badenian) to recent (Pipík 2001; Witt 2011).
Palaeoecology: Extant F. balatonica favours shallow
ephemeral pools and swampy marginal lake settings but also
occurs in the littoral of lakes, in rivers and densely vegetated
rivulets. Potentially, it is able to cope with slightly elevated
(oligohaline) salinity (Meisch 2000; Pipík 2001; Gross 2004).
Fabaeformiscandona sp. 1
Fig. 8 E–H
Material: layer 5a: 2 valves, 6 fragments; layer 5b: 112
juvenile valves, 14 fragments (UMJGP 211.491).
Dimensions: Lj: l = 0.61– 0.65 mm, h = 0.31– 0.33 mm; Rj:
l = 0.61– 0.62mm, h = 0.31– 0.32 mm.
Remarks: These juvenile (?A-2) candoninae shells are the
most frequent ostracod remains occurring at the Badenbrunn
locality. Perhaps, these valves represent larval stages of F. cf.
balatonica (see Gross 2004).
Palaeoecology: See F. cf. balatonica.
Fabaeformiscandona sp. 2
Fig. 8 I
Material: layer 5a: 1 R fragment (UMJGP 211.492).
Dimensions: l = ? mm, h = 0.42 mm.
Remarks: This specimen displays some similarity with
Fabaeformiscandona fabaeformis (Fischer, 1851), which is,
however, more elongated (Janz 1997; Fuhrmann 2012). By
neglecting its significantly larger size, Fabaeformiscandona
hyalina (Brady & Robertson, 1870) match very well solely
based on the outline (see Fuhrmann 2012). As the posterior
end of the shell is missing and only one fragment is available,
specific identification is unfeasible.
Palaeoecology: See F. cf. balatonica.
Genus: Pseudocandona Kaufmann, 1900
Pseudocandona cf. praecox (Straub, 1952)
Fig. 8 J
Material: layer 5a: 1 Lj, 1 Rj; layer 5b: 1 Rj, 1 fragment
(UMJGP 211.493).
Dimensions: Rj: l = 0.63 mm, h = 0.32 mm.
42
DOUBRAWA, GROSS and HARZHAUSER
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Fig. 8. A–C: Fabaeformiscandona cf. balatonica (Daday, 1894) (A: L; B: R; C: R, central muscle scars);
D: Ilyocypris ?ex gr. gibba (Ramdohr,
1808), R;
E–H: Fabaeformiscandona sp. 1 (E: L; F: L; G: R; H: R); I: Fabaeformiscandona sp. 2, R; J: Pseudocandona cf. praecox (Straub,
1952), R; K–L: Paralimnocythere rostrata (Straub, 1952) (K: L; L: L).
43
LIFE IN THE FLUVIAL HINTERLAND OF THE LATE SARMATIAN SEA IN THE STYRIAN BASIN (AUSTRIA)
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Remarks: These valves probably represent the A-2 stage of
a Pseudocandona species and resemble juveniles of P. praecox
(e.g., Witt 1998; Schäfer 2005), which is suggested to be
a junior synonym (Janz 1992, 1997; Witt 2000) or not (Schäfer
2005, 2011) of Pseudocandona steinheimensis (Sieber, 1905).
The material available at present does not enable a more pre-
cise identification as well as a decision about this possible
synonymy (see Janz & Matzke-Karasz 2001; Gross 2004).
Distribution: (of P. praecox): lower Oligocene to late mid-
dle Miocene (Sarmatian) in the Swiss Molasse Basin, North
Alpine Foreland Basin and Třeboň Basin (see Witt 1998,
2000; Schäfer 2005, 2011).
Palaeoecology: Presumably a dweller in stagnant fresh-
waters (ponds and lakes; see Janz 1997).
Family: Ilyocyprididae Kaufmann, 1900
Subfamily: Ilyocypridinae Kaufmann, 1900
Genus: Ilyocypris Brady & Norman, 1889
Ilyocypris ?ex gr. gibba (Ramdohr, 1808)
Fig. 8 D
Material: layer 5b: 3 R, 1 fragment (UMJGP 211.494).
Dimensions: R: l = 0.75mm, h = 0.44 mm.
Remarks: Only few, to some degree fragmented Ilyocypris
valves are available. This hampers a closer determination by
considering the variability of Ilyocypris’ shell characters (e.g.,
nodes). Partly species-specific “inner marginal ripplets” are
not preserved in the current specimens (Janz 1994; Meisch
2000; Mazzini et al. 2014).
Among Central European Miocene Ilyocypris species, I. gibba
of Witt (2000; although somewhat more elongated and larger)
and the valve fragment (Ilyocypris sp.) of Gross (2004) are
most similar. Equally, extant I. gibba (e.g., Triebel 1941;
Fuhrmann 2012) resembles the present specimens. Due to the
lack of knowledge about marginal ripplets neither species
identification nor a profound allocation to the “gibba”-group
are possible (Witt 2002).
Distribution: (I. gibba of Witt 2000 and Ilyocypris sp. of
Gross 2004): early (Ottnangian) to late Miocene (early Panno-
nian) in the North Alpine Foreland Basin and Styrian Basin.
Palaeoecology: Ilyocypris occurs in shallow, fresh to slightly
saline (oligohaline) water bodies (ponds, lakes, springs, rivers;
e.g., Morkhoven 1963; Meisch 2000; Karanovic 2012).
Superfamily: Cytheroidea Baird, 1850
Family: Limnocytheridae Klie, 1938
Subfamily: Limnocytherinae Klie, 1938
Genus: Paralimnocythere Carbonnel, 1965
Paralimnocythere rostrata (Straub, 1952)
Fig. 8 K–L
* 1952 Limnocythere rostrata n. sp. – Straub, p. 499–500, pl. C, figs.
55–56, text-fig. 23.
1965 Limnocythere rostrata Straub 1952 – Lutz, p. 310–311, text-
fig. 26.
1989 Limnocythere rostrata Straub 1952 – Reichenbacher, pl. 3, fig. 13.
2000 Paralimnocythere rostrata (Straub 1952) – Witt, p. 119, pl. 1, fig. 3.
2011 Paralimnocythere rostrata (Straub 1952) – Pirkenseer &
Berger, p. 32–33, pl. 3, figs. 4a–b.
Material: layer 5a: 1 Lf; layer 5b: 1 Lf, 3 fragments
(UMJGP 211.495).
Dimensions: Lf: l = 0.56–0.58 mm, h = 0.31 mm.
Remarks: These very fragile, female valves match well
with P. rostrata in Pirkenseer & Berger (2011). In the latter,
however, the reticulum forms low, oblique ridges in the antero-
ventral area, which are not developed in the specimens of
Badenbrunn. Given the variation in ornamentation of this spe-
cies (fide synonyms included by Pirkenseer & Berger 2011),
this slight difference is suggested to represent intraspecific
variability. Potentially, Paralimnocythere sp. of Pipík (1998:
Danube Basin; middle Pannonian) also belongs to P.
rostrata
but the provided (tilted) illustration does not allow a detailed
comparison. The late Miocene (Tortonian) species Para
limnocythere bouleigensis Carbonnel, 1965 is similar (com-
pare figures of paratypes in Martens 1992) but diverges by its
more pronounced ventrolateral swelling and pitted surface
(see also Carbonnel 1969). Paralimnocythere tenera Sokač,
1972 from the Pannonian of Croatia is similar but its anterior
and posterior parts are finely punctuated. Sokač (1972) has not
mentioned a reticulum, which is clearly visible in our speci-
mens. Unfortunately, the illustrations of this author do not
allow a detailed examination and a reinvestigation of P. tenera
is needed to clarify its relation to P. rostrata.
Distribution: (fide Pirkenseer & Berger 2011): Oligocene
to middle Miocene (Badenian) in the Upper Rhine Graben,
Swiss Molasse Basin, Aquitaine Basin, North Alpine Foreland
Basin and North Croatian Basin (see Witt 2000; Hajek-Tadesse
et al. 2009).
Palaeoecology: P. rostrata probably inhabits shallow
freshwater environments, like ponds, lakes and rivers (see
Pirkenseer & Berger 2011 and ecological demands summa-
rized for extant Paralimnocythere species in Meisch 2000).
Vertebrates
The taxonomy used here follows Böhme (2010: fishes),
Prieto et al. (2010: rodents) and Aiglstorfer et al. (2014:
proboscideans).
Class: Actinopterygii Cope, 1887
Order: Cypriniformes Bleeker, 1859
Family: Cyprinidae Bonaparte, 1832
Subfamily: Cyprininae Bonaparte, 1840
Genus: Palaeocarassius Obrhelová 1970
Palaeocarassius sp.
Fig. 9 A–C
Material: layer 5b: 2 posterior teeth, 1 pharyngeal tooth
(UMJGP 211.496; provisional determination M. Böhme).
44
DOUBRAWA, GROSS and HARZHAUSER
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Distribution: Early Miocene to late Miocene (late
Pannonian) in the Central Paratethys (Böhme & Ilg 2003;
Böhme 2010 and personal comment M. Böhme, 2015).
Palaeoecology: An inhabitant of lacustrine environments
(Böhme 1999, 2010).
Subfamily: Leuciscinae Howes, 1991
Genus: Palaeoleuciscus Obrhelová 1969
Palaeoleuciscus sp.
Fig. 9 D–E
Material: layer 5b: 7 pharyngeal teeth (UMJGP 211.497;
provisional determination M. Böhme).
Remarks: Four teeth have a pointed terminal hook, the
other three show marks of usage or are fragments. The teeth
show a wide tooth crown, narrowing towards the shaft.
Distribution: Early Miocene to late Miocene (middle
Pannonian) in the Central Paratethys (Böhme & Ilg 2003 and
personal comment M. Böhme, 2015).
Palaeoecology: Probably an inhabitant of rivers and asso-
ciated backwaters (Böhme 2002).
Class: Mammalia Linne, 1758
Order: Rodentia Bowdich, 1821
Family: Muridae Illiger, 1811
Subfamily: Copemyinae Jacobs & Lindsay, 1984
Genus: Democricetodon Fahlbusch, 1964
Democricetodon sp. nov. (sensu Kälin & Engesser, 2001)
Fig. 9 F
Material: layer 4b: one isolated m1 (UMJGP 211.498;
provisional determination J. Prieto).
Remarks: Democricetodon sp. nov. (sensu Kälin &
Engesser, 2001) is a small hamster. The anterocone of the m1
is round to ovaloid, the long mesolophid reaches the edge of
the tooth. The molar has two roots (see Prieto et al. 2010).
Distribution: in Central Europe: late middle Miocene (Sarma-
tian) to late Miocene (middle Pannonian; Gross et al. 2011a).
Palaeo-ecology: A wet and wooded habitat has been sug-
gested for Democricetodon sp. nov. (sensu Kälin & Engesser,
2001) (Wessels & Reumer 2009; Prieto et al. 2014).
Order: Proboscidea Illiger, 1811
Family: Deinotheriidae Bonaparte, 1845
Genus: Deinotherium Kaup, 1829
Deinotherium levius Jourdan, 1861
Remarks: Mottl (1958) described a tusk, which previously
was found at Badenbrunn (UMJGP 54.563). She identified it
as Deinotherium levius Jourdan, 1861 and suggested a late
Sarmatian age for the specimen. Nevertheless, Mottl (1958)
mentioned that Deinotherium species identifications, solely
based on tusks, are difficult and are not conclusive for strati-
graphic purposes (see Aiglstorfer et al. 2014).
Discussion
Sedimentology
Due to poor outcrop conditions, observations are confined
to vertical section analysis, which renders an interpretation
difficult (Bridge 1993). Our tentative facies interpretation is
based on Miall (1996) and Einsele (2000). More detailed
sedimentological investigations are needed, but this is beyond
our possibilities.
The basal (bed 1), cross-bedded gravels (Gt) with some
cross-bedded sand lenses (St) indicate sedimentation from
unidirectional currents and are interpreted as channel-fill or
transverse bar deposits and remnants of 3-D dunes of a fluvial
system. Up-section (bed 2), trough cross- and subordinately
ripple-bedded sands (St, Sr) refer to the development of 3-D
dunes/ripples. Bed 2 is overlain by silt–fine sand alternations
(beds 3–4a; Fl), which display obliquely (~30°) oriented
bedding planes. Towards the hanging wall (top of layer 4a),
the bedding gradually converts into subhorizontal stratifi-
cation (layer 4b). The lithofacies of these beds implies deposi-
tion from suspension and low velocity currents, for example,
in overbank areas or abandoned channels. Due to the upward
decrease in the dip of bedding planes, deposition by lateral
accretion (point bar) can be suspected. The massive, blocky
structured layer 4b (Fr) is rich in terrestrial gastropods and
notably affected by roots, penetrating up to ~0.8 m into layer
4a. This indicates incipient pedogenic overprint of suspension
load deposits in an overbank environment or abandoned
channel. Bed 4b is topped by pelitic sediments (bed 5; Fm),
containing aquatic biota (molluscs, ostracods, fishes). These
characteristics refer to deposition from suspension in standing
waters like in floodplain ponds or oxbow lakes.
Above (beds 6–9), the sediments coarsen upwards pointing
to increased influx of the trunk river. Partly plant-rich
Fig. 9. A–C: Palaeocarassius sp. (A: pharyngeal tooth; B–C:
posterior teeth); D–E: Palaeoleuciscus sp., pharyngeal teeth;
F: Democricetodon sp. nov. (sensu Kälin & Engesser, 2001), m1.
45
LIFE IN THE FLUVIAL HINTERLAND OF THE LATE SARMATIAN SEA IN THE STYRIAN BASIN (AUSTRIA)
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
sand–silt alternations (beds 6–7; Fl), indicative for inter-
changes of weak traction current and suspension sedimenta-
tion, might be deposited during floodwaters (distal crevasse
splays?) in a vegetated floodplain. Upper plane bed conditions
are indicated by the horizontal bedded fine–medium sand (Sh)
of bed 8, possibly formed during the initial stage of a crevasse
splay and followed by ripple-bedded sand deposition (layer
9a; Sr) as the flow diminished. The outcrop-wide, approxi-
mately uniformly low-angle bedded (Sl) layer 9b could repre-
sent a sandy channel fill (crevasse channel?). Cm-thick calcic
layers of nodules in the ripple- and horizontal bedded layer 9c,
possibly mark further individual sheet flooding/crevasse splay
events.
Palaeoecology
In total, 15 gastropod and 1 bivalve species were found at
the Badenbrunn section. Terrestrial taxa derive mainly from
the palaeosol-layer 4b, where aquatic gastropods (only 3
Bithynia-opercula) are almost absent. As discussed in the
taxonomic descriptions, most taxa indicate humid woodland,
which is especially supported by the high number of Triptychia
specimens. The presence of two Carychium species suggests
moist bottom conditions, for example, with leaf litter.
Therefore, the gastropod assemblage points to a vegetated,
moist environment with nearby water bodies (e.g., poorly
drained floodplain or abandoned
channel). A rather moist forest
habitat has been assumed for the
late Sarmatian Democricetodon
species (Prieto et al. 2010, 2014).
However, a single tooth is cer-
tainly not significant for palaeo-
environmental reconstructions.
The ostracod fauna of the over-
lying bed 5 gives a clear indica-
tion for a freshwater environment,
most probably of a shallow pond or
lake. No indications of any marine
influx — for example, of ostracods,
typical for the Sarmatian Sea —
have been found. In accordance,
the recovered fish remains and
molluscs (Anodonta, Ferrissia)
point to a limnic setting. Although
more detailed sampling and taxo-
nomic works are necessary concer-
ning the plant material, abun dant
Glyptostrobus remains in layer 5b
hint at a swampy, very shallow
environment (Averyanov et al.
2009). Shallow, nutrient-rich lentic
waters are indicated by the pre-
sence of plentiful occurrences of
Trapa nuts (Kovar-Eder et al.
2002, Meller & Hofmann 2004).
With the gradual return to a more proximal setting in respect
to the trunk river (beds 6–7), only plant fragments of the
riparian forest (e.g., Alnus, Salix) can be found.
Biostratigraphy
The mollusc assemblage indicates a middle Miocene age.
Taxa, such as Ferrissia cf. wittmanni, Triptychia sarmatica,
Klikia giengensis, Archaeozonites costatus and Palaeotachea
sylvestrina, are unknown from the late Miocene (Fig. 10).
The frequent Pseudochloritis gigas persisted into the early
Pannonian but is most widespread during the Sarmatian.
Therefore, the stratigraphic ranges of the taxa exclude
a Pannonian age (late Miocene) and point to a Sarmatian (late
middle Miocene) age. A Badenian age can be excluded due to
the presence of P. gigas.
The ostracod fauna found in layer 5, overlying the palaeosol
(4b), is quite poor both in abundance and in diversity.
Specimens are largely fragmented and juvenile valves are
common. Due to these limitations, the biostratigraphic value
of the ostracod fauna is restricted. However, there are some
implications, arguing for a middle Miocene, possibly
Sarmatian age: Fairly speculatively, Fabaeformiscandona
balatonica s.l. appears in middle Badenian times as a des-
cendant of Fabaeformiscandona pokornyi and may, hence,
hint at an age not older than the middle Badenian.
Fig. 10. Stratigraphic distribution of the Badenbrunn gastropod fauna.
46
DOUBRAWA, GROSS and HARZHAUSER
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Fabae formiscandona sp. 1 (possibly juveniles of F. cf.
balatonica) and Fabaeformiscandona sp. 2 do not offer
a biostratigraphic implication to date. Pseudocandona praecox
would point to an age not younger than Sarmatian times,
however, this claim remains tentative due to ambiguous
taxonomic identification of the current material. Due to
similarities with Ottnangian–early Pannonian Ilyocypris
species, Ilyocypris ?ex. gr. gibba vaguely refers to this time
range. Despite some similarities with Pannonian
Paralimnocythere species, Paralimnocythere rostrata is
reported until now only from the Oligocene to the middle
Miocene.
The isolated tooth of Democricetodon sp. nov. (sensu Kälin
& Engesser, 2001) provides only a rough biostratigraphic indi-
cation. This species is found in Central Europe during the late
Sarmatian to the middle Pannonian times.
Palaeocarassius and Palaeoleuciscus teeth have been found
co-occurring in layer 5b. Both are reported separately through-
out the Miocene, up to the Pannonian. However, coexisting
records of both are unknown from the late Miocene so far
(personal comment, M. Böhme, 2015).
In consideration of the hitherto available palaeontological
indications and the available geologic information, a late
middle Miocene (late Sarmatian) age is plausible for the
Badenbrunn locality.
Conclusions
The Badenbrunn pit exposes fluvial sediments ranging from
thalweg to point bar and overbank deposits (e.g., poorly
drained floodplain, floodplain pond or abandoned channel,
crevasse splay/channel).
A moderately diverse terrestrial gastropod fauna settling
a moist and vegetated floodplain or abandoned channel has
been found. Subsequently, this fauna was replaced by
an aquatic (freshwater) assemblage due to floodplain pond or
oxbow lake development.
The recovered mollusc assemblage points to a Sarmatian
age (late middle Miocene). Ostracod, fish and rodent remains
support this dating. Based on the geolo gical background
(marine upper Sarmatian sediments in the footwall; brackish
lower Pannonian sediments in the hangingwall), a late
Sarmatian age of this site is most plausible.
Terrestrial deposits, close to the Sarmatian/Pannonian
boundary in age are virtually undescribed from the Styrian
Basin and rare in the Central Paratethyan realm as well. For
example, the localities of Gratkorn (~ 42 km NW) and
Mataschen (~14 km E) are slightly older or younger respec-
tively (Fig. 1). Both sites differ notably in their faunal compo-
sition (Gross 2008; Gross et al. 2011a, b, 2014; Harzhauser et
al. 2008).
Our study contributes to a better understanding of the palae-
obiological and palaeoecological characteristics of the
Paratethyan hinterland around the middle–late Miocene tran-
sition, which is, however, a period of critical turnovers in
aquatic and continental ecosystems of Central Europe (e.g.,
Piller & Harzhauser 2005, Böhme et al. 2008, 2011).
Acknowledgements: We thank Madelaine Böhme (Tübingen)
and Jérôme Prieto (Munich) for determination of the fish and
rodent material. Thanks go to Norbert Winkler (UMJ, Graz)
for his assistance during fieldwork and sample preparation.
For access to the pit, we are grateful to Rupert Niederl (Baden-
brunn). We are thankful to two anonymous reviewers for their
helpful comments.
References
Absolon A. 1970: Fossiles Vorkommen von Candona balatonica
Daday 1894 (Ostracoda). Mitt.
Bayer. Staatssamml. Paläontol.
Hist. Geol. 10, 199–204.
Aiglstorfer M., Göhlich U.B., Böhme M. & Gross M. 2014: A partial
skeleton of Deinotherium (Proboscidea, Mammalia) from the
late Middle Miocene Gratkorn locality (Austria). Palaeobio.
Palaeoenv. 94, 49–70.
Andreae A. 1902: Zweiter Beitrag zur Binnenconchylienfauna des
Miocäns von Oppeln in Schlesien. Mitt. Roemer. Mus.,
Hildesheim 18, 1–31.
Averyanov L.V., Phan K.L., Nguyen T.H., Nguyen S.K., Nguyen T.V.,
Pham T.D. 2009: Preliminary Observation of Native Glypto
strobus pensilis (Taxodiaceae) Stands in Vietnam. Taiwania 54,
191–212.
Binder H. 2002: Die Land- und Süßwassergastropoden aus dem
Karpatium des Korneuburger Beckens. Beitr. Paläont. 27,
161–203.
Binder H. 2004: Terrestrial, freshwater and brachyhaline Gastropoda
from the Lower Miocene deposits of Oberdorf (Styria, Austria).
Ann. Naturhist. Mus. Wien 105 A, 189–229.
Binder H. 2008: The systematic positions of the genera Pseudo
chloritis C. Boettger 1909 and Joossia Pfeffer 1929 (Gastro-
poda: Pulmonata: Heliocoidea: Helicidae). Arch. Molluskenkd.
137, 167–193.
Böhme M. 1999: Die miozäne Fossil-Lagerstätte Sandelzhausen. 16.
Fisch- und Herpetofauna - Erste Ergebnisse. N. Jahrb. Paläont.
Geol. Abh. 214, 487–495.
Böhme M. 2002: New approaches investigating freshwater
palaeoecosystems. Eur
op. Palaeont. Ass.: Workshop on Fresh
water and Brackish (Palaeo)ecosystems, Fribourg University,
23–30.
Böhme M. 2010: Ectothermic vertebrates (Actinopterygii, Allo-
caudata, Urodela, Anura, Crocodylia, Squamata) from the Mio-
cene of Sandelzhausen (Germany, Bavaria) and their implica-
tions for environment reconstruction and palaeoclimate. Paläont.
Z. 84, 3–41.
Böhme M. & Ilg A. 2003: Database of Lower Vertebrates. fosFAR
base: http://www.wahre-staerke.com.
Böhme M., Ilg A. & Winklhofer M. 2008: Late Miocene “washhouse”
climate in Europe. Earth Planet. Sci. Lett. 275, 393–401.
Böhme M., Winklhofer M. & Ilg A. 2011: Miocene precipitation in
Europe: Temporal trends and spatial gradients. Palaeogeogr.
Palaeoclimatol. Palaeoecol. 304, 212–218.
Bouchet P., Rocroi J.P., Frýda J., Hausdorf B., Ponder W., Valdés Á.
& Warén A. 2005: Classification and nomenclator of gastropod
families. Malacologia. 47, 1–2, 1–397.
Bourguignat J.R. 1857: Aménités Malacologiques, LXIV. Du genre
Carychium. Rev. et Mag. de Zool. 2, 9, 5, 209–232.
Brady G.S. & Robertson D. 1870: The Ostracoda and Foraminifera of
tidal rivers. Part 1. Ann. Mag. Nat. Hist. 4, 6, 1, 1–33.
47
LIFE IN THE FLUVIAL HINTERLAND OF THE LATE SARMATIAN SEA IN THE STYRIAN BASIN (AUSTRIA)
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Bridge J.S. 1993: Description and interpretation of fluvial deposits:
a critical perspective. Sedimentology 40, 801–810.
Brusina S. 1892: Über die Gruppe der Congeria triangularis.
Z. dtsch. Geol. Ges. 44, 488–497.
Carbonnel G. 1965: Sur un nouveau genre (Paralimnocythere) et une
nouvelle espèce (P. bouleigensis) d’ostracodes du Tortonien.
Arch. Sci. 18, 1, 146–150.
Carbonnel G. 1969: Les Ostracodes du Miocène Rhodanien (systéma-
tique, biostratigraphie, écologique, paéobiologie). Doc. Lab.
Géol. Fac. Sci. Lyon. 32, 1, 1–228.
Daday J. 1894: Weitere Erkenntnisse zur Balaton-Mikrofauna (Nema-
toda, Rotaoria, Entomostraca) [Ujabb adatok a Balaton mikro-
faunájának ismeretéhez (Nematoda, Rotatoria, Entomostraca)].
Math. és Term. Ert. 12, 122–145 (in Hungarian).
Desmarest A.G. 1814: Note sur les Ancyles ou Patelles d’eau douce,
et particulièrement sur deux espèces de ce genre, non encore
décrites, l’une fossile et l’autre vivante. Bull. Soc. Philom. Paris
3, 18–20.
Einsele G. 2000: Sedimentary Basins: Evolution, Facies, and Sedi-
ment Budget. SpringerVerl., Berlin/Heidelberg, 1–792.
Ferrero-Mortara E., Montefameglio L., Novelli M., Opesso G., Pavia
G. & Tampieri R. 1984: Katalog von Typen und Exemplaren der
Bellardi und Sacco Sammlung [Catalogo dei tipi e degli
esemplari figurati della collezione Bellardi e Sacco]. Parte II.
Mus. Reg. Sci. nat. Cat. 7, 1–484 (in Italian).
Fischer S. 1851: Abhandlung über das Genus Cypris und dessen in
der Umgebung von St. Pétersbourg und von Fall bei Reval
vorkommenden Arten. Mém. Acad. Imp. Sci. St. Pétersbourg 7,
127–167.
Fischer K. & Wenz W. 1914: Die Landschneckenkalke des Mainzer
Beckens und ihre Fauna. Jahrb. Nassau. Ver. Naturkd. 67, 1–154.
Frank C. 2006: Plio-pleistozäne und holozäne Mollusken Österreichs.
Mitt. Prähist. Komm. Österr. Akad. Wiss. 62, 1–860).
Fuhrmann R. 2012: Atlas quartärer und rezenter Ostrakoden Mittel-
deutschlands. Altenbg. naturwiss. Forsch. 15, 1–320.
Fuhrmann R. & Goth K. 2011: Neue und weitere bemerkenswerte
Ostrakoden aus dem Quartär Mitteldeutschlands. Palaeontogr. A
294, 95–201.
Gaál I. 1911: Eine sarmatische Schneckenfauna aus dem ungarischen
Rákosd [A Hunyadmegyei Rákosd szarmatakorú csigafaunája].
Magy. Földt. Intéz. Èvk. 18, 1–108 (in Hungarian).
Gobanz J. 1854: Die fossilen Land- und Süsswasser-Mollusken des
Beckens von Rein in Steiermark. Sitzb. K. Akad. Wiss.,
math.naturwiss. Cl. Wien 13, 1, 180–201.
Goldfuss G.A. 1837: Petrefacta Germaniae tam ea, quae in museo
universitatis regiae Borussicae Fridericiae Wilhelmiae Rhenanae
servantur quam alia quaecunque in Museis Hoeninghusiano
Muensteriano aliisque extant ; iconibus et descriptionibus illus-
trata. Abbildungen und Beschreibungen der Petrefacten Deutsch-
lands und der angränzenden Länder. Unter Mitwirkung des
Herrn Grafen Georg zu Münster; herausgegeben von August
Goldfuss. 2 (1834–40), Divisio quarta: Molluscorum Acephali-
corum Reliquiae — Muschelthiere der Vorwelt, I. Bivalvia. Arnz
& Co., Düsseldorf, 65–286.
Gottschick F. & Wenz W. 1916: Die Sylvanaschichten von Hohen-
memmingen und ihre Fauna. Nachr. Bl. dtsch. malak. Ges. 48,
17–31, 55–71, 97–113.
Gottschick F. & Wenz W. 1919: Die Land- und Süßwassermollusken
des Tertiärbeckens von Steinheim am Aalbuch. 1. Die Vertigi-
niden. Nachr. Bl. dtsch. malak. Ges. 51: 1–23.
Gross M. 2004: Zur Ostracodenfauna (Crustacea), Paläoökologie und
Stratigrafie der Tongrube Mataschen (Unter-Pannonium, Stei-
risches Becken, Österreich). Joannea Geol. Paläont. 5, 49–129.
Gross M. 2006: Mittelmiozäne Ostracoden aus dem Wiener Becken
(Badenium/Sarmatium, Österreich). Österr. Akad. Wiss.,
Schriftenr. Erdwiss. Komm., spec. issue 1, 1–224.
Gross M. 2008: A limnic ostracod fauna from the surroundings of the
Central Paratethys (Late Middle Miocene/Early Late Miocene;
Styrian Basin; Austria). Palaeogeogr. Palaeoclimatol. Palaeo
ecol. 264, 263–276.
Gross M., Fritz I., Piller W.E., Soliman A., Harzhauser M., Hubmann
B., Moser B., Scholger R., Suttner T.J., Bojar H.-P. 2007: The
Neogene of the Styrian Basin - Guide to Excursions. Joannea.
Geol. Paläont. 9, 117–193.
Gross M., Böhme M. & Prieto J. 2011a: Gratkorn: A benchmark
locality for the continental Sarmatian sensu stricto of the Central
Paratethys. Int. J. Earth Sci. 100, 1496–1913.
Gross M., Piller W.E., Scholger R. & Gitter F. 2011b: Biotic and
abiotic response to palaeoenvironmental changes at Lake
Pannons’ western margin (Central Europe, Late Miocene).
Palaeogeogr., Palaeoclimatol., Palaeoecol. 312, 181–193.
Gross M., Böhme M., Havlik P. & Aiglstorfer M. 2014: The late
Middle Miocene (Sarmatian s.str.) fossil site Gratkorn — the
first decade of research, geology, stratigraphy and vertebrate
fauna. Palaeobiodivers. Palaeoenvir
on. 94, 5–20.
Hajek-Tadesse V., Belak M., Sremac J., Vrsaljko D. & Wacha L.
2009: Early Miocene ostracods from the Sadovi section (Mt
Požeška gora, Croatia). Geol. Carpathica 60, 3, 251–262.
von Halaváts G. 1903: Die Fauna der Pontischen Schichten in der
Umgebung des Balatonsees. Result. wiss. Erforsch. Balatonsees
1, 1, 1–80.
von Halaváts G. 1923: Eine baltavarsische Molluskenfauna vom
oberen See [A baltavarsi felsöpontusi koru Molluszka fauna].
Magy. Földt. Intéz. Èvk. 24, 395–407 (in Hungarian).
Handmann P.R. 1887: Die fossile Conchylienfauna von Leobersdorf
im Tertärbecken von Wien. Verl. Aschendorff’schen Buchhandl.,
Münster, 1–55.
Harzhauser M. & Binder H. 2004: Synopsis of the Late Miocene
mollusc fauna of the classical sections Richardhof and Eich-
kogel in the Vienna Basin (Austria, Pannonian, MN 9-MN11).
Arch. Molluskenkd. 133, 1–2, 1–57.
Harzhauser M. & Piller W.E. 2004: Integrated stratigraphy of the
Sarmatian (Upper Middle Miocene) in the western Central Para-
tethys. Stratigraphy 1, 65–86.
Harzhauser M., Gross M. & Binder H. 2008: Biostratigraphy of
Middle Miocene (Sarmatian) wetland systems in an Eastern
Alpine intramontane basin (Gratkorn Basin, Austria): the terres-
trial gastropod approach. Geol. Carpathica 59, 1, 45–58.
Harzhauser M., Neubauer T.A., Gross M. & Binder H. 2014: The
early Middle Miocene mollusc fauna of Lake Rein (Eastern
Alps, Austria). Palaeontogr. A 302, 1–71.
Harzhauser M., Neubauer T.A., Georgopoulou E., Esu D.,
D’Amico C., Pavia G., Giuntelli P. & Carnevale G. 2015: Late
Messinian continental and Lago-Mare gastropods from the
Tertiary Piedmont Basin, NW Italy. Boll. Soc. Paleontol. Ital. 54,
1, 1–53.
Höltke O. & Rasser M. 2015: Pseudochloritis insignis — A peculiar
large land-snail from the Miocene of SW Germany: Taxonomic
status and census of morphologically related forms. J. Conchol.
42, 1, 1–12.
Höltke O. & Rasser M. 2016: The Palaeotachea complex (Gastropo-
da: Pulmonata) in the Miocene of SW Germany: A morpho-
metric approach. J. Conchol. 42, 4, 239–256.
Janz H. 1992: Die miozänen Süßwasserostrakoden des Steinheimer
Beckens (Schwäbische Alb, Süddeutschland). Stutt. Beitr.
Naturkd. B 183, 1–117.
Janz H. 1994: Zur Bedeutung des Schalenmerkmals „Marginalrip-
pen“ der Gattung Ilyocypris (Ostracoda, Crustacea). Stutt. Beitr.
Naturkd. B, 206, 1–19.
Janz H. 1997: Die Ostrakoden der kleini-Schichten des miozänen
Kratersees von Steinheim am Albuch (Süddeutschland). Stutt.
Beitr. Naturkd. B 251, 1–101.
48
DOUBRAWA, GROSS and HARZHAUSER
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Janz H. & Matzke-Karasz R. 2001: Holozäne Ostrakoden aus Karbo-
natablagerungen im Bereich der neolithischen Feuchtboden-
siedlung Unfriedshausen (Bayern) als ökologische Indikatoren.
Mitt. Bayer. Staatssamml. Paläontol. Hist. Geol. 41, 33–63.
Jörger K.M., Stöger I., Kano Y., Fukuda H., Knebelsberger T. &
Schrödl M. 2010: On the origin of Acochlidia and other enig-
matic euthyneuran gastropods, with implications for the sys-
tematics of Heterobranchia. BMC Evol. Biol. 10, 232, 1–20.
Jourdan C. 1861: Description de restes fossiles de deux grands mam-
mifères constituant deux genres, l’un le genre Rhizoprion de
l’ordre des Cétecés et du groupe des Delphinoides; l’autre le
genre Dynocyon de l’ordre des Carnassiers et de la familie des
Canidés. Ann. Sci. Nat. 16, 369–374.
Kälin D. & Engesser B. 2001: Die jungmiozäne Säugertierfauna vom
Nebelbergweg bei Nunningen (Kanton Solothurn, Schweiz).
Schweiz. palaeontol. Abh. 121, 1–61.
Karanovic I. 2012: Recent Freshwater Ostracods of the World.
Springer, Heidelberg, 1–608.
Katona L., Magyar I., Berta T., Varga A., Sztanó O. 2015: Panno-
nische Molluskenfauna aus zwei Ausgrabungen der Fűzfõ-Bucht
[Pannóniai puhatestű fauna a Fűzfői-öböl környékének két
feltárásából]. Földt. Közl. 145, 127–150 (in Hungarian).
Kheil J. 1964: Die Ostrakoden der Mydlovary-Schichtenfolge im süd-
böhmischen Třeboň-Becken. Sbor. Geol. Věd., Geol. 4, 7–46.
von Klein A. 1846: Conchylien der Süsswasserkalkformation Wür-
ttembergs. Jahresh. Ver. vaterl. Nat. kd. Württ. 2, 60–116.
von Klein A. 1853: Conchylien der Süsswasserkalkformation Wür-
ttembergs. Jahresh. Ver. vaterl. Nat. kd. Württ. 9, 203–223.
Kókay J. 2006: Nonmarine mollusc fauna from the Lower and Middle
Miocene, Bakony Mts, W Hungary. Geol. Hun., ser. Palaeont.
56, 1–196.
Kollmann K. 1965: Jungtertiär im Steirischen Becken. Mitt. Geol.
Ges. Wien 57, 479–632.
Kováč M., Baráth I., Harzhauser M., Hlavatý I. & Hudáčková N.
2004: Miocene depositional systems and sequence stratigraphy
of the Vienna Basin. Cour. Forsch.Inst. Senckenberg 246, 187–212.
Kovar-Eder J., Schwarz J. & Wójcicki J.J. 2002: The predominantly
aquatic flora from Pellendorf, Lower Austria, Late Miocene,
Pannonian – a systematic study. Acta Palaeobot. 42, 125–151.
Kowalke T. & Reichenbacher B. 2005: Early Miocene (Ottnangian)
Mollusca of the Western Paratethys - ontogenetic strategies and
palaeoenvironments. Geobios 38, 609–635.
Kröll A., Flügel H.W., Seiberl W., Weber F., Walach G. & Zych D.
1988: Erläuterungen zu den Karten über den prätertiären Unter-
grund des Steirischen Beckens und der Südburgenländischen
Schwelle. Geol. Bundesanst., Wien, 4 maps.
Lueger J.P. 1981: Die Landschnecken im Pannon und Pont des Wiener
Beckens, I. Systematik, II. Fundorte, Stratigraphie, Faunen-
provinzen. Oesterr. Akad. Wiss. Math.
Natwiss. Kl., Denkschr.
120, 1–157.
Lutz A.K. 1965: Jungtertiäre Süßwasser-Ostracoden aus Süddeutsch-
land. Geol. Jahrb. 82, 271–330.
Magyar I., Geary D.H. & Müller P. 1999: Paleogeographic evolution
of the Late Miocene Lake Pannon in Central Europe. Palaeo
geogr. Palaeoclimatol. Palaeoecol. 147, 151–167.
Martens K. 1992: A reassessment of Paralimnocythere Carbonnel,
1965 (Crustacea, Ostracoda, Limnocytherinae), with a descrip-
tion of a new genus and two new species. Bull. Inst. R. Sci. Nat.
Belg. Biol. 62: 125–158.
Mazzini I., Gliozzi E., Rossetti G. & Pieri V. 2014: The Ilyocypris
puzzle: A multidisciplinary approach to the study of phenotypic
variability. Int. Rev. Hydrobiol. 99: 1–14.
Meisch C. 2000: Freshwater Ostracoda of Western and Central
Europe. In: Schwoerbel, J. & Zwick, P. (Eds.): Süßwasserfauna
von Mitteleuropa, 8, 3. Spektrum Akad. Verl., Heidelberg/Berlin,
1–522.
Meller B. & Hofmann C.-C. 2004: Paläoökologische Interpretation
von Diasporen- und Palynomorphen-Vergesellschaftungen aus
obermiozänen Seesedimenten (Mataschen bei Fehring, Oststeier-
mark, Österreich). Joannea Geol. Paläont. 5: 177–217.
Miall A.D. 1996: The Geology of Fluvial Deposits. SpringerVerl.,
Berlin/Heidelberg/New York, 1–582.
Modell, H. 1957: Najaden aus dem limnischen Tertiär Kärntens.
Carinthia 2, 67, 95–100.
Morkhoven van F.P.C.M. 1963: Post-Palaeozoic Ostracoda, their
morphology, taxonomy, and economic use. Volume 2, Generic
descriptions. Elsevier Publishing Company, Amsterdam, 1–478.
Mottl M. 1958: Neue Säugetierfunde aus dem Jungtertiär der Steier-
mark. Mitt. Mus. Bergbau, Geol. Tech. Landesmus. „Joanneum“
19, 16–41.
Neubauer T.A., Harzhauser M., Kroh A., Georgopoulou E. & Mandic
O. 2015a. A gastropod-based biogeographic scheme of the
European Neogene freshwater systems. EarthSci. Rev. 143,
98–116.
Neubauer T.A., Georgopoulou E., Kroh A., Harzhauser M., Mandic
O. & Esu D. 2015b: Synopsis of European Neogene freshwater
gastropod localities: updated stratigraphy and geography.
Palaeontol. Electr
on. 18.1.3T, 1–7.
Neubauer T.A., Harzhauser M., Mandic O., Jovanović G. 2017: The
late middle Miocene non-marine mollusk fauna of Vračević
(Serbia): filling a gap in Miocene land snail biogeography.
Bulletin of Geoscience, 91, 4, 731–778.
Papp A. 1951: Das Pannon des Wiener Beckens. Mitt. Geol. Ges.
Wien 39–41, 99–193.
Papp A. 1957: Landschnecken aus dem limnischen Tertiär Kärntens.
Mitt. naturwiss. Ver. Kärnten 67, 85–94.
Papp A., Marinescu F. & Senes J. 1974a: M5. Sarmatien. Chrono
stratigr. Neostratotypen, Miozän Zentr. Paratethys 4, 1–707.
Papp A. 1974b: Landschnecken im Sarmatien der Zentralen Paratethys.
In: Papp A., Marinescu F. & Senes, J. (Eds.): Chrono stratigr.
Neostratotypen, Miozän Zentr. Paratethys 4, 377–395.
Pfeffer G. 1930: Zur Kenntnis tertiärer Landschnecken. Geol.
Paläont. Abh. 3, 1–230.
Piller W.E. & Harzhauser M. 2005: The Myth of the Brackish Sarma-
tian Sea. Terra Nova, 17, 450-455.
Piller W., Harzhauser M. & Mandic O. 2007: Miocene Central Para-
tethys stratigraphy — current status and future directions. Strati
graphy 4, 2, 3, 151–168.
Pilsbry H.A. 1909–1910: Caecilioides, Glessula and Partulidae. Man.
Conchol., Second Ser.: Pulmonata 20, 1–336, Acad. Nat. Sci.,
Philadelphia.
Pipík R. 1998: Salinity changes recorded by ostracoda assemblages
found in Pannonian sediments in the western margin of the
Danube Basin. Bull. centr. rech. explor.prod. ElfAquitaine,
Mém. 20, 167–177.
Pipík R. 2001: Les ostracodes d’un lac ancient et ses paléobiotopes au
Miocène supérieur: le basin de Turiec (Slovaquie). Université
ClaudeBernard, Lyon, thesis, 1–337.
Pirkenseer C. & Berger J.P. 2011: Paleogene Ostracoda from the
southern Upper Rhine Graben: Taxonomy, palaeoecology and
palaeobiogeography. Palaeontogr. A 295, 1–152.
Prieto J., Böhme M. & Gross M. 2010: The cricetid rodents from
Gratkorn (Austria, Styria): a benchmark locality for the conti-
nental Sarmatian sensu stricto (late Middle Miocene) in the
Central Paratethys. Geol. Carpath. 61, 5, 419–436.
Prieto J., Angelone C., Casanovas-Vilar I., Gross M., Hír J., van den
Hoek Ostende L.W., Maul L.C. & Vasilyan D. 2014: The small
mammals from Gratkorn: an overview. Palaeobiodivers.
Palaeoenviron. 94, 1, 135–162.
Ramdohr F.A. 1808: Über die Gattung Cypris Müll. und drei zu
derselben gehörige neue Arten. Mag. neuesten Entd. Naturkd.
Berlin 2, 83–93.
49
LIFE IN THE FLUVIAL HINTERLAND OF THE LATE SARMATIAN SEA IN THE STYRIAN BASIN (AUSTRIA)
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Reichenbacher B. 1989: Feinstratigraphische Gliederung der Kirch-
berger Schichten (Unter-Miozän) an der Typuslokalität
Illerkirchberg bei Ulm. Geol. Bavarica 94, 135–177.
Reischenbacher D., Rifelj H., Sachsenhofer R.F., Jelen B., Coric S.,
Gross M. & Reichenbacher B. 2007: Early Badenian paleo-
environment in the Lavanttal Basin (Mühldorf Formation):
Evidence from geochemistry and paleontology. Austrian J.
Earth Sci. 100, 202–229.
Reuse C. 1983: On the taxonomic significance of the internal shell in
the identification of European slugs of the families Limacidae
and Milacidae (Gastropoda, Pulmonata). Biol. Jaarb. Dodonaea
41, 180–200.
Sacco F. 1886: Neue Arten von terrestrischen Mollusken des Tertiärs
aus den Süß- und Brackwassern des Piemonts [Nuove specie ter-
ziarie di molluschi terrestri, l’acqua dolce e salmastra del
Piemonte]. Atti soc. ital. sci. nat. 29, 427–476 (in Italian).
Salvador R.B. 2013: The fossil pulmonate snails of Sandelzhausen
(Early/Middle Miocene, Germany): Ellobiidae, Pupilloidea, and
Clausilioidea. Paläont. Z. 89, 37–50.
Salvador R.B. & Rasser M.W. 2014: The fossil pulmonate snails of
Sandelzhausen (Early/Middle Miocene, Germany) (Hygrophila,
Punctoidea and limacoids). Arch. Molluskenkd. 143, 187–202.
Salvador R.B. & Rasser M.W. 2016: The fossil land and freshwater
snails of Oggenhausen (Middle Miocene, Germany). Rev. Bras.
paleont. 19, 41–52.
Salvador R.B., Rasser M.W. & Höltke O. 2015a: Fossil gastropods
from Miocene Lake Randeck Maar and its hinterland (SW
Germany). N. Jahrb. Geol. Paläont. 277, 251–273.
Salvador R.B., Sach V.J. & Valentas-Romera B.L. 2015b: The fossil
continental mollusks in the Upper Freshwater Molasse (Middle
Miocene) of the districts of Biberach, Ravensburg and Neu-Ulm,
Germany. Rev. bras. paleont. 18, 201–216.
Salvador R.B., Prieto J., Mayr C. & Rasser M.W. 2016: New gastro-
pod assemblages from the Early/Middle Miocene of Riedens-
heim and Adelschlag-Fasanerie, southern Germany. N. Jahrb.
Geol. Paläont. 279, 127–154.
Sames B. 2011: Glossary of morphologic terms of late Mesozoic non-
marine Ostracoda, relevant to Theriosynoecum Branson 1936
and Cypridea Bosquet 1852. Micropaleont. 57, 433–454.
Sandberger F. 1858–1863: Die Conchylien des Mainzer Tertiär-
beckens. livr. 1, 1–40, pl.1–5 (1858); livr. 2, 41–72, pl. 6–10
(1858); livr. 3, 73–112, pl. 11–15 (1859); livr. 4, 113–152, pl.
16–20; livr. 5, 153–192, pl. 21–25; livr. 6: 193–232, pl. 26–30;
livr. 7, 233–270, pl. 31–35 (1862); livr. 8, 271–468 (1863), C.W.
Kreidel, Wiesbaden.
Sandberger F. 1870–1875: Die Land- und Süsswasser-Conchylien der
Vorwelt. livr. 1, 1–48, pl. 1–4 (1870); livr. 2–3, 49–96, pl. 5–12
(1870); livr. 4–5, 97–160, pl. 13–20 (1871); livr. 6–8, 161–256,
pl. 21–32 (1872); livr. 9–10, 257–352, pl. 33–36 (1873); livr. 11,
353–616 (1875); livr. 12, 617–1000, (1875), C.W. Kreidel,
Wiesbaden.
Schäfer P. 2005: Beiträge zur Ostracoden- und Foraminiferen-Fauna
der Unteren Süßwassermolasse in der Schweiz und in Savoyen
(Frankreich). 2. La Chaux (Kanton Waadt, Schweiz). Sencken
bergiana leth. 85, 1, 95–117.
Schäfer P. 2011: Beiträge zur Ostracoden- und Foraminiferen-Fauna
der Unteren Süßwassermolasse in der Schweiz und in Savoyen
(Frankreich). 3. Das Findreuse-Profil (Départment Haute-
Savoie, Frankreich). Zitteliana A. 51, 255–283.
Schlickum W.R. 1964: Die Molluskenfauna der Süßbrackwasser-
molasse Niederbayerns. Arch. Molluskenkd. 93, 1–70.
Schlickum W.R. 1976: Die in der pleistozänen Gemeindekiesgrube
von Zwiefaltendorf a. d. Donau abgelagerte Molluskenfauna der
Silvanaschichten. Arch. Molluskenkd. 107, 1–31.
Schlickum W.R. 1978: Zur oberpannonen Molluskenfauna von Öcs,
I. Arch. Molluskenkd. 108, 245–261.
Schlosser M. 1907: Die Land- und Süsswassergastropoden vom
Eichkogel bei Mödling. Nebst einer Besprechung der Gastro-
poden aus dem Miocän von Rein in Steiermark. Jahrb. K. K.
Geol. Reichsanst. 57, 753–792.
Schlotheim E.F. 1820: Die Petrefaktenkunde auf ihrem jetzigen
Standpunkte durch die Beschreibung seiner Sammlung verstei-
nerter und fossiler Ueberreste des Thier- und Pflanzenreichs der
Vorwelt erläutert. Becker, Gotha, 1–438.
Schnabel T. 2006: Die känozoischen Filholiidae Wenz 1923.
Teil 3: Die miozänen Vertreter der Gattung Triptychia. (Gastrop-
oda, Pulmonata, Clausilioidea). Arch. Molluskenkd. 135,
133–203.
Schnabel T. 2007: The Cenozoic Filholiidae Wenz 1923. Part 4:
The eo- and oligocene species of the genus Triptychia, inclu-
ding remarks on ecology, geo- and stratigraphic distribution of
the Filholiidae and evolution of the genus Triptychia (Gastro-
poda, Pulmonata, Clausilioidea). Arch. Molluskenkd. 136,
25–57.
Schreilechner M.G. & Sachsenhofer R.F. 2007: High Resolution
Sequence Stratigraphy in the Eastern Styrian Basin (Miocene,
Austria). Austrian J. Earth Sci. 100, 164–184.
Schultz O. 2001: Catalogus Fossilium Austriae, Band 1/Teil 1
(Bivalvia neogenica). Oesterr. Akad. Wiss. Wien, XLVIII +
1–379.
Schütt H. 1967: Die Landschnecken der untersarmatischen Rissoen-
schichten von Hollabrunn. Arch. Molluskenkd. 96, 199–222.
Sieber E. 1905: Fossile Süßwasser-Ostrakoden aus Würtemberg.
Jahresh. Ver. vaterl. Naturkd. Württ. 61, 321–346.
Sokač A. 1972: Pannonian and Pontian ostracode Fauna of Mt. Med-
vednica. Palaeont. Jugoslav. 11, 1–140.
Straub E.W. 1952: Mikropaläontologische Untersuchungen im Tertiär
zwischen Ehingen und Ulm a. d. Donau. Geol. Jahrb. 66, 433–523.
Strauch F. 1977: Die Entwicklung der europäischen Vertreter der
Gattung Carychium O.F. Müller seit dem Miozän. Arch.
Molluskenkd. 107, 149–193.
Stworzewicz E. 1999a: Miocene land snails from Belchatów (Central
Poland), III: Carychiinae (Gastropoda; Pulmonata: Ellobiidae).
Paläont. Z. 73, 261–276.
Stworzewicz E. 1999b: Miocene land snails from Belchatów (Central
Poland). IV: Pupilloidea (Gastropoda Pulmonata). Systematic,
biostratigraphic and palaeoecological studies. Fol. Malacol. 7, 3,
133–170.
Stworzewicz E., Prisyazhnyuk V.A. & Górka M. 2013: Systematic
and palaeoecological study of Miocene terrestrial gastropods
from Zwierzyniec (South Poland). Ann. Soc. Geol. Pol. 83,
179–200.
Triebel E. 1941: Zur Morphologie und Ökologie der fossilen Ostra-
coden. Mit Beschreibung einiger neuer Gattungen und Arten.
Senckenbergiana 23, 4–6, 294–400.
Troll O.R. von 1907: Die pontischen Ablagerungen von Leobersdorf
und ihre Fauna. Jahrb. Geol. Reichsanst. 57, 33–90.
Welter-Schultes F. 2012: European non-marine molluscs, a guide
for species identification. Planet Poster Edition, Göttingen,
1–760.
Wenz W. 1911: Gonostoma (Klikia) osculum Thom. und ihre Ver-
wandten im mitteleuropäischen Tertiär. Jahrb. Nassau. Ver.
Naturkd. 64, 75–101.
Wenz W. 1914: II. Paläontologischer Teil. Jahrb. Nassau. Ver.
Naturkd. 67, 30–154.
Wenz W. 1923–1930: Fossilium Catalogus I: Animalia. Gastropoda
extramarina tertiaria. 17, 1, 1–352 (1923), 18/2, 353–736
(1923), 20/3, 737–1068 (1923), 21/4, 1069–1420 (1923),
22/5, 1421–1734 (1923), 23/6, 1735–1862 (1923), 32/7,
1863–2230 (1926), 38/8, 2231–2502 (1928), 40/9, 2503–2886
(1929), 43/10, 2887–3014 (1929), 46/11, 3015–3387 (1930),
W. Junk, Berlin.
50
DOUBRAWA, GROSS and HARZHAUSER
GEOLOGICA CARPATHICA
, 2018, 69, 1, 30–50
Wenz W. 1927: Weitere Beiträge zur Fauna der pontischen Schichten
von Leobersdorf. Senckenbergiana 9, 41–48.
Wenz W. & Edelauer A. 1942: Die Molluskenfauna der oberpon-
tischen Süßwassermergel vom Eichkogel bei Mödling, Wien.
Arch. Molluskenkd. 74, 82–98.
Wessels W. & Reumer B.M. 2009: Democricetodon and Mega
cricetodon (Mammalia, Cricetidae) from the Miocene of
Sandelzhausen, Germany. Paläont. Z. 83, 187–205.
Wiktor A. & Likharev M. 1979: Phylogenetische Probleme bei
Nacktschnecken aus den Familien Limacidae und Milacidae
(Gastropoda, Pulmonata). Malacologia 18, 123–131.
Winkler A. 1927: Über die sarmatischen und pontischen Ablagerun-
gen im Südostteil des steirischen Beckens. Jahrb. Geol. Bundes
anstalt 77, 393–456.
Witt W. 1998: Die miozäne Fossil-Lagerstätte Sandelzhausen. 14.
Ostracoden. Mitt. Bayer
. Staatssamml. Paläonotol. Hist. Geol.
38, 135–165.
Witt W. 2000: Süßwasserostracoden der miozänen Vorlandmolasse
Süddeutschlands. Mitt. Bayer. Staatssamml. Paläonotol. Hist.
Geol. 40, 109–151.
Witt W. 2002: Zur Süßwasserostracodenfauna der oligo-miozänen
Vorlandmolasse Süddeutschlands. Mitt. Bayer. Staatssamml.
Paläonotol. Hist. Geol. 42, 35–49.
Witt W. 2011: Mixed ostracod faunas, co-occurrence of marine Oligo-
cene and non-marine Miocene taxa at Pınarhisar, Thrace, Turkey.
Zitteliana A. 51, 237–254.
von Zieten C.-H. 1830–1832: Die Versteinerungen Württembergs.
I–VIII, 1–16, 1–12 pl. (1830); 17–32, 13–24 pl. (1831), 33–64,
25–48 pl. (1832), 65–102, 49–72 pl. (1833), Verlag and Litho
graphie der Expedition des Werkes unserer Zeit, Stuttgart.