191
MARINE AND TERRESTRIAL VERTEBRATES FROM THE MIDDLE MIOCENE OF GRUND
GEOLOGICA CARPATHICA, 55, 2, BRATISLAVA, APRIL 2004
191197
MARINE AND TERRESTRIAL VERTEBRATES FROM THE MIDDLE
MIOCENE OF GRUND (LOWER AUSTRIA)
GUDRUN DAXNER-HÖCK
1
, PETRA M. MIKLAS-TEMPFER
1
, URSULA B. GÖHLICH
2
,
KATI HUTTUNEN
3
, EMESE KAZÁR
4
, DORIS NAGEL
5
, GERTRUD E. ROESSNER
2
,
ORTWIN SCHULTZ
1
and REINHARD ZIEGLER
6
1
Museum of Natural History Vienna, Burgring 7, A-1014 Vienna, Austria; gudrun.hoeck@nhm-wien.ac.at;
ortwin.schultz@nhm-wien.ac.at; petra.tempfer@nhm-wien.ac.at
2
Department für Geo- und Umweltwissenschaften, Sektion Paläontologie, Richard-Wagner-Str. 10, D-80333 München, Germany;
u.goehlich@lrz.uni-muenchen.de; u.goehlich@web.de; g.roessner@lrz.uni-muenchen.de
3
Museum of Natural History Vienna, Burgring 7, A-1014 Vienna, Austria; Kati_jh@yahoo.de
4
Geological Institute of Hungary, Stefánia út 14, H-1143 Budapest, Hungary; kazar@mafi.hu
5
Department of Paleontology, University of Vienna, Geozentrum, Althanstrasse 14, A-1090 Vienna; doris.nagel@univie.ac.at
6
Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, D-7000 Stuttgart; r.ziegler.smns@naturkundemuseum-bw.de
(Manuscript received June 5, 2003; accepted in revised form December 16, 2003)
Abstract: The Grund assemblage comprises 45 vertebrate species from marine and terrestrial ecosystems. A whale,
sharks, rays, bony fishes, and sea-birds hint at marine conditions, whereas a salamander, a pheasant bird, turtles, snakes
and land-mammals indicate various terrestrial paleoenvironments. Biostratigraphically the upper part of MN5 (mam-
mals) and the upper part of the Lower Lagenidae Zone (foraminifers) are indicated. The assemblage of Grund therefore
serves as a reference point for the continental/marine correlation in the Central Paratethys.
Key words: Badenian, Central Paratethys, fishes, amphibians, reptiles, birds, mammals.
Introduction
In Grund near Hollabrunn in Lower Austria, extended excava-
tions were carried out by the University of Viennas Institute
of Paleontology in the field seasons 19982000.
diverse in species (45). The scarce, disarticulated terrestrial
fossils are strongly eroded. They hint at long-distance trans-
port from the hinterland, at reworking and dislocation, and at
a final deposition in channels filled with sand and molluscan
debris.
Fig. 1. Geographical and geological position of the locality Grund (slightly modified from Roetzel
& Pervesler 2004).
The geological setting of the type
area of the Grund Formation and the
sedimentology of the Grund sec-
tions are investigated and discussed
in detail by Roetzel & Pervesler
(2004) (Fig. 1). The sediments are
dominated by yellowish fine sands
and silts; the channels are filled with
coarser sand, fine gravel, and with
molluscan debris, occasionally inter-
mixed with vertebrate remains. For
localization of the vertebrate-bear-
ing layers in the sections (GRU-B1-1,
GRU-B1-3, GRU-F-11) see Roetzel
& Pervesler (2004: Fig. 4ad).
In addition to the excavations,
three large bulk samples were
washed by members of the Natural
History Museum Vienna and by stu-
dents, focusing on smaller molluscs
and vertebrates. For wet screening
sieves with mesh sizes of 0.5, 2.5
and 5.0 mm were used. These inves-
tigations yielded a vertebrate as-
semblage that is expectedly poor in
specimens (180) but surprisingly
192 DAXNER-HÖCK et al.
Table 1: Middle Miocene vertebrates from the locality Grund (sections GRU-B1 and GRU-F) and Guntersdorf. The specimens are housed in
the collections: Natural History Museum Vienna, Geol.-Paleontol. Department (Nat. Hist. Mus. Vienna), Institute of Paleontology, Universi-
ty of Vienna
1)
, Niederösterr. Landesmuseum
2)
, H. Schaffer
3)
, Geological Survey Vienna
4)
,
A. Kroh
5)
.
Terrestrial and marine vertebrates
The vertebrate fossil from two localities of the Grund Beds
in the Molasse Basin of Lower Austria, that is the fossil sites
of Mühlbach and Grund, were recently investigated and pub-
lished in a special volume (104 A) of the Ann. Nat. Hist. Mus.
Wien (2003). These two seemingly unspectacular assemblag-
es merit special attention because terrestrial and marine or-
ganisms occur together in isochronous marine sediments.
They not only provide insight into various ecosystems, but
also allow a correlation to be established between continental
and marine biozonations.
The present paper summarizes our present knowledge of
the vertebrates from the Grund sections, from some wine cel-
lars in Grund, and from the neighbouring village of Gunters-
dorf (Table 1). A total of 45 taxa were described and figured in
the above-mentioned special volume (104 A): 19 fishes
(Schultz 2003), 1 amphibian and 6 reptiles (Miklas-Tempfer
2003), 4 birds (Göhlich 2003), 1 insectivore (Ziegler 2003), 2
rodents (Daxner-Höck 2003), 4 carnivores (Nagel 2003), 5 ru-
minants (det. Roessner), 2 proboscideans (Huttunen 2003) and
1 whale (det. Kazár).
Pisces Schultz (2003): The fish fauna is poor, even
though the Grund area has been sampled for over 150 years.
The total yield is 19 marine fish-taxa: 8 sharks/Squalomorphii,
6 rays/Batomorphii and 5 bony fishes/Osteichthyes. The small-
sized sharks as well as the rays and the bony fishes indicate a
shallow sea. The habitat can be classified as neritic. The shark
G
R
U
-B
1-
1 (
1998)
G
R
U
-B
1-
3 (
1998)
G
R
U
-F
-11 (
1999)
N
at
. H
is
t. M
us
.
Vi
enna
Uni
Vi
enna
1)
N
Ö
L
an
d. M
us
.
2)
Co
ll.
SCHAF
FE
R
3)
Ge
ol.
Sur
v.
4)
GRU-KROH
5)
Pisces (Schultz 2003)
Squalomorphii indet.
2 vertebrae
Notorynchus primigenius (Agassiz, 1843)
2 teeth
Carcharias acutissimus (Agassiz, 1844)
9 teeth
22+3? teeth
Carcharias cuspidatus (Agassiz, 1844)
9 teeth
Carcharias sp.
2 teeth
Scyliorhinus distans (Probst, 1879)
1 tooth
Carcharhinus priscus (Agassiz, 1843)
1? tooth
1+2? teeth
11 teeth
Galeocerdo aduncus Agassiz, 1843
2 teeth
Sphyrna? sp.
1 tooth
Dasyatis cf. rugosa (Probst, 1877)
1 tooth
1 tooth
Dasyatis sp.
1 tooth
1 tooth
Myliobatis sp.
1 toothplate
Rhinoptera sp.
1 toothplate
Myliobatis/Rhinoptera sp.
1 toothplate
2 toothplates
5 toothplates
Aetobatus arcuatus Agassiz, 1843
1 toothplate
2 toothplates
Myliobatidae/Dasyatidae
2 tail spines
1 tail spine
Diplodus incisivus (Gervais, 1852)
2 teeth
Sparus umbonatus (Münster, 1846)
6 teeth
10 teeth
Pagrus cinctus (Agassiz, 1836)
1 tooth
1+?1 teeth
Sparidae indet.
13 teeth
10 teeth
Sphyraena sp.
1 tooth
Amphibia (Miklas-Tempfer 2003)
Salamandra sansaniensis Lartet, 1851
1 vertebra
Reptilia (Miklas-Tempfer 2003)
Testudo sp.
1 humerus
2)
1 shellfrag.
1)
shellfrag.
3)
Ptychogaster grundensis Bachmayer et
Schaffer, 1959
2 shellfrag.
cast of
Holotypus
1 carapax
(Holotypus)
3)
1 shellfrag.
Trionyx vindobonensis Peters, 1855
shellfrag.
3)
Anguidae indet.
1 vertebra
Elaphe kohfidischi Bachmayer et Szyndlar,
1985
1 vertebra
1 vertebra
2 vertebrae
Naja romani Hofstetter, 1939
1 vertebra
3 vertebrae
1 vertebra
193
MARINE AND TERRESTRIAL VERTEBRATES FROM THE MIDDLE MIOCENE OF GRUND
Table 1: Continued
Sphyrna and the rays Rhinoptera and Aetobatis indicate tropi-
cal to subtropical conditions. Living relatives of all remaining
taxa inhabit tropical to temperate waters.
Amphibia and Reptilia Miklas-Tempfer (2003: Figs. 2
14; Plates 15): The herpetofauna points to a wooded
paleoenvironment and partly to some standing or slowly flow-
ing freshwater habitats. The reptiles are: 3 tortoises
(Ptychogaster grundensis, Testudo sp. and Trionyx vindobo-
nensis), 1 species of Anguidae indet. and 2 snakes (Elaphe
kohfidischi and Naja romani). The amphibians are represented
by only one member of the salamanders, Salamandra
sansaniensis. Its living relative is Salamandra salamandra,
which prefers hilly or mountainous wooded habitats up to
2000 m. This terrestrial salamander needs damp environments
with bushes and deciduous woods, but can also be found in
semiarid environments. While modern herbivorous Testu-
dinidae live in dry environments, members of the fossil turtle
Ptychogaster were terrestrial-aquatic animals inhabiting
bushes and woods near water bodies. Terrapins of the genus
Trionyx require standing or slowly flowing, shallow freshwa-
ters with some sandy ground for burrowing. The ecological re-
quirements of living Anguidae span from dry (e.g. Pseudopus
apodus) to moist (e.g. Anguis fragilis). The snakes from
Grund definitely point to ecologically dry conditions, such as
G
R
U
-B
1-
1 (
1998)
G
R
U
-B
1-
3 (
1998)
G
R
U
-F
-11 (
1999)
N
at
. H
is
t. M
us
.
Vi
enna
Uni
Vi
enna
1)
N
Ö
L
an
d. M
us
.
2)
Co
ll.
S
CHAF
FE
R
3)
Ge
ol.
Sur
v.
4)
GRU-KROH
5)
Aves (Göhlich 2003)
Microsula pygmaea (Milne-Edwards, 1874)
1 femur
1 humerus
Phalacrocorax intermedius (Milne-Edwards, 1867) 1 carpometa-carpus
cf. Palaeortyx intermedia Ballmann, 1969
1 coracoid
Laridae indet.
1 ulna
Aves indet.
1 carpometa-
carpus
Insectivora (Ziegler 2003)
Schizogalerix pristinus Ziegler, 2003
1 M2 sin.
Rodentia (Daxner-Höck 2003)
Cricetodon meini Freudenthal, 1963
1 maxilla
1 jaw
1 M1 sin.
Democricetodon mutilus Fahlbusch, 1964
1 M2 dext.
Carnivora (Nagel 2003)
Semigenetta sansaniensis (Lartet, 1851)
maxilla P3-4
Caniformia gen. et spec. indet.
1 phalange
Mustelidae gen. et spec. indet.
1 caput femoris
Carnivora indet.
1 vertebraefrag.
Ruminantia (det. Rössner)
Ruminantia indet.
1 atlas
1 frag. vert.
1 intermediale
1 m inf.
1 humerus-
frag.
Pecora indet.
1 ulna prox.
2 phalanges
2 phalanges
Micromeryx sp.
1 M2 dext.
Micromeryx aff. flourensianus Lartet, 1851
1 M1 sin.
1 M2 dext.
1 m1 sin.
1 P4 dext.
Palaeomeryx eminens von Meyer, 1847
1 p4 sin.
4)
Proboscidea (Huttunen 2003)
Gomphotherium angustidens Cuvier, 1817
1 incisorfrag.*
1 maxillafrag.*
Prodeinotherium bavaricum von Meyer, 1841
1 d4 sin.
Proboscidea indet.
1 sesamoid
Cetacea (det. Kazár)
Mysticeti indet.
1 mandiblefrag.
*deciduous dental material from Guntersdorf near Grund
194 DAXNER-HÖCK et al.
Table 2: Stratigraphy and correlation of Early to Middle Miocene vertebrate faunas from Central Europe (slightly modified from Dax-
ner-Höck 2003).
in the case of Elaphe kohfidischi, which is a relative to the liv-
ing Aesculapian snake. As a member of the Colubrinae it pre-
fers dry and wooded environments, even without permanent
water nearby. The living Aesculapian snake is a good climber
on trees and bushes. Members of the genus Naja, a poisonous
snake, are restricted to tropical areas today; they inhabit dry
places and spend the day concealed in the vegetation and
crevices.
Aves Göhlich (2003: Plate1): The avifauna contains 4
taxa of different ecological habitats: a pheasant, a boobie, a
cormorant and a gull. An additional carpometacarpus from an
as yet undetermined bird was also found. Whereas the pheas-
ant (cf. Palaeortyx intermedia) is a terrestrial bird, the boobie
(Microsula pygmaea), the cormorant (Phalacrocorax interme-
dius) and the gull (Laridae indet.) are aquatic taxa, most of
them probably marine. Modern boobies are sea-birds known
worldwide from pelagic to coastal environments. The piscivo-
rous cormorants are also cosmopolitan in marine and freshwa-
ter environments, preferring temperate to tropical waters of
coasts, lakes, open swamps and slow-flowing rivers. Extant
gulls (Laridae) are typically sea-birds, but are also found in in-
land waters. Phalacrocorax intermedius was previously
known only from its Early Miocene type locality (Faluns de
lOrléanais, France) and is herewith recorded for the first time
in the Middle Miocene (MN5).
Insectivora Ziegler (2003: Tafel 12) and Rodentia
Daxner-Höck (2003: Plates 13): Small mammal fossils are
extremely rare in the Grund fauna. The hedgehog Schizo-
galerix pristinus, the small hamster Democricetodon mutilus
and the large hamster Cricetodon meini are represented by
one molar each. Furthermore, one maxilla and a mandible
without teeth can be assigned to the large hamster Cricetodon.
These few small mammal fossils are identical to those of the
hedgehog S. pristinus and the hamsters C. meini and D. muti-
lus, that is the most numerous mammals from Mühlbach.
They are not only the dominating mammal species of the
Mühlbach assemblage, but also highly relevant for strati-
graphic interpretations. As a burrowing rodent, Cricetodon
points to rather dry woodland and a low groundwater level;
Schizogalerix is also indicative for a woodland habitat but
hints at close-by freshwater habitats.
S. pristinus is the most primitive species of Schizogalerix,
and Mühlbach is the earliest record of the genus in Central
Europe. The large hamster Cricetodon migrated from Anato-
lia to SE-Europe in the Early Miocene (MN4). The first record
of C. meini is Komotini in Greece (MN5). It reached Central
and Western Europe and ultimately became extinct at the end
of the Mammal Zone MN5. In Central Europe, C. meini marks
a short time interval in the upper part of MN5. This is shown
by several occurrences immediately below the Brock-hori-
195
MARINE AND TERRESTRIAL VERTEBRATES FROM THE MIDDLE MIOCENE OF GRUND
zon (indicating the Ries impact at 14.9 Ma) of the Upper
Freshwater Molasse in Germany (Table 2). Above the
Brock-horizon, C. meini was no longer present, but was re-
placed by its larger relative C. aff. aureus.
Carnivora Nagel (2003: Plate 1): The carnivore fossils
from Grund are: a maxilla with P4, a caput femori, a phalanx
and some postcranial remains. Due to the size and the charac-
teristic premolar, the maxilla could be identified as Semige-
netta sansaniensis. It is the first record of this species in Aus-
tria. The caput femori is probably from a mustelid, while the
phalanx could fit into the morphology of an amphicyonid.
Some scarce postcranial remains can only be identified as
Carnivora indet.
Ruminantia det. Roessner: The investigated material
from Grund sections comprises six isolated teeth and eight
postcranial elements. While a species determination of the
Table 3: Early to Late Miocene geochronology and biostratigraphy (Harzhauser et al. 2003). The shaded area indicates the stratigraphic
position of vertebrate-bearing samples of the Grund sections.
teeth is possible, for the time being the postcranials and a
strongly worn lower molar were assigned to Ruminantia in-
det. and to Pecora indet., respectively. Micromeryx aff. flou-
rensianus is indicated by 4 isolated teeth. One M2 of Mi-
cromeryx sp. is slightly larger than the former. One P4 of
Palaeomeryx eminens is listed in Pia & Sickenberg (1934).
Detailed investigations on the ruminants are in progress.
Proboscidea Huttunen (2003: Plate 1): Gomphotherium
angustidens is represented by a fragmentary maxilla with left
and right deciduous teeth (D2-4), permanent premolars (P2-3)
and the M1, and by an isolated deciduous incisor. This excep-
tional finding from the locality of Guntersdorf near Grund
was investigated for the first time by Schlesinger (1917: 17
22; Plate1: Fig. 2a; Plate 2: Fig. 1). The present study yielded
new details of deciduous and permanent tooth morphology
and the existence of a permanent P2 of G. angustidens. The
196 DAXNER-HÖCK et al.
second proboscidean, Prodeinotherium bavaricum, was iden-
tified on the basis of a deciduous lower d4 from the locality
Grund. Furthermore, a sesamoid from Grund most likely be-
longs to a small-sized proboscidean.
Cetacea det. Kazár: A mandible fragment of a baleen
whale from Grund shows marked similarities to lower jaws of
the genus Mesocetus, an insufficiently known Middle Mio-
cene cetothere (Cetacea: Mysticeti: Cetotheriidae) reported
from several localities of the Carpathian and Vienna Basins
(Van Beneden 1884; Kadiæ 1904; Kellogg 1925; Pia 1937).
Due to the lack of diagnostic cranial features, however, the
fossil cetacean remains from Grund can only be referred to as
Mysticeti indet.
Middle Miocene species classified as cetotheres were ba-
leen-bearing, filter-feeding mysticetes structurally close to
present-day balaenopterids (Fordyce & Muizon 2001). The
most influential factor in baleen whale distribution is food
supply: large whales today depend on dense patches of zoop-
lankton or fish swarms (Gaskin 1976). Although the mandible
fragment from Grund belonged to a small individual (estimat-
ed mandible length: 100120 cm, and thus the estimated total
body length falls well below the size of the smallest recent ba-
leen whale, the minke whale, Balaenoptera acutorostrata),
the presence of a mysticete in the Grund fauna indicates a
high-productivity environment. Living baleen whales are dis-
tributed worldwide in pelagic marine environments and it is
possible that Middle Miocene forms favoured similar habitats.
Paleoecology, stratigraphy and correlations
Paleoecology
The Grund assemblage combines elements of marine and
terrestrial ecosystems. The shallow sea was inhabited by dif-
ferent sharks, rays and bony fishes. The presence of a baleen
whale (Mysticeti) points to abundant planktonic organisms
and/or fish swarms. Fishes were the food supply for the sea-
birds, the cormorant (Phalacrocorax) and the boobie (Micro-
sula), while the gull (Laridae) might have fed on eggs, on
small birds and on the fish-prey that was lost by cormorants
and boobies. The amphibians, reptiles, the pheasant bird and
the land mammals represent different terrestrial environments.
More dry areas along the coast and on land slopes covered
with dense bushes, shrubs and smaller trees were inhabited
by the tortoises (Testudo), the snakes (Elaphe and Naja), the
pheasant bird (cf. Palaeortyx), and by the hamsters (Crice-
todon and Democricetodon). On the other hand, the sala-
mander (Salamandra), the terrapin (Trionyx), the semi-aquat-
ic turtle (Ptychogaster), and the hedgehog (Schizogalerix)
preferred wooded environments close to standing or flowing
freshwater. The musk deer-like (Micromeryx) and the large
ruminant (Palaeomeryx) most probably inhabited swampy
forests and floodplains along rivers, as did the proboscideans
(Gomphotherium and Prodeinotherium). All these herbivo-
rous mammals, birds and the reptiles were potential prey of
the carnivore-omnivore hunters, that is the viverrid (Semige-
netta) and the amphicyonid and mustelid carnivores.
Stratigraphy and correlation
According to Daxner-Höck (2003) the terrestrial vertebrate
assemblages with Cricetodon meini are representative of the
late MN5 and are definitely older than the Ries event, which
is dated at ~14.9 million years (Table 2). The marine fauna of
the Grund Formation corresponds to the foraminiferal Zone
Lower Lagenidae Zone. It is interpreted by Rögl et al. (2002)
and Rögl & Spezzaferri (2003) to span the upper part of the
planktonic foraminiferal Zone M5b (indicated by Praeorbuli-
na glomerosa circularis) to Zone M6 (FAD of Orbulina su-
turalis) at 15.1 million years (Table 3). The co-occurrence
with rodents, indicating the late MN5, agrees with this dating.
Therefore, the fauna from Grund serves as a reference tie
point for marine/continental correlation in the Central Para-
tethys.
Acknowledgments: The fossils were collected during field
investigations by the Institute of Paleontology of the Universi-
ty of Vienna, under the leadership of P. Pervesler and R. Roetzel,
and supported by students and preparators. M. Harzhauser
helped with the computer graphics. M. Stachowitsch correct-
ed the English. The scientific investigations were funded by
the FWF-Project P-15724-N06. All these institutions and per-
sons are sincerely acknowledged.
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