GEOLOGICA CARPATHICA, AUGUST 2005, 56, 4, 337345
Ruminantia from the Grund Formation: correlation of
continental and marine deposits and climatic development
during early Middle Miocene of the Central Paratethys
GERTRUD E. RÖSSNER
Department für Geo- und Umweltwissenschaften, Sektion Paläontologie, und GeoBio-Center der Ludwig-Maximilians-Universität
München, Richard-Wagner-Str. 10, D-80333 München, Germany; email@example.com
(Manuscript received March 29, 2004; accepted in revised form December 9, 2004)
Abstract: The Badenian marine littoral Grund Formation at Grund in the Molasse Zone of Lower Austria provided
marine and terrestrial faunas in the same strata. They allowed correlation between continental and marine biozonations
of the European Middle Miocene. In the present paper for the first time the included fossil remains of Ruminantia are
described in detail. Six isolated teeth and nine postcranial skeletal elements are determined as Ruminantia indet., Pecora
indet., cf. Micromeryx flourensianus Lartet, 1851, Micromeryx cf. styriacus Thenius, 1950, and cf. Eotragus clavatus
(Gervais, 1850). The biochronological age indicated by cf. M. flourensianus, M. cf. styriacus and cf. E. clavatus is
restricted to the European mammal zones late MN5 to MN6. Consequently the hitherto identified age of late MN5 for
these sediments is confirmed by the occurrence of the Ruminantia represented. The taxonomic composition further
indicates paleoecological conditions compatible with earlier reconstructions of the environment as either islands or the
elevated parts of a peninsula or rising terrain of the hinterland. In any of these settings groundwater levels would have
been low and the vegetation a covering of woods, bushes and shrubs. Comparisons with Austrian ruminant faunas of the
early MN5/late Karpatian (Obergänserndorf and Teiritzberg) support the interpretation of a change during MN5 towards
a drier climate under more or less stable temperatures in Central Europe.
Key words: Middle Miocene, Lower Badenian, Molasse Zone, Grund Formation, paleoecology, biostratigraphy, taxonomy,
dry wooded habitat, Ruminantia.
Excavations in the type area of the Lower Badenian marine
littoral Grund Formation at Grund near Hollabrunn in Lower
Austria (Fig. 1) by the Institute of Paleontology of Vienna
University from 1998 to 2000 provided a vertebrate assem-
blage poor in specimens but diverse in species (Roetzel 2003;
Daxner-Höck et al. 2004). Its outstanding character is the syn-
occurrence of marine and terrestrial vertebrates in isochronous
sediments. Biostratigraphically the Grund-Formation belongs
to the regional Lower Lagenid Zone (Lower Badenian) based
on the evolutionary level of Praeorbulina glomerosa circu-
laris and therewith to the top of the Planktonic Foraminifera
Zone M5b/Mt5b (Rögl & Spezzaferri 2003) (Fig. 2). The co-
occurrence of the nannofossils Sphenolithus heteromorphus
and Helicosphaera waltrans in the section Mühlbach of the
Gaindorf Formation (Æoriæ 2003), which is a lateral equivalent
(Roetzel 2003) indicates the Calcareous Nannoplankton Zone
NN5 (Harzhauser et al. 2002) (Fig. 2). According to Daxner-
Höck (2003) the terrestrial vertebrate assemblages are repre-
sentative for the later part of the European Land Mammal
Zone MN5. A major part of the vertebrate remains have been
investigated and are already published in a special volume of
Annalen des Naturhistorischen Museums in Wien, 104/A.
The remains of Ruminantia and a preliminary determination
were hitherto only briefly mentioned in Daxner-Höck et al.
(2004). In the present paper the ruminant specimens from
Grund are described in detail, determined as exactly as possi-
ble, and assessed biochronologically.
Material and methods
The available material of Ruminantia from Grund comprises
six isolated teeth and nine postcranial specimens. Determina-
tion at species level can be attempted for the teeth, based on
extensive and reliably determined material from different Eu-
ropean localities. Comparisons of the postcranial specimens
are difficult or even impossible, because postcranials have
been described more rarely than dentitions and because reli-
able species determination of postcranials in taxonomically di-
verse collections have not been done. Therefore the ruminant
postcranials from Grund are described and compared within
these limitations, mainly based on material housed in the
Bavarian State Collection of Paleontology and Geology Mu-
nich, and are determined only at a higher systematic level. The
Grund fossils are housed in the collection of the Geological-
Paleontological Department, Museum of Natural History,
All measurements are given in mm. Manner of measure-
ment in teeth and tooth crown terminology is according to
Abbreviations: apd largest anterior-posterior diameter
distal; app largest anterior-posterior diameter proximal;
aw anterior width; l largest length; pw posterior
width; td largest transversal diameter distal; tp largest
transversal diameter proximal; w largest width.
Suborder Ruminantia Scopoli, 1777
Ruminantia indet. 1
Material: NHMW GRU-B/-1 13, atlas (l = 26.8; td = min.
42.3); Fig. 7A.
Description: Nearly completely preserved atlas of a
small ruminant. The most lateral and most delicate parts of the
wings are missing and the specimen is slightly corroded. The
Fig. 1. Geological sketch of the Alpine-Carpathian Foredeep in
northeastern Austria, and the position (asterisk) of the investigated
site (taken from Harzhauser et al. 2003).
shape is trapezoidal. The dorsocaudal Tuberculum dorsale is
blunt. Dorsally laterocranially the Foramen alare and Foramen
vertebrale laterale are close together and well rounded. There
is no Foramen transversarium developed.
Discussion: The absence of a Foramen transversarium
clearly indicates a member of the Ruminantia. A more precise
determination is impossible.
Ruminantia indet. 2
Material: NHMW GRU-B/-1 15, left intermediale
(l = 11.8; td = 14.5); Fig. 7D.
Description: The over all specimen is slightly corroded.
The proximal articular surface extends far palmarly. It is dif-
ferentiated into a large volar and a small palmar part, which
consist both of a convex dorsal and a concave palmar mor-
phology. The distal articular surface consists of a dorsal con-
vex and a volar concave part. At the convex surface a blunt
median crest separates the articular surfaces for Carpale 3 and
Carpale 4. Generally the lateral and medial articular outlines
are concave. The size is comparably small.
Discussion: All characters indicate Ruminantia. A more
precise determination is impossible.
Infraorder Pecora Linné, 1758
Pecora indet. 1
Material: NHMW GRU-F 11, proximal phalange III or
IV (l = 33.7; tp = 11.6; app = 13.4; td = 7.3; apd = 7.0); Fig. 7E.
Description: The small to medium sized phalange is ro-
bust. The surface of the internal side is very rough and marks
the insertion of a strong interdigital ligament. The dorsal pro-
file line is convex. The transversal diameter is relatively short
and produces a slender shape. Respectively the proximal artic-
ular surface is slender with a wide incision for the metapodial
Verticillus running from plantar/palmar to dorsal; interdigital
as well as exterior part have the same width. Proximointerdigi-
tally a shallow ligament groove for the proximal interdigital
ligament is developed.
Interdigital and exterior part of the distal articular surface
are of the same size. The interdigital part extends a little more
dorsally than the exterior part. The median groove runs slight-
ly from exterior to interdigital. The lateral outline of the pulley
is well rounded volarly/plantarly and distally, but flat dorsally.
The exterior ligament groove is shallow and rounded, the in-
terdigital one is deeper and elongated.
Discussion: The morphology differs clearly from tra-
gulids in which the incision for the metapodial Verticillus
does not reach the dorsal edge as well as the pulley is relative-
ly shorter and wider and has a different shape of the articular
surface. Within higher pecoran groups Cervidae show the
most similar morphotype (see Köhler 1993). The exterior part
of the proximal articular surface clearly extends more proxi-
mally than in the small to medium sized Cervidae Procervulus
dichotomus (Rauscheröd, Germany, MN4, Rössner 1995) and
Heteroprox (Sandelzhausen, Germany, MN5, Rössner 1997,
2002, 2004). The distal articular surface is distally less wide,
possibly a result of corrosion but this is not certain. For the
RUMINATIA FROM THE GRUND FORMATION OF THE MIDDLE MIOCENE (AUSTRIA) 339
Fig. 2. Early to Late Miocene geochronology and biostratigraphy (taken from Daxner-Höck 2004 after Harzhauser et al. 2003). The shad-
ed area indicates the stratigraphic position of vertebrate-bearing samples from the Grund Formation.
time being the most precise determination possible is Pecora
Pecora indet. 2
Material: NHMW GRU-F 12, distal fragment of proxi-
mal phalange (tp = 14.4; app = 11.9); Fig. 3.
The interdigital part of the distal articular surface is wider
than the exterior part and extends more distally and dorsally.
The facet of the distal articulation is well visible from the dor-
sal aspect. The outline of the pulley is flattened from the distal
and the volar aspect. The exterior ligament groove is shallow
and rounded, the interdigital one is deeper and elongated. The
dorsal transversal diameter of the pulley is wider than in speci-
men GRU-F 11.
Discussion: In tragulids the pulley is proximally short-
er. The size is similar to Heteroprox n. sp. from Sandelz-
hausen (Germany, MN5, Rössner 1997, 2002, 2004), but the
specimen is too incomplete to allow a more detailed compari-
son. Hence within Pecora a precise determination is not yet
Pecora indet. 3
Material: NHMW GRU-B/-1 10, proximal fragment of
right ulna; Fig. 7B.
Description: The specimen is slightly corroded and be-
longed to a small ruminant. The proximal Tuber olecrani is ab-
sent. It is not possible to decide, if this either might indicate an
immature stage or simply an incomplete preservation. Cranial-
ly the articular area is characterized by a prominent proximo-
cranial ridge, by a shallow lateral groove in the middle part,
by a prominent laterodistal appendage, and a small non-prom-
inent mediodistal articular surface.
Discussion: The morphology differs by a less prominent
laterodistal appendage from small tragulids.
Pecora indet. 4
Material: NHMW GRU-B/-1 14 proximal fragment of
proximal phalange, (td = 7.7; apd = 6.8); Fig. 7C.
Description: The specimen is slightly corroded and me-
dium sized. The transversal diameter is comparably small and
produces a slender proximal articular surface. The incision for
the metapodial Verticillus is shallow and wide. The interdigi-
tal part of the articular surface is slender and shows the stron-
gest palmo/plantointerdigital extension of the whole articula-
tion area. Laterally no articulation surface is developed.
Laterodorsally a ligament groove is present. This proximal
phalange differs clearly from specimen GRU-F 11, Pecora in-
det. 1, in morphology and a little larger size.
Discussion: The subdivided articular surface is typical
for pecorans in contrast to the tragulids with an incompletely
divided articular surface. The lack of marginal articular surfac-
es is untypical for moschids, cervids, bovids and palaeomeryc-
ids. This might indicate a phalange of a side digit (II or V),
what further would indicate a large individual, considered the
medium size of the specimen and consequently larger phalan-
ges of ray III or IV. For the time being the specimen can only
be set in Pecora indet.
Fig. 3. NHMW GRU-F 12, Pecora indet. 2, distal fragment of
proximal phalange. A external view, B internal view, C dis-
tal view, D dorsal view. Scale bar 5 mm.
Family Moschidae Gray, 1821
Genus Micromeryx Lartet, 1851
cf. Micromeryx flourensianus Lartet, 1851
Material: NHMW GRU-F 1, sin M1 (l = 6.1; aw = 6.8;
pw = 6.8); Fig. 6A. NHMW GRU-F 2, dex M2 (l = 6.4;
aw = 7.0; pw = 6.8); Fig. 6B. NHMW GRU-F 3, sin m1 (l = ;
aw = ; pw = 4.3); Fig. 6C. NHMW GRU-F 4, dex P4 (l = 4.2;
w = 5.2); Fig. 6D.
Description: Specimens GRU-F 1 and GRU-F 2 are me-
dium worn. Their morphology is seleno-brachyodont and their
size is relatively small. Parastyle, mesostyle and the paracone
column are strong and they are not connected at the base.
There are weak anterior cingula and entostyles. Specimen
GRU-F 1 shows a spur of the posthypocrista and a neocrista.
GRU-F 3 and GRU-F 4 are strongly worn and have lost
their morphological details. Moreover specimen GRU-F 3
lacks its most anterior part and specimen GRU-F 4 lacks its
most lingual part.
Discussion: Based on the small size those teeth might be-
long to the selenodont-brachyodont European ruminants
Pomelomeryx gracilis (Pomel, 1853) (Moschidae) (Rössner &
Rummel 2001), Tuscomeryx huerzeleri Abbazzi, 2001
(Moschidae), Micromeryx flourensianus Lartet, 1851
(Moschidae), Micromeryx styriacus Thenius, 1950
(Moschidae), Hispanomeryx aragonensis Azanza, 1986 (Bo-
voidea), Hispanomeryx duriensis Morales, Moyà-Solà et Soria,
1981 (Bovoidea), Lagomeryx pumilio Roger, 1896 (Cervidae),
and Lagomeryx parvulus (Roger, 1898) (Cervidae). Distinctive
morphological details are no longer preserved except for the
shape of the teeth. Compared with P. gracilis, L. pumilio and
H. duriensis the teeth show a different shape in the upper mo-
lars (less square) and the P4 (more symmetrical). Moreover L.
pumilio is smaller than the specimens from Grund. In Mi-
cromeryx styriacus, Tuscomeryx huerzeleri, and L. parvulus
the shape is the same, but the size is larger. Hispanomeryx
aragonensis is only known from lower teeth, but estimated
from those, it might come close in size. M. flourensianus show
the same shape and size. Hence several of the postcranial ele-
ments differ from Hispanomeryx but resemble M. flourensian-
us therefore the specimens GRU-F 1, GRU-F 2, GRU-F 3,
and GRU-F 4 most likely belong to M. flourensianus. But be-
cause of further missing morphological species characters
(crown height, closed anterior valley in p4, third lingual conid
of m3, dorsodistally closed gulley of the metatarsal III+IV;
Gentry et al. 1999) the present determination is cf. M. flou-
Material: NHMW GRU-Kroh 9, middle phalange
(l = 13.5; tp = 4.7; td = 3.1); Fig. 4A.
Description: The phalange is small as well as compara-
tively long and slender within ruminants (see Köhler 1993). In
exterior view the proximal articular surface is strongly con-
cave. The Plateau Postarticulaire (Heintz 1970) is proximally
elongated. The dorsal extensor process is short. In interdigital
view the outline of the distal articular surface is well rounded.
Volarly it extends only a little and dorsally strongly. The inter-
digital part of the distal articulation is slightly elongated. The
volar surface includes distally a shallow sagittal groove.
RUMINATIA FROM THE GRUND FORMATION OF THE MIDDLE MIOCENE (AUSTRIA) 341
Fig. 4. cf. Micromeryx flourensianus. A NHMW GRU-Kroh 9, middle phalange. A
external view, A
dorsal view, A
nal view, A
plantar or palmar view, A
proximal view, A
distal view. B NHMW GRU-Kroh 8, right Condylus humeri.
cranial view, B
medial view, B
caudal view, B
lateral view. Scale bar 5 mm.
The outstanding slenderness of the proximal articular sur-
face fits with the slenderness of the distal articular surface of
specimen GRU-F 11. But the size is a little bit too small. Pos-
sibly specimen GRU-Kroh 9 is an anterior phalange and spec-
imen GRU-F 11 is a posterior phalange of the same species.
Discussion: Generally this specimen differs from Tragul-
idae by its slenderness. Moreover in Tragulidae the proximal
articular surface does not have an elongated Plateau Postar-
ticulaire. Its delicate size combined with its slenderness is
comparable to a phalange of Micromeryx flourensianus fig-
ured by Filhol (1891, Pl. 25, Figs. 12), but unlike Hispano-
meryx duriensis Morales et al. 1981 (Pl. 1, Fig. 7). All Middle
Miocene Bovidae can be eliminated, because of a general larg-
er body size. The smaller cervids from that time might be a
possible source. Lagomeryx parvulus as well as Lagomeryx
pumilio from Sandelzhausen (Germany, MN5, Rössner 1997,
2002, 2004) show little differences in size and significant in
shape. From Euprox minimus (Toula, 1884) no middle pha-
lange is known, but it would be probably most similar to L.
parvulus. Therefore the most precise determination what can
be given is cf. Micromeryx flourensianus.
Material: NHMW GRU-Kroh 7, distal fragment of left
humerus (td = 13.9; apd = 11.4); Fig. 7F. NHMW GRU-Kroh 8,
right Condylus humeri (td = 13.5; apd = 10.3); Fig. 4B.
Description: The specimens belong to a comparatively
small ruminant. The body is slender without prominent crests
or obvious rough areas. The mediodistal Epicondylus medialis
is strong with a sagittal ligament groove distally. The Condy-
lus humeri shows medially a narrow rounded shallow liga-
ment groove and the Capitulum humeri shows laterally a wide
rounded deep ligament groove. The diameter of the Trochlea
medially is twice as large as the one of the Capitulum laterally.
The sagittal crest of the Condylus is rounded.
Discussion: The size of the specimen indicates a small ru-
minant. The small tragulid Dorcatherium guntianum Meyer,
1846 differs by a larger size and a different morphology of the
Capitulum humeri (Hambach, Germany, MN5, Mörs 2000).
The moschids Pomelomeryx gracilis and P. boulangeri (Pomel,
1853) as well as the cervids Lagomeryx pumilio and L. parvulus
can be excluded from the determination: In P. gracilis and P.
boulangeri from Wintershof-West (Germany, MN3, Rössner
1997, 2002) the Condylus medially has the same width but a
larger height, in L. pumilio and L. parvulus from Sandelzhausen
(Germany, MN5, Rössner 1997, 2002, 2004) the Condylus is
morphologically identical, but larger. Hispanomeryx duriensis
differs in morphology with a more pronounced Verticillus (Mo-
rales et al. 1981, Pl. I, Fig. 11). Morphologically and in size this
specimen is most similar to Micromeryx flourensianus (Fihol,
1851, Pl. XXV, Figs. 3, 4) with some uncertainties left from the
comparison with a figure.
Micromeryx cf. styriacus Thenius, 1950
Material: NHMW GRU-Kroh 5, dex M2 (l = 7.4; aw = 7.6;
pw = 7.3); Fig. 5A.
Description: The seleno-brachyodont tooth is very well
preserved and only weakly worn. It is relatively small. Its
shape is nearly square and usur facets are developed anteriorly
and posteriorly, which indicates an M2 (see Rössner 1995: Ta-
ble 2). Its styles and the column of the paracone are slender
and not connected at the base. A short but clear anterior cingu-
lum is developed. The spur of the posthypocrista is strong and
the Neocrista is poorly developed sitting at the end of the post-
protocrista. The entostyle is small.
Discussion: This small-sized brachyodont ruminant is
similar to the European Early to Late Miocene moschids
Pomelomeryx gracilis, Micromeryx flourensianus, Microme-
ryx styriacus, and Tuscomeryx huerzeleri, the Early to Middle
Miocene cervids Lagomeryx parvulus and L. pumilio, as well
as the Middle and Late Miocene bovoids Hispanomeryx ara-
gonensis and H. duriensis. It differs from P. gracilis, L. parvu-
lus and L. pumilio by a slightly higher crown as well as more
slender styles and paracone-column. The labial crown base is
less bulging than in the latter species. Specimen GRU-Kroh 5
is a little bit larger than M. flourensianus from Sansan (type
locality, France, MN6) and a little bit smaller than T. huerze-
leri as well as H. duriensis. It might fit in the size class of M.
styriacus from Göriach (type locality, Austria, MN5), which is
slightly larger than M. flourensianus, but only recorded by an
m3, an m2 and a p4 (Hofmann 1893; Thenius 1950) and still
discussed as sexual dimorphism within M. flourensianus.
Hence direct comparison is not possible for the time being
specimen GRU-Kroh 5 is described as Micromeryx cf. styri-
acus. It can not be compared with incomplete remains of H.
Fig. 5. A NHMW GRU-Kroh 5, Micromeryx cf. styriacus,
dex M2. A
occlusal view, A
labial view. B NHMW
GRU-Kroh 6, cf. Eotragus clavatus, dex m1/2/3. B
labial view. Scale bar 5 mm.
Fig. 6. A NHMW GRU-F 1, cf. Micromeryx flourensianus, sin M1.
occlusal view, A
labial view. B NHMW GRU-F 2, cf. Mi-
cromeryx flourensianus, dex M2. B
occlusal view, B
view. C NHMW GRU-F 3, cf. Micromeryx flourensianus, sin m1,
occlusal view. D NHMW GRU-F 4, cf. Micromeryx flourensianus,
dex P4, occlusal view. Scale bar 5 mm.
aragonensis (see above), but the exclusively Spanish abun-
dance of the latter one, makes this assessment improbable.
Family Bovidae Gray, 1821
Genus Eotragus Pilgrim, 1939
cf. Eotragus clavatus (Gervais, 1850)
Material: NHMW GRU-Kroh 6, dex m1/2/3 (l = ;
aw = 9.1; pw = ); Fig. 5B.
Description: The tooth is strongly worn and only par-
tially preserved. The morphology is selenodont-brachyodont
with a barely recognizeable metastylid characteristic for selen-
odont ruminants. There is no trace of a Palaeomeryx-fold.
Discussion: Compared to its wear stage the tooth is still
relatively high crowned. Metastylid and praeentoconid are
fused, which indicates also a relatively higher crown within
the selenodont-brachyodont ruminants. Moreover, the lack of
the Palaeomeryx-fold limits the possible determinations to the
RUMINATIA FROM THE GRUND FORMATION OF THE MIDDLE MIOCENE (AUSTRIA) 343
Fig. 7. A NHMW GRU-B/-1 13, Ruminantia indet. 1, atlas. A
cranial view, A
ventral view, A
caudal view, A
B NHMW GRU-B/-1 10, Pecora indet. 3, proximal fragment of right ulna. B
lateral view, B
cranial view, B
medial view. C
NHMW GRU-B/-1 14, Pecora indet. 4, proximal fragment of proximal phalange, proximal view. D NHMW GRU-B/-1 15, Ruminantia indet.
2, left intermediale. D
lateral view, D
proximal view, D
medial view, D
distal view, D
dorsal view, D
plantar view. E
NHMW GRU- F 11, Pecora indet. 1, proximal phalange III or IV. E
internal view, E
dorsal view, E
external view. E
or palmar view, E
proximal view, E
distal view. F NHMW GRU-Kroh 7, cf. Micromeryx flourensianus, distal fragment of left hu-
lateral view, F
cranial view, F
medial view, F
caudal view, F
distal view. Scale bar 10 mm.
group of early comparably low crowned European Bovidae:
Eotragus artenensis Ginsburg et Heintz, 1968, Eotragus clava-
tus (Gervais, 1850), Eotragus cristatus (Biedermann, 1873),
Pseudoeotragus seegrabensis van der Made, 1989. E. artenen-
sis is a little smaller and lower crowned. P. seegrabensis and E.
cristatus are smaller, too. E. clavatus is of the same size and
therefore specimen GRU-Kroh 6 is assigned to it. However, the
incompleteness of the tooth makes it necessary to restrict the de-
termination to cf. Eotragus clavatus. The bovoid Amphimos-
chus can be excluded because of a stronger ectostylid.
Determinations exact enough for biochronological interpre-
tation of the Ruminantia from Grund comprise Micromeryx cf.
styriacus, cf. Micromeryx flourensianus, and cf. Eotragus
clavatus. M. styriacus, M. flourensianus and E. clavatus are
known from the stratigraphical context of Europe from the
Mammal Zone MN5 onwards (Gentry et al. 1999). However
their total occurrence spans differ: while M. flourensianus is
known in Europe until MN11, E. clavatus persisted only to
MN6, and M. styriacus is restricted to late MN5. In a whole,
the biochronological age indicated by the ruminant fossils
from Grund is restricted from MN5 to MN6, if we are careful
with the M. cf. styriacus, or even to late MN5. Consequently
the hitherto identified age of late MN5 for the terrestrial verte-
brate assemblage from Grund (Daxner-Höck 2003), mainly
based on the micromammal elements, is confirmed by the oc-
currence of the Ruminantia represented.
The marine microfauna from Grund corresponds to the Cal-
careous Nannoplankton Zone NN5, based on the co-occur-
rence of Sphenolithus heteromorphus and Helicosphaera wal-
trans (Æoriæ 2003), and more precisely to the top of Planktonic
Foraminifera Zone M5b/Mt5b, based on the evolutionary lev-
el of Praeorbulina glomerosa circularis and therewith to the
top of the regional Lower Lagenid Zone (Lower Badenian)
(Rögl & Spezzaferri 2003). The co-occurrence with the mam-
mal assemblage placed in late MN5 makes Grund a reference
tie point for marine/continental correlation in the Middle Mi-
ocene of the Central Paratethys (Harzhauser et al. 2003; Dax-
ner-Höck et al. 2004).
The allochthonous nature of the remains of terrestrial verte-
brates in the studied marine sections of Grund (Roetzel 2003;
Roetzel et al. 1999) limits paleoecological interpretations to
purely taxonomically related evidence. Even the poor rumi-
nant record, as much in specimens as in species number, from
Grund gives an insight into the composition of this assem-
blage. Micromeryx and Eotragus are typical representatives
for terrestrial faunas from the early Middle Miocene of Eu-
rope. Micromeryx is an extremely long legged moschid (Gen-
try et al. 1999) with relatively long metapodials, what is typi-
cal for bovids living in open country (Scott 1985). For
Eotragus a duiker-like habitus and mode of life in more or less
dense forests (Köhler 1993; Rössner 2002) can be reconstruct-
ed. Their abundance usually is rare to medium depending on
the local habitat structure. But in the case of Grund, they seem
to be the major faunal elements. Interestingly cervids, which
usually are the major component in European Miocene forest
communities (Rössner 1997, 2004), are not recorded by teeth.
Of course they might be represented within the Pecora indet.
or Ruminant indet. specimens. Moreover tragulids are miss-
ing, which are usually also a major component in European
Miocene forest communities, but indicate strongly moist
ground conditions in a habitat with lots of small open standing
waterbodies (Köhler 1993; Rössner 1997, 2004). Their ab-
sence might be due to dry ground conditions. Considering all
that, the deduced environment agrees with the reconstruction
of Daxner-Höck (2003). She suggested a habitat with low
groundwater level at either an elevated region of a Peninsula
or islands or small mountains of the hinterland, which are cov-
ered with woods, bushes and shrubs.
Significant differences in species composition among Aus-
trian faunas of the early MN5/late Karpatian (Obergänserndorf
and Teiritzberg) and the late MN5 /early Badenian (Grund)
were demonstrated by Daxner-Höck (2003) for the rodents.
These led to the conclusion of changes from a humid to a more
dry climate under similar temperature conditions (subtropical
climate with a mean annual temperature of 1417, frost free
winters, a minimal cold month temperature of 38 °C, annual
precipitation of approximately 2000 mm, Harzhauser et al.
2002) in Central Europe. The ruminants of Obergänserndorf
and Teiritzberg comprise remains of Ruminantia indet., the
tragulid Dorcatherium crassum and a cervid Procervulinae in-
det. (Rössner 1998), which support the idea of a more humid
environment with dense forests for the late Karpatian.
The extraordinary case of a locality yielding marine and ter-
restrial faunas in the same strata made Grund a reference tie
point for marine/continental correlation in the early Middle
Miocene Central Paratethys (Lower Badenian). The terrestrial
fauna includes fossil remains of Ruminantia, which strengthen
the biostratigraphical correlation. The identification of the
species cf. Micromeryx flourensianus, Micromeryx cf. styria-
cus and cf. Eotragus clavatus indicates an age from European
Land Mammal Zone MN5 to MN6 and strongly supports the
already established ages of late MN5 (micromammals) and the
top of Planktonic Foraminifera Zone M5/Mt5b (top of the
Lower Lagenid Zone).
The interpretation of the preferred environment of the repre-
sented Ruminantia supports the reconstruction of a woodland
habitat and the previously assumed successive drying during
MN5 in Central Europe.
Acknowledgments: G. Daxner-Höck (Vienna), A. W. Gentry
(London), J. van der Made (Madrid), and F. F. Steininger
(Frankfurt am Main) gave helpful comments on earlier drafts
of the manuscript. G. Janssen (Munich) did the photographs.
The sketches for the locality map and the stratigraphic table
were kindly offered by G. Daxner-Höck.
Abazzi L. 2001: Cervidae and Moschidae (Mammalia, Artiodactyla)
from the Baccinello V-3 Assemblage (Late Miocene, Late Turo-
RUMINATIA FROM THE GRUND FORMATION OF THE MIDDLE MIOCENE (AUSTRIA) 345
lian, Grosseto Italy). Riv. Ital. Paleont. Stratigr. 107, 1, 107123.
Azanza B. 1986: Estudio geológico y paleontológico del Mioceno
del sector oeste de la comarca de Borja (Provincia de Zarago-
za). Cuad. Estudios Borjanos, XVII-XVIII, 63126.
Biedermann W.G.A. 1873: Petrefacten aus der Umgegend von Win-
therthur. 4. Reste aus Veltheim. Buchdruckerei von J. Westfe-
hling, Wintherthur, 116.
Æoriæ St. 2003: Calcareous nannofossil biostratigraphy of the Mühl-
bach section. Ann. Naturhist. Mus., Wien 104 A, 1522.
Daxner-Höck G. 2003: Cricetodon meini and other rodents from
Mühlbach and Grund (late MN5). Ann. Naturhist. Mus., Wien
104 A, 267291.
Daxner-Höck G., Göhlich U.B., Huttunen K., Kazár E., Nagel D., Röss-
ner G.E., Schultz O., Miklas-Tempfer P.M. & Ziegler R. 2004:
Marine and terrestrial vertebrates from the Middle Miocene of Gr-
und (Lower Austria). Geol. Carpathica 55, 2, 191197.
Filhol H. 1891: Études sur les mammifères fossiles de Sansan. Li-
braire de lAcadémie de Médecine, Paris, 1319.
Gentry A.W., Rössner G.E. & Heizmann E.P.J. 1999: Suborder Ru-
minantia. In: Rössner G.E. & Heissig K. (Eds.): The Miocene
Land Mammals of Europe. Verlag Dr. Friedrich Pfeil,
Gervais M.P. 18481852: Zoologie et paléontologie francaises (ani-
maux vertébrès) ou nouvelles recherches sur les animaux vi-
vants et fossiles de la France. 1, 1271.
Ginsburg L. & Heintz É. 1968: La plus ancienne antilope dEurope,
Eotragus artenensis du Burdigalien dArtenay. Bull. Mus. Nat.
Hist. Natur. 2, 40, 4, 837812.
Gray J.E. 1821: On the natural arrangement of vertebrose animals.
London Medical Repository 15, 296310.
Harzhauser M., Böhme M., Mandic O. & Hofmann Ch.-Ch. 2002:
The Karpatian (Late Burdigalian) of the Korneuburg Basin. A
paleoecological and biostratigraphical synthesis. Beitr.
Paläont. 27, 441456.
Harzhauser M., Daxner-Höck G., Boon-Kristkoiz E., Æoriæ S., Man-
dic O., Miklas-Tempfer P., Roetzel R., Rögl F., Schultz O.,
Spezzaferri S., Ziegler R. & Zorn I. 2003: Paleoecology and
biostratigraphy of the section Mühlbach (Gaindorf Formation,
lower Middle Miocene, Lower Badenian, Austria). Ann.
Naturhist. Mus. Wien, 104 A, 323334.
Heintz E. 1970: Les cervidés Villafranchiens de France et
dEspagne. Volume 1: Texte et planches, Volume 2: Figures et
Tableaux. Mém. Mus. Nat. Hist. Natur. C, XXII, 1, 1303.
Hofmann A. 1893: Die Fauna von Göriach. Abh. K.-Kön. Geol. Re-
ichsanst. XV, 6, 187.
Köhler M. 1993: Skeleton and Habitat of recent and fossil Rumi-
nants. Müncher Geowiss. Abh. A, 25, 188.
Lartet E. 1851: Notice sur la Colline de Sansan, suivie dune récapit-
ulation des diverses espèces danimaux vertébrés fossiles, trou-
vés soit a Sansan, soit dans dautres gisements du terrain
tertiaire Miocène dans le Bassin Sous-Pyrénéen. J.-A. Portes,
Auch, France, 146.
Linnaeus C. von 1758: Systema naturae per regna tria naturae, se-
cundum classes, ordines, genera, species, cum characteribus,
differentiis, synonymis, locis. 1: Regnum animale. Laurentii
Salvii, Stockholm, 1823.
Made J.v.d. 1989: The bovid Pseudoeotragus seegrabensis nov. gen.,
nov. sp. from the Aragonian (Miocene) of Seegraben near Leoben
(Austria). Proc. K. Nederl. Akad. Wet., Ser. B, 92, 3, 215240.
Meyer H. von 1846: Mittheilungen an Professor Bronn. Neu. Jb.
Mineral., Geol., Geognäsie und Petrefaktenkunde, Jg. 1846,
Mörs T., von der Hocht F. & Wutzler B. 2000: Die erste Wirbeltierfau-
na aus der miozänen Braunkohle der Niederrheinischen Bucht
(Ville-Schichten, Tagebau Hambach). Paläont. Z. 74, 45170.
Morales J., Moyà-Solà S. & Soria D. 1981: Presencia de la familia
Moschidae (Artiodactyla, Mammalia) en el Vallesiense de Es-
pana: Hispanomeryx duriensis nov. gen. nov. sp. Estud. Geol.
Pilgrim G.E. 1939: The fossil Bovidae of India. Mem. Geol. Surv.
India, New Ser. 26, 1, 1356.
Pomel A. 1853: Catalogue méthodique et descriptif des Vertébrés
fossiles découverts dans le bassin Hydrographique supérieur de
la Loire et surtout dans la Vallé de son affluent principal
Rögl F., Reiser H., Rupp Ch. & Wenger W.F. 1998: BavarianAus-
trian Molasse Basin. In: Cicha I., Rögl F., Rupp Ch. & Ètyroká
J. (Eds.): OligoceneMiocene foraminifera of the Central
Paratethys. Abh. Senckenberg.Naturforsch. Gessell. 549, 715.
Rögl F. & Spezzaferri S. 2003: Foraminiferal paleoecology and bio-
stratigraphy of the Mühlbach section. Ann. Naturhist. Mus.,
Wien 104 A, 2376.
Rössner G.E. 1995: Odontologische und schädelanatomische Unter-
suchungen an Procervulus (Cervidae, Mammalia). Müncher
Geowiss. Abh. A, 29, 1127.
Rössner G.E. 1997: Biochronology of ruminant assemblages in the
Lower Miocene of Southern Germany. In: Aguilar J.-P., Leg-
endre S. & Michaux J. (Eds.): Actes du Congrès Biochrom97.
Mém. Trav. de lE.P.H.E., Inst. Montpellier 21, 609618.
Rössner G.E. 1998: Säugetiere (Mammalia) aus dem Karpat des Ko-
rneuburger Beckens. 4. Ruminantia (Artiodactyla). Beitr.
Paläont. 23, 409413.
Rössner G.E. 2002: Miozäne Ruminatia Süddeutschlands. Taxo-
nomie und Ökologie. Habilitationsschrift, unpublished, Lud-
wig-Maximilians-Universität München, München, 1160.
Rössner G.E. 2004: Community structure and regional patterns in
late Early to Middle Miocene Ruminantia of Central Europe.
In: Steininger F.F., Kovar-Eder J. & Fortelius M. (Eds.): The
Middle Miocene environments and ecosystem dynamics of
the Eurasian Neogene (EEDEN). Courier Forschungsinst.
Senckenberg 249, 91100.
Rössner G.E. & Rummel M. 2001: Pomelomeryx gracilis (Pomel,
1853) (Mammalia, Artiodactyla, Moschidae) from the Lower
Miocene karstic fissure filling complex Rothestein 10/14 (Ger-
many, Bavaria). Lynx, New Ser. 32, 323353.
Roetzel R. 2003: Zur Geologie der mittelmiozänen Fossilfundstelle
Mühlbach am Manhartsberg (Niederösterreich). Ann. Naturhist.
Mus., Wien 104 A, 313.
Roetzel R., Pervesler P., Daxner-Höck G., Harzhauser M., Mandic
O., Zuschin M. & Cicha I. 1999: C4 Grund Kellergasse. Ar-
beitstagung der Geologische Bundesanstalt. Exkursionen A,
Geologie der Umgebung Retz. Geologische Bundesanstalt,
Roger O. 1896: Vorläufige Mittheilungen über Säugethierreste aus
dem Dinotheriensand von Stätzling bei Augsburg. Ber. Natur-
wiss. Ver. Schwaben und Neuburg 32, 547552.
Roger O. 1898: Wirbelthierreste aus dem Dinotheriensande der bay-
erisch-schwäbischen Hochebene. Ber. Naturwiss. Ver. Schwa-
ben und Neuburg 33, 146.
Scopoli G.A. 1777: Introductio ad Historiam naturalem, sistens gen-
era Lapidum, Plantarum et Animalium hactenus detecta, carac-
teribus essentialibus donata, in tribus divisa, subinde ad leges
Naturae. Gerle, Prague, Czechia, X+1506+134.
Scott K.M. 1985: Allometric trends and locomotor adaptations in the
Bovidae. Bull. Amer. Mus. Natur. Hist. 179, 2, 197288.
Thenius E. 1950: Die tertiären Lagomeryciden und Cerviden der
Steiermark. Beiträge zur Kenntnis des steierischen Tertiärs V.
Sitzu. Österr. Akad. Wiss., Math.-Naturwiss. Kl., Abt. I, 159, 6
Toula F. 1884: Ueber einige Säugethierreste von Göriach bei Turnau
(Bruck a. M. Nord) in Steiermark. Jb. K.-Kön. Geol. Reich-
sanst. 34, 3, 49401.