GEOLOGICA CARPATHICA, 53, 4, BRATISLAVA, AUGUST 2002
269—279
TEMPORAL AND SPATIAL DISTRIBUTION
OF MIOCENE MAMMALS IN THE WESTERN CARPATHIANS
(SLOVAKIA)
MARTIN SABOL and PETER HOLEC
Department of Geology and Paleontology, Faculty of Science, Comenius University, Mlynská dolina, 842 15 Bratislava, Slovak Republic
(Manuscript received, May 15, 2001; accepted in revised form December 13, 2001)
Abstract: Mammal fossils are found relatively rarely in the Miocene deposits of Slovakia. So far, eleven Miocene
mammal localities are known. They cover a time span from the Middle Badenian to the Pannonian. Miocene mammals
have been found mostly at the NE margin of the Vienna Basin (Zapfe’s fissures, Sandberg, Bonanza, Wait Quarry,
Glavica, Dúbravská hlavica, and Borský Svätý Jur). However, some localities are also known from the northern part of
the Danube Basin (Pezinok, Topo čany-Kalvária, and Nováky) and from the northern margin of the Transcarpathian
(East Slovak) Basin (Košice-Bankov). The article provides a review of Slovak Miocene sites of fossil mammals and
updates some older stratigraphical data.
Key words: Miocene, Western Carpathians, Slovakia, mammals, biostratigraphy.
Introduction
Fossil findings of mammals are found relatively rarely in the
Miocene deposits of Slovakia. In total, eleven localities are
known which yielded Miocene mammals. The mammal fau-
nas date the single sites to MN 6 until MN 10—11 biozones of
the Neogene European Land Mammal Zones (Table 1).
From the biostratigraphic and paleoecological point of view,
localities on the slopes of Devínska Kobyla Hill near Devíns-
ka Nová Ves (the former German name of this modern subur-
ban part of Bratislava was “Neudorf an der March” or only
“Neudorf”) in the SW Slovakia are the most important and the
best known. The sediments of these sites contain a large quan-
tity of Miocene marine and terrestrial mammals. Holec & Sab-
ol (1996) gave the first detailed list of the fossil taxa known
from Devínska Kobyla localities.
However, a large quantity of new data on the Neogene eco-
systems and environmental changes has been published re-
cently (e.g. “EEDEN Project”). The result of that are already
well-known “high-resolution” time intervals for the Eurasian
realm. The results of many paleovertebratologists have yielded
new data including major changes in the composition of ma-
rine and terrestrial vertebrate communities (e.g. Rössner &
Heissig 1999).
These new data have been used for the revision of the Mi-
ocene mammal faunas from Slovak territory. The older data
have been restudied and some of them corrected. This article
provides a detailed review of the time distribution of Miocene
mammal faunas in the territory of Slovakia.
Localities
The Slovak localities with Miocene mammal fossils are
situated in three basins. Most of these sites is situated at the
NE margin of the Vienna Basin (Zapfe’s fissures, Bonanza,
Sandberg, Wait Quarry, Glavica, Dúbravská hlavica, and
Borský Svätý Jur). However, fossil findings of Miocene
mammals are also known from the northern part of the
Danube Basin (Pezinok, Topo čany-Kalvária, and Nováky)
and from the northern margin of the Transcarpathian (East
Slovak) Basin (Košice-Bankov) (Fig. 1 and Table 2).
The Zapfe’s fissures site (also known as Neudorf-Spalte) is
the biostratigraphically oldest locality with mammal fossils in
Slovakia. It is situated in the northern margin of the Vienna
Basin near Bratislava on the northern slope of Devínska Koby-
la Hill in the Stockerau limestone pit (the exact geographical
position: longitude (LG) is 17° 01´ and latitude (LA) is 48°
12´) (Fig. 1). The locality consists of some fissures in the dark
recrystallized Jurassic limestone of the Tatric Devín Succes-
sion of the Malé Karpaty Mts (in sense of Plašienka et al.
1991). They are often filled by the sinter, cave sediments, “ter-
ra rossa” and “terra fusca” (Mišík 1976) with the marine sandy
deposits of the Sandberg Member overlying them. The site is
dated to the lower part of the MN 6 biozone (Astaracian, the
uppermost part of the Middle Badenian) on the basis of fossil
mammal assemblages which were found in the terrestrial sedi-
ments of two of them (Cícha et al. 1972; Fejfar 1974) (Table 2).
The Bonanza locality was discovered in 1984. It is situated in
the territory of the same quarry as the Zapfe’s fissures site
(Stockerau limestone pit; the exact geographic position is
same), but on its eastern margin (Fig. 1). The site is represented
by a broad fissure in the protective limestone wall oriented to-
wards the railway line from Bratislava to Prague. Marine
sands, sandstone and large limestone boulders fill the fissure.
The sandy sediments contain fossil remnants of marine and
terrestrial vertebrates and invertebrates (Holec et al. 1987;
Ivanov 1998). Hereby, frog fossils (Špinar et al. 1993) suggest
the presence of a fresh-water environment in the near vicinity
of the locality during the period of deposition of the fossilifer-
270 SABOL and HOLEC
ous sediments. Their Badenian age has been determined on the
basis of the presence of the bivalve Pecten aduncus Eichwald
only (Holec et al. 1987). However, new findings of mammals
(e.g. Dinosorex zapfei Engesser, 1975 or Democricetodon vin-
dobonensis (Schaub et Zapfe, 1953)), indicate the Late Bade-
nian age of the site (MN 6b) (Tables 2 and 3).
Approximately 2 km westwards from the Stockerau lime-
stone pit on the north-western slope of Devínska Kobyla Hill,
the Sandberg locality is situated on the southern border of
Devínska Nová Ves (LG: 17° 00´, LA: 48° 12´) (Fig. 1). This
significant paleontological site was determined as the facios-
tratotype for the Bulimina-Bolivina Zone of the Late Badenian
(Švagrovský 1978). The locality consists of shallow littoral
sediments of the Sandberg Member (basal coarse clastics, mi-
caceous sands, clayey sands, calcareous sandstone, organo-
gene and organodetritic limestone, breccias, and gravel layers
(Baráth et al. 1994)) which are situated erosively on older se-
quences of the Mesozoic carbonates of the Tatric Devín Suc-
cession of the Malé Karpaty Mts (in sense of Plašienka et al.
1991). They are products of the Late Badenian transgressive
event on the eastern margin of the Vienna Basin partly con-
trolled by transtensive tectonics (Baráth et al. 1994). Abundant
terrestrial and marine fauna of the vertebrates and inverte-
brates has been found here especially in the fine-grained grav-
el layers (Švagrovský 1978). On the basis of large terrestrial
mammal fossils (fossils of micromammals have not been
found in these marine sediments), the locality is dated to the
Astaracian (MN 6b to MN 7+8?) (Table 2).
The Wait Quarry is the next locality in the vicinity of
Devínska Nová Ves. It is an abandoned quarry on the western
slope of Devínska Kobyla Hill, approximately 600—700 metres
southwards from the Sandberg site (LG: 17° 00´, LA: 48° 12´)
(Fig. 1). The site consist of transgressive horizontal layers of
the Upper Badenian marine sands (Sandberg Member) situat-
ed in Lower Jurassic carbonates, especially in the upper parts
of the quarry. Only findings of fishes, turtles and seals (Ta-
ble 3) have been found here in the marine sands which fill the
space of one little submarine cave (Zágoršek 1985). The age of
the site probably corresponds to the age of the Sandberg site
(MN 6b to MN 7+8?, Astaracian, the Late Badenian) (Ta-
ble 2).
Fossils of whales (Mesocetus sp.) together with an undeter-
mined mammalian rib have been found in the Glavica locality
(Holec 2001). The locality is also situated near the Sandberg
site approximately 1 km eastwards from Devínska Nová Ves
(LG: 17° 00´, LA: 48° 12´) (Fig. 1). Like the Wait Quarry site,
it consists of Upper Badenian marine sands in which layers of
fine and coarse fractions alternate (Sandberg Member). The
age of the locality was determined on the basis of the poor
mollusc assemblages and it also corresponds with the Sand-
berg age (MN 6b to MN 7+8?, Astaracian) (Table 2).
The Nováky-Mier Mine locality is situated in the territory
of the Danube Basin approximately 10 km south-west of
Prievidza town in the central part of Slovakia (LG: 18° 35´,
LA: 48° 42´) (Fig. 1). It is a coal mine where dark clays with
coal and dark-green tuffaceous clays to sandy tuffites underlie
the main coal bed with grey to dark-green tuffaceous clays
overlying it (Takáč 1970). Andesite flows and pyroclastic
rocks cover this Middle Miocene (Nováky Formation; Late
Badenian—Early Sarmatian) coal complex. Besides plant fos-
sils (especially fossils of the families Taxodiaceae, Cupresace-
ae, Myriacaceae, Junglandaceae, Fagaceae, Hamamelidace-
ae, Lauraceae, and others (Sitár & Takáč 1993)), one tooth of
the Zygolophodon turicensis (Schinz, 1824) and odd fossils of
rhinoceros have been found in the lignite bed too (Table 3).
The more exact age of the fossils is not known. However on
the basis of the flora findings, it is assumed the Late Bade-
nian—Early Sarmatian (Sitár et al. 1987; Sitár & Takáč 1993).
From the MN-zonation point of view it corresponds especially
to the MN 7+8 biozone (Astaracian) (Table 2).
The Dúbravská hlavica locality is situated near Bratislava
approximately 1-km south-eastwards from the Stockerau lime-
stone pit and approximately 300 metres southwards from the
Dúbravská hlavica trigonometric point (LG: 17° 01´, LA: 48°
11´) (Fig. 1). It consists of the Sarmatian clayey sediments
(Karlová Ves Member) deposited above Late Badenian marine
Fig. 1. Location of Slovak localities where fossil remains of Mi-
ocene mammals have been found. 1 – Zapfe’s fissures, 2 – Bo-
nanza, 3 – Sandberg, 4 – Wait Quarry, 5 – Glavica, 6 –
Nováky-Mine Mier, 7 – Dúbravská hlavica, 8 – Košice- Bankov,
9 – Pezinok, 10 – Topo čany-Kalvária, 11 – Borský Svätý Jur.
TEMPORAL AND SPATIAL DISTRIBUTION OF MIOCENE MAMMALS 271
Table 1: Middle to Late Miocene circum-Mediterranean marine-continental chronological correlations of the European Mammal Units
and Zones (after Steininger 1999).
272 SABOL and HOLEC
sands with fossils of the gastropod genera Turitella and Conus
(Sandberg Member). Fossils of one rhinoceros individual have
been found in the Sarmatian clayey deposits only (Holec &
Sabol 1996) (Table 2).
The fossils of the Miocene mammals have been described
from one locality in the territory of Eastern Slovakia only. It is
the magnesite mine of Bankov (Fig. 1) at the north-western
border of Košice town (LG: 21° 13´, LA: 48° 44´) where the
Neogene rhyodacitic tuffs and tuffites fill paleokarst fissures
in the metasomatic Carboniferous carbonates of the Ochtiná
Formation. The fossils of the rare viverrid Lophocyon car-
pathicus Fejfar, Schmidt-Kittler et Zacharov, 1987 and pro-
boscidean tusk fragments have been found in one of the fis-
sures only (Table 3). Fejfar et al. (1987) suppose a Sarmatian
age for the findings (MN 7+8, Astaracian) (Table 2).
Lower Pannonian fossiliferous strata are exposed by the
Pezinok brickyard situated in the WNW part of the Danube
Basin approximately 10 km northwards from Bratislava (LG:
17° 16´, LA: 48° 16´) (Fig. 1). The fossiliferous sediments
(Ivánka Formation) consist of fine pelitic clays, green-grey
sandy clays, fine sands and isolated little lignite beds in clays.
Light and dark clays with isolated carbonate concretions are
situated above this sequence. Besides findings of inverte-
brates, fossils of the Hippotherium primigenium H. v. Meyer,
1829 and Trogontherium have been found especially in the
overlying clays. On the basis of the mollusc assemblage (with
Congeria subglobosa subglobosa Partsch, C. subglobosa
longitesta Papp, Dreissenomya primiformis Papp, and Mono-
dacna viennesis Papp) (Fordinál 1997), the whole sedimentary
complex has been dated to the upper part of the E zone (in
sense of Papp 1951) (Early Pannonian, MN 9, Vallesian; Ta-
ble 2).
The next locality in the territory of the Danube Basin is situ-
ated on the south-western border of Topo čany town in the part
known as Kalvária (LG: 18° 09´, LA: 48° 33´) (Fig. 1). The
Upper Miocene coarse grey sands underlie the Quaternary
sediments. Mandible fragments of Hippotherium primigenium
have been found in the psammites in 1967 (Holec 1981) to-
gether with the proboscidean molars of a so-called “grandinci-
sivoid” taxon (described by Silnický (1930) as Stegotetrabelo-
don grandincisivus). On the basis of these mammal fossils, the
sediments of the site have been dated to the MN 9—10 biozone
(Vallesian) which indicate the Pannonian age (Table 2).
The biostratigraphically youngest locality with findings of
Miocene mammals in the territory of Slovakia (Table 2) is
situated in the northern margin of the Slovak part of the Vien-
na Basin approximately 75 km north of Bratislava. It is loam
pit of the brickyard in Borský Svätý Jur (LG: 17° 02´, LA:
48° 36´) (Fig. 1). The locality consist of clayey silts with some
sandy beds, silty sands and fine sands which contain fossils of
the ostracods, molluscs and vertebrates, especially mammals
(e.g. Eomellivora wimani Zdansky, 1924; Ictitherium viverri-
num Roth et Wagner, 1854; and “Hipparion” s.s.) (Pipík &
Holec 1998). On the basis of mammal fossils, the locality is
Table 2: Age of individual Slovak localities where the fossil remains of the Miocene mammals have been found.
Locality
Basin
Age
European Land Mammal
References
Cent. Paratethys
Mega-zone
MN unit
Zapfe's fissures
Vienna Basin
Middle Badenian
Astaracian
6a
Zapfe 1949, 1950a,b,
1951, 1952, 1953, 1954,
1960, 1976, 1979, 1983
Thenius 1952
Wettstein-Westtersheimb 1955
Estes 1969
Cícha et al. 1972
Fejfar 1972, 1974, 1990
Špinar 1975
Holec 1986
Holec & Sabol 1996
Bonanza
Vienna Basin
Late Badenian
Astaracian
6b
Holec et al. 1987
Špinar et al. 1993
Holec & Sabol 1996
Ivanov 1998
Sandberg
Vienna Basin
Late Badenian
Astaracian
6b – 7+8?
Hoernes 1848
Pia & Sickenberg 1934
Thenius 1952
Holec 1986
Holec & Sabol 1996
Wait Quarry
Vienna Basin
Late Badenian
Astaracian
6b – 7+8?
Zágoršek 1985
Holec & Sabol 1996
Glavica
Vienna Basin
Late Badenian
Astaracian
6b – 7+8?
Holec & Sabol 1996
Holec 2001
Nováky-Mier Mine
Danube Basin
Late Badenian–
Astaracian
7+8
Takáč 1970
Early Sarmatian
Sitár et al. 1987
Sitár & Takáč 1993
Dúbravska hlavica
Vienna Basin
Sarmatian
Astaracian
7+8
Holec & Sabol 1996
Košice-Bankov
East-Slovak Basin
Sarmatian
Astaracian
7+8
Fejfar et al. 1987
Pezinok
Danube Basin
Pannonian
Vallesian
9
Fordinál 1997
Pipík 2000
Topoľčany-Kalvária
Danube Basin
Pannonian
Vallesian
9 – 10
Silnický 1930
Borský Svätý Jur
Vienna Basin
Pannonian
Vallesian–
10 – 11?
Lupták 1995a,b
Turolian
Pipík & Holec 1998
TEMPORAL AND SPATIAL DISTRIBUTION OF MIOCENE MAMMALS 273
preliminarily dated to the MN 11 biozone (Turolian, Middle
Pannonian), but it is not excluded that they are older (Table 2).
Faunal History
The oldest fauna of Tertiary mammals in the territory of Slo-
vakia is known from the period of the Middle Badenian (MN
6a). The fossils of single vertebrate taxa of this Astaracian fau-
na have been found in terrestrial sediments of the Zapfe’s fis-
sures locality on Devínska Kobyla Hill. During this period, the
hill was probably a part of peninsula or it was part of a territo-
ry of insular character. The presence of this Middle Badenian
fauna (Table 3) is result of migration waves into the territory
of Europe during this period, especially from Africa. The most
important elements of this African immigration are pliopithec-
ids (Pliopithecus (Epipliopithecus) vindobonensis Zapfe et
Hürzeller, 1957). According to Rögl (1999), they could still
migrate into Europe before the short Langhian re-opening of
the Tethyan Seaway to the Mediterranean. However, the older
mammal elements which are present in the European fauna
since the Early Badenian, are represented in the taxocenoses
too. Many of these elements reached the territory of Europe
from Africa (Zygolophodon, Prodeinotherium, Aureliachoe-
rus, Dorcatherium and others) in the time of the so-called
“Gomphotherium Landbridge”, whereas the presence of other
mammal taxa is a result of the Early Miocene invasion from
Asia (Eotragus, Megacricetodon, Eumyarion and others)
(Agustí et al. 2001; Rögl 1999).
Before the short transgressive event in the Late Badenian,
which reached the Vienna Basin, the terrestrial fauna of mam-
mals probably still persisted in the territory of Devínska Koby-
la Hill. This fauna has been described from the marine sands
of the Sandberg locality, and its composition is very similar to
that from the Zapfe’s fissures site (Table 3). However, the
Sandberg taxocenoses shows also the presence of some differ-
ent, especially younger elements (e.g. Griphopithecus suessi
Abel, 1902, Hemicyon goeriachensis (Toula, 1884), Tapirus
telleri Hofmann, 1893), whereas other ones (e.g. insectivores,
rodents or chalicotheres) are missing here. It is not out of ques-
tion that fauna of this composition also lived in the territory of
Devínska Kobyla Hill near of the sea-shore during the trans-
gression or in the time after this short Late Badenian marine
cycle reached its maximum level. In this period, seals (Devi-
nophoca claytoni (tentative name) and Pristiphoca vetusta,
Zapfe) occupied the marine coast (Bonanza, Sandberg and
Wait Quarry) and sirens (Halitherium cordieri Christol, Thala-
tosiren petersi (Abel), and Metaxitherium sp.) together with
whales (Mesocetus hungaricus Kadic) and dolphins (Schizo-
delphis cf. sulcatus (Gervais, 1861)) lived in waters of this
Late Badenian sea as well (Sandberg, Glavica) (Table 3).
The closure of the seaway between the Indo-Pacific and the
Paratethys in the Early Sarmatian caused the change of salinity
in the Paratethys (Rögl 1999) which was accompanied by a
gradual regression, especially in its central part. Fossils of rhi-
noceros (Brachypotherium sp. from the Dúbravská hlavica
site, and Rhinocerotidae gen. et spec. indet. from the Nováky-
Mier Mine site), mammutids (Zygolophodon turicensis from
the Mier Mine too) and the viverrid Lophocyon carpathicus
together with undetermined remnants of “mastodons” from the
Košice-Bankov site in the territory of Eastern Slovakia are
known from this period only (MN 7+8) (Table 3). However,
these finds are evidence of some intermittent faunal migra-
tions in the time of changing environmental conditions, caused
by the both marine regression and increased volcanic activity
in the area of the Central Paratethys.
The beginning of the MN 9 biozone (Vallesian) is well de-
fined by the first occurrence of “hipparions” in Europe and
their overall expansion in the Northern Hemisphere (Rögl
1999). In this time, the Pannonian Lake had been situated in
the territory of Central Paratethys. Its existence was a result of
paleogeographical changes, which happened in this area be-
fore and at the beginning of the Late Miocene. A diferent con-
sequence of these changes was the immigration of new faunal
elements from Asia (including “hipparions” too) and the ex-
tinction of the Middle Miocene elements. The first appearance
of “hipparions” in European territory is observed from the
Late Bessarabian layers of the Nessebar site on the Bulgarian
coast of the Black Sea (Bakalov & Nikolov 1962). However,
the oldest findings of “hipparions” (Hippotherium primi-
genium) in the Slovak territory are known only from younger
layers of the Pannonian (Pezinok, Topo čany). Apart from the
equid findings, the fossils of rodents (Trogontherium sp.) and
a so-called “grandincisivoid” taxon (gen. indet. grandicisivus)
have been found in the Lower Pannonian sediments in Slovak
territory (Table 3).
The reduction of the Paratethys aquatic environment also
continued after the Early Pannonian (Daxner-Höck 1996). The
mammal assemblage in the sediments of the Borský Svätý Jur
site is the only one of this period found in Slovakia. Fossil in-
sectivores, carnivores, rodents, artiodactyls and perissodactyls
have been found here. Finds of Eomellivora wimani (Lupták
1995a), Ictitherium viverrinum (Lupták 1995b), and “Hippari-
on” s.s. (described as Hipparion sp.; Pipík & Holec 1998) are
among the most important of them (Table 3). The finding of
Eomellivora wimani from the Borský Svätý Jur site is the
northernmost occurrence of this giant mustelid species
(Lupták 1995a).
So far, no fossils of terrestrial mammals from the younger
deposits of the Late Miocene are known in the territory of Slo-
vakia. However, it is not excluded that mammalian fauna of
these periods (Late Pannonian—Pontian—Early Dacian) was
also present here. Its presence in Europe is connected with the
isolation of the Paratethys with nearly fresh-water conditions
and with the migration pathways between Eurasia and Africa
which remained open in the Near East area (Rögl 1999).
Discussion
The biostratigraphy of the Cainozoic continental sediments
is established on the fossil assemblages of the terrestrial mam-
mals, especially on the basis of rodent findings. In spite of
good characterization of single mammalian biozones for the
Neogene by the key-fossils, some differences exist in the opin-
ions of scientists on the determination of the chronological
calibration of single biozones, mainly for the Miocene Period
(Table 4). These differences can also be observed in the dating
274 SABOL and HOLEC
Table 3: The Miocene mammal taxa from Slovak localities. “Sandberg” – presence of the taxon in the site, “Sandberg” – type locality
for present taxon, “Sandberg” – site of the taxon with the problematic taxonomical name.
Taxon
Locality
Age of Locality
MN range
References
Comments
Insectivora
"Allosorex" gracilidens (Viret et Zapfe, 1951)
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 6
Viret & Zapfe, 1951
Amphechinus intermedius (Gaillard, 1899)
Zapfe´s Fissures
Middle Badenian
MN 6a
7+8
Zapfe, 1951
Dinosorex zapfei Engesser, 1975
Zapfe´s Fissures
Middle Badenian
MN 6a
6
Zapfe, 1951
Bonanza
Late Badenian
MN 6b
Lanthanotherium sansaniense (Lartet, 1851)
Zapfe´s Fissures
Middle Badenian
MN 6a
6 - 7+8
Zapfe, 1951
Lartetium dehmi (Viret et Zapfe, 1951)
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 7+8
Viret & Zapfe, 1951
Plesiodimylus chantrei Gaillard, 1897
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 11
Zapfe, 1951
Bonanza
Late Badenian
MN 6b
"Scaptonyx" edwardsi Gaillard, 1899
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 9
Zapfe, 1951
Talpa minuta Blainville, 1840
Zapfe´s Fissures
Middle Badenian
MN 6a
3 - 9
Zapfe, 1951
?Urotrichus dolichochir (Gaillard, 1899)
Zapfe´s Fissures
Middle Badenian
MN 6a
7+8
Zapfe, 1951
Erinaceidae gen. et spec. Indet.
Bonanza
Late Badenian
MN 6b
Soricidae gen. et spec. indet.
Bonanza
Late Badenian
MN 6b
Talpidae gen. et spec. indet.
Zapfe´s Fissures
Middle Badenian
MN 6a
Zapfe, 1951
Bonanza
Late Badenian
MN 6b
Holec et al., 1987
Insectivora gen. et spec. indet.
Bonanza
Late Badenian
MN 6b
Borský Svätý Jur
Pannonian
MN 10 - 11?
Pipík & Holec, 1998
Chiroptera
Megaptera lugdunensis (Depéret, 1892)
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 6
Zapfe, 1950b
Miniopterus fossilis Zapfe, 1950
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 13
Zapfe, 1950b
Paleptesicus priscus (Zapfe, 1950)
Zapfe´s Fissures
Middle Badenian
MN 6a
6 - 7+8
Zapfe, 1950b
Rhinolophus delphinensis Gaillard, 1899
Zapfe´s Fissures
Middle Badenian
MN 6a
3 - 13
Zapfe, 1950b
Rhinolophus grivensis (Depéret, 1892)
Zapfe´s Fissures
Middle Badenian
MN 6a
3 - 10
Zapfe, 1950b
Chiroptera gen. et spec. indet.
Bonanza
Late Badenian
MN 6b
Primates
Griphopithecus suessi Abel, 1902
Sandberg
Late Badenian
MN 6b - 7+8? 6
Thenius, 1952
Pliopithecus antiquus Blainville, 1839
Sandberg
Late Badenian
MN 6b - 7+8? 5 - 8
Thenius, 1952
Pliopithecus vindobonensis Zapfe et Hürzeller, 1957
Zapfe´s Fissures
Middle Badenian
MN 6a
6
Zapfe, 1952
Carnivora
Alopecocyon leptorhynchus (Filhol)
Zapfe´s Fissures
Middle Badenian
MN 6a
?
Zapfe, 1950a
Amphicyon major Blainville, 1841
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 7+8
Zapfe, 1950a
Amphicyon cf. steinheimensis Fraas, 1885
Zapfe´s Fissures
Middle Badenian
MN 6a
6 - 7
Zapfe, 1950a
Amphicyon sp.
Sandberg
Late Badenian
MN 6b - 7+8? 1 - 11
Thenius, 1952
Devinophoca claytoni Koretsky (tentative name)
Bonanza
Late Badenian
MN 6b
6
Eomellivora wimani Zdansky, 1924
Borský Svätý Jur
Pannonian
MN 10 - 11?
11 - 12
Lupták, 1995a
Hemicyon goeriachensis (Toula, 1884)
Sandberg
Late Badenian
MN 6b - 7+8? 6 - 9
Thenius, 1952
Hemicyon sansaniensis Lartet, 1851
Zapfe´s Fissures
Middle Badenian
MN 6a
5b - 6
Zapfe, 1950a
Hemicyon cf. sansaniensis Lartet, 1851
Zapfe´s Fissures
Middle Badenian
MN 6a
5b - 6
Zapfe, 1950a
Ictitherium viverrinum Roth et Wagner, 1854
Borský Svätý Jur
Pannonian
MN 10 - 11?
9 - 12
Lupták, 1995b
Lartetictis dubia (Blainville, 1842)
Sandberg
Late Badenian
MN 6b - 7+8? 6
Thenius, 1952
Lophocyon carpathicus Fejfar et al., 1987
Košice-Bankov
Sarmatian
MN 7+8
7+8
Fejfar et al., 1987
Potamotherium miocenicum (Petters, 1868)
Sandberg
Late Badenian
MN 6b - 7+8? 4a - 6
Thenius, 1952
Pristiphoca vetusta (Zapfe)
Bonanza
Late Badenian
MN 6b
6 - 7+8?
Holec et al., 1987
Sandberg
Late Badenian
MN 6b - 7+8?
Holec & Sabol, 1996
Wait Quarry
Late Badenian
MN 6b - 7+8?
Zágoršek, 1985
Pseudaelurus lorteti Gaillard, 1899
Sandberg
Late Badenian
MN 6b - 7+8? 4a - 7+8
Thenius, 1952
Pseudaelurus turnauensis (Hoernes, 1882)
Sandberg
Late Badenian
MN 6b - 7+8? 3 - 9
Thenius, 1952
Pseudaelurus sp.
Zapfe´s Fissures
Middle Badenian
MN 6a
3 - 9
Zapfe, 1950a
Pseudarctos aff. bavaricus Schlosser, 1899
Sandberg
Late Badenian
MN 6b - 7+8? 4 - 9
Thenius, 1952
Sansanosmilus jourdani (Filhol, 1883)
Zapfe´s Fissures
Middle Badenian
MN 6a
6b - 9
Zapfe, 1950a
Trocharion albanense F. Major, 1903
Zapfe´s Fissures
Middle Badenian
MN 6a
5 - 9
Zapfe, 1950a
Bonanza
Late Badenian
MN 6b
Holec et al., 1987
Sandberg
Late Badenian
MN 6b - 7+8?
Thenius, 1952
Ursavus brevirhinus (Hofmann, 1887)
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 6
Zapfe, 1950a
Sandberg
Late Badenian
MN 6b - 7+8?
Thenius, 1952
Phocidae gen. et spec. indet.
Bonanza
Late Badenian
MN 6b
Mustelidae gen. et spec. indet.
Zapfe´s Fissures
Middle Badenian
MN 6a
Sandberg
Late Badenian
MN 6b - 7+8?
Holec & Sabol, 1996
revised
Viverrinae gen. et spec. indet.
Bonanza
Late Badenian
MN 6b
Lupták, 1999
Rodentia
Anomalomys gaudryi Gaillard, 1900
Zapfe´s Fissures
Middle Badenian
MN 6a
6 - 9
Schaub & Zapfe, 1953
Blackia miocenica Mein, 1970
Zapfe´s Fissures
Middle Badenian
MN 6a
2 - 11
Fejfar, 1974
Bransatoglis astaracensis (Baudelot, 1970)
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 9
Fejfar, 1974
Bonanza
Late Badenian
MN 6b
Cricetodon sansaniensis Lartet, 1851
Zapfe´s Fissures
Middle Badenian
MN 6a
6 - 7+8
Zapfe, 1949
Democricetodon vindobonensis (Schaub et Zapfe, 1953)
Zapfe´s Fissures
Middle Badenian
MN 6a
6
Schaub & Zapfe, 1953
Bonanza
Late Badenian
MN 6b
Democricetodon sp.
Bonanza
Late Badenian
MN 6b
4 - 9
Eomyops sp.
Zapfe´s Fissures
Middle Badenian
MN 6a
5 - 14
Fejfar, 1974
Bonanza
Late Badenian
MN 6b
Eumyarion latior (Schaub et Zapfe, 1953)
Zapfe´s Fissures
Middle Badenian
MN 6a
6
Schaub & Zapfe, 1953
Eumyarion weinfurteri (Schaub et Zapfe, 1953)
Zapfe´s Fissures
Middle Badenian
MN 6a
(4) - 6
Schaub & Zapfe, 1953
Eumyarion sp.
Bonanza
Late Badenian
MN 6b
4 - 9
Holec et al., 1987
Keramidomys carpathicus (Schaub et Zapfe, 1953)
Zapfe´s Fissures
Middle Badenian
MN 6a
(5) - 6
Schaub & Zapfe, 1953
Lartetomys cf. zapfei Mein et Freudenthal, 1971
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 6
Fejfar, 1974
Megacricetodon gregarius (Schaub, 1925)
Zapfe´s Fissures
Middle Badenian
MN 6a
7+8
Zapfe, 1949
Megacricetodon schaubi Fahlbusch, 1964
Zapfe´s Fissures
Middle Badenian
MN 6a
6
Fejfar, 1974
Megacricetodon sp.
Bonanza
Late Badenian
MN 6b
4 - 9
Microdyromys miocaenicus (Baudelot)
Zapfe´s Fissures
Middle Badenian
MN 6a
?
Fejfar, 1974
Miodyromys hamadryas F. Major, 1899
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 7+8
Zapfe, 1949
Muscardinus (Eomuscardinus) sansaniense (Lartet, 1851)
Zapfe´s Fissures
Middle Badenian
MN 6a
6 - 7+8
Fejfar, 1974
Myoglis larteti Baudelot, 1965
Zapfe´s Fissures
Middle Badenian
MN 6a
?
Fejfar, 1974
Neocometes brunonis Schaub et Zapfe, 1953
Zapfe´s Fissures
Middle Badenian
MN 6a
6 - 8
Schaub & Zapfe, 1953
Bonanza
Late Badenian
MN 6b
Spermophilinus bredai (H. von Meyer, 1848)
Zapfe´s Fissures
Middle Badenian
MN 6a
6 - 7+8
Fejfar, 1974
Bonanza
Late Badenian
MN 6b
Trogontherium (Euroxenomys) minutum (H. v. Meyer, 1838) Borský Svätý Jur
Pannonian
MN 10 - 11?
(3)4 - 13
Pipík & Holec, 1998
revised
Trogontherium sp.
Pezinok
Pannonian
MN 9
(3)4 - 13
Borský Svätý Jur
Pannonian
MN 10 - 11?
Pipík & Holec, 1998
TEMPORAL AND SPATIAL DISTRIBUTION OF MIOCENE MAMMALS 275
Gliridae gen. et spec. indet.
Bonanza
Late Badenian
MN 6b
Sciuridae gen. et spec. indet.
Bonanza
Late Badenian
MN 6b
Rodentia gen. et spec. indet.
Borský Svätý Jur
Pannonian
MN 10 - 11?
Pipík & Holec, 1998
revised
Perissodactyla
Aceratherium sp.
Sandberg
Late Badenian
MN 6b - 7+8? 9 - 10
Thenius, 1952
Anchitherium aurelianense Cuvier, 1812
Sandberg
Late Badenian
MN 6b - 7+8? 3 - 10
Thenius, 1952
Brachypotherium sp.
Dúbravská hlavica
Sarmatian
MN 7+8
6 - 9
Holec & Sabol, 1996
revised
Chalicotherium grande (Blainville, 1849)
Zapfe´s Fissures
Middle Badenian
MN 6a
5 - 7
Zapfe, 1976
Dicerorhinus stenheimensis (Jager, 1839)
Zapfe´s Fissures
Middle Badenian
MN 6a
7 - 9
Zapfe, 1949
Sandberg
Late Badenian
MN 6b - 7+8?
Thenius, 1952
Haploaceratherium cf. tetradactylum (Lartet, 1851)
Zapfe´s Fissures
Middle Badenian
MN 6a
6
Zapfe, 1949
"Hipparion" s. s.
Borský Svätý Jur
Pannonian
MN 10 - 11?
11 - 13
Pipík & Holec, 1998
revised
Hippotherium primigenium H. v. Meyer, 1829
Pezinok
Pannonian
MN 9
9 - 10
Holec, 1981
revised
Topoľčany-Kalvária
Pannonian
MN 9 - 10
Holec, 1981
revised
Tapirus telleri Hofmann, 1893
Sandberg
Late Badenian
MN 6b - 7+8? 6
Thenius, 1952
Rhinocerotidae gen. et spec. indet.
Zapfe´s Fissures
Middle Badenian
MN 6a
Zapfe, 1949
Nováky - Mier Mine
Late Badenian -
MN 7+8
Early Sarmatian
revised
Proboscidea
Deinotherium laevius (Jourdan)
Sandberg
Late Badenian
MN 6b - 7+8? ?
Thenius, 1952
Prodeinotherium bavaricum (H. v. Meyer, 1831)
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 9
Zapfe, 1954
Zygolophodon turicensis (Schinz, 1824)
Zapfe´s Fissures
Middle Badenian
MN 6a
3b - 10
Zapfe, 1954
Sandberg
Late Badenian
MN 6b - 7+8?
Thenius, 1952
Bonanza
Late Badenian
MN 6b
Holec et al., 1987
Nováky - Mier Mine
Late Badenian -
MN 7+8
Early Sarmatian
gen. indet. grandincisivus (Schlesinger, 1917)
Topoľčany-Kalvária
Pannonian
MN 9 - 10
7+8 - 10
Silnický, 1930
revised
Proboscidea gen. et spec. indet.
Košice-Bankov
Sarmatian
MN 7+8
Fejfar et al., 1987
Sirenia
Halitherium cordieri Christol
Sandberg
Late Badenian
MN 6b - 7+8? 7+8?
Thenius, 1952
Metaxitherium sp.
Sandberg
Late Badenian
MN 6b - 7+8? 7+8?
Thenius, 1952
Thalatosiren petersi (Abel)
Sandberg
Late Badenian
MN 6b - 7+8? 7+8?
Thenius, 1952
Artiodactyla
Albanohyus pygmaeum (Depéret, 1892)
Sandberg
Late Badenian
MN 6b - 7+8? 4 - 9
Thenius, 1952
Aureliachoerus aurelianensis (Stehlin, 1899)
Zapfe´s Fissures
Middle Badenian
MN 6a
3+4
Zapfe, 1983
Conohyus simorrensis (Lartet, 1851)
Sandberg
Late Badenian
MN 6b - 7+8? 5 - 9
Thenius, 1952
Dicrocerus elegans Lartet, 1837
Zapfe´s Fissures
Middle Badenian
MN 6a
5 - 6
Zapfe, 1949
Sandberg
Late Badenian
MN 6b - 7+8?
Thenius, 1952
Dorcatherium naui Kaup, 1834
Sandberg
Late Badenian
MN 6b - 7+8? 4 - 10
Thenius, 1952
Dorcatherium vindobonense H. v. Meyer, 1846
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 6(9)
Zapfe, 1949
Sandberg
Late Badenian
MN 6b - 7+8?
Meyer, 1846
Eocetrus sp.
Sandberg
Late Badenian
MN 6b - 7+8? ?
Thenius, 1952
Eotragus haplodon (H. v. Meyer)
Zapfe´s Fissures
Middle Badenian
MN 6a
?
Zapfe, 1949
Sandberg
Late Badenian
MN 6b - 7+8?
Thenius, 1952
Heteroprox larteti (Filhol, 1891)
Zapfe´s Fissures
Middle Badenian
MN 6a
5 - 7+8
Zapfe, 1949
Sandberg
Late Badenian
MN 6b - 7+8?
Thenius, 1952
Hyotherium soemeringi H. v. Meyer, 1829 sive 1934
Zapfe´s Fissures
Middle Badenian
MN 6a
4 - 6(9)
Zapfe, 1983
Sandberg
Late Badenian
MN 6b - 7+8?
Thenius, 1952
Lagomeryx parvulus (Roger, 1898)
Sandberg
Late Badenian
MN 6b - 7+8? 4 - 6
Thenius, 1952
Bonanza
Late Badenian
MN 6b
Holec et al., 1987
Palaeomeryx kaupi H. v. Meyer, 1834
Sandberg
Late Badenian
MN 6b - 7+8? 4 - 6
Thenius, 1952
Palaeomeryx magnus Lartet, 1851
Zapfe´s Fissures
Middle Badenian
MN 6a
5 - 6
Zapfe, 1993
Sandberg
Late Badenian
MN 6b - 7+8?
Thenius, 1952
Taucanamo sansaniense (Lartet, 1851)
Sandberg
Late Badenian
MN 6b - 7+8? 5 - 6
Thenius, 1952
Listriodontinae gen. et spec. indet.
Sandberg
Late Badenian
MN 6b - 7+8?
Thenius, 1952
revised
Bovidae (Boselaphini) gen. et spec. indet.
Sandberg
Late Badenian
MN 6b - 7+8?
Thenius, 1952
revised
Artiodactyla gen. et spec. indet.
Borský Svätý Jur
Pannonian
MN 10 - 11?
Pipík & Holec, 1998
revised
Cetacea
Mesocetus hungaricus Kadic
Sandberg
Late Badenian
MN 6b - 7+8? 7+8?
Holec & Sabol, 1996
Mesocetus sp.
Glavica
Late Badenian
MN 6b - 7+8?
Holec, 2001
Schizodelphis cf. sulcatus (Gervais, 1861)
Sandberg
Late Badenian
MN 6b - 7+8? 7+8?
Holec & Sabol, 1996
Mammalia
Mammalia gen. et spec. indet.
Glavica
Late Badenian
MN 6b - 7+8?
Holec & Sabol, 1996
of some Slovak sites where the fossils of the Miocene mam-
mals have been found.
The oldest fossil findings of mammals in Slovakia were de-
scribed from the Zapfe’s fissures locality. However, the more
exact determination of their age is more or less problematical,
because the biostratigraphical range of the most taxa of this
terrestrial mammal assemblage is mainly from MN 5 to MN 8
biozones. On the other hand, the typical taxa of the MN 6 bio-
zone (e.g. Dinosorex zapfei Engesser, 1975 (Ziegler 1999),
Eumyarion latior (Schaub et Zapfe, 1953), E. weinfurteri
(Schaub et Zapfe, 1953), Democricetodon vindobonensis
(Schaub et Zapfe, 1953) (Kälin 1999), Keramidomys carpathi-
cus (Schaub et Zapfe, 1953) (Engesser 1999), Megacricetodon
schaubi Fahlbusch, 1964 (Kälin 1999) and others) are domi-
nating. In spite of that, some scientists (e.g. Rögl 1999) con-
template the dating of this locality to the end of the MN 5 bio-
zone. Their conclusions are especially based on the findings of
the primate species Pliopithecus (Epipliopithecus) vindobon-
ensis a representative of pliopithecids which migrated to Eu-
rope from Africa during the period of the MN 5 biozone. How-
ever, others (Köhler et al. 1999) date the fossil remnants of the
pliopithecids from these fissures to the MN 6 biozone! Here-
by, the rodent fauna of this site is distinctly younger than that
from the MN 5 sites (e.g. Langenmoosen, Franzensbad, Teir-
itzberg, Ober Gänserndorf and others). It corresponds well
with the opinion of Fejfar (1974, 1990, 1997). He dates the lo-
cality on the basis of the fossil rodent findings to the upper
part of the Middle Badenian and excludes its dating to the MN
5 biozone (pers. commun.). An unanswered question is the ex-
act species determination of some mammal fossils which do
not fall into the assumed age of the locality by their biostrati-
graphic range. These findings have been determined as Am-
Table 3: Continued.
276 SABOL and HOLEC
phechinus intermedius (Gaillard, 1899) (MN 7+8), Urotrichus
dolichochir (Gaillard, 1899) (MN 7+8), Megacricetodon gre-
garius (Schaub, 1925) (MN 7+8), Dicerorhinus steinheimen-
sis (Jager, 1839) (MN 7—9), Aureliachoerus aurelianensis
(Stehlin, 1899) (MN 3—4), and others (Table 3). It is not out of
question that the fossils are either incorrectly determined or
their original biostratigraphic range is larger. In spite of these
discrepancies, we can conclude from the lithological circum-
stances that the Zapfe’s fissures locality is older than the Sand-
berg site which is dated to the Late Badenian. On this basis,
the locality is dated to the lower part of the MN 6 biozone (as
MN 6a), whereas the dating to the MN 5 biozone can be ex-
cluded by the different character of the index small mammali-
an taxa.
The Sandberg site is another more or less problematic local-
ity. According to the latest researches (Baráth et al. 1994), the
marine sediments of the Sandberg Member have been deposit-
ed during the transgressive cycle. In spite of this, some opin-
ions about the regression origin of these marine deposits are
still always occurring in some papers (e.g. Rögl 1999). The
transgression started in the Late Badenian period (14—13 Ma
in sense of Rögl 1998), which is in good agreement with the
age obtained on the basis of the Sr-dating of some Sandberg
mollusc shells (13.6—13.5 Ma; Kováč et al. 1994). From the
mammal-zones point of view, this time level corresponds to
the end of the MN 6 biozone (Table 1). On the other hand, the
findings of nannoplankton suggest the NN 6 biozone (Grigor-
ovič et al., in press). After Steininger (1999), this nannoplank-
Table 4: Comparison of opinions on the dating of single Middle to
Late Miocene MN-zones (modified after Bernor et al. 1996; van
Dam 1997; Kempf et al. 1997; Mein 1999; Rögl 1999; Steininger
1999; Steininger et al. 1996).
ton biozone corresponds to the final phase of the MN 6 bio-
zone, but mainly to the relevant part of the MN 7+8 biozone.
Thus, a discrepancy exists between the age of this site deter-
mined on the basis of nannoplankton fossils and the age deter-
mined on the basis of the terrestrial mammals. From the mam-
mal-zones point of view, the dating of the locality to the MN 6
biozone (Fejfar 1990) is especially supported by the findings
of primate species Griphopithecus suessi Abel, 1902 (G. dar-
wini (Abel, 1902) after Andrews et al. 1997) (African immi-
grant) and cervid species Dicrocerus elegans Lartet, 1837
(Asian immigrant). The fossils of the Sandberg suid, formerly
determined to the species Listriodon lockharti, probably be-
long either to the species Bunolistriodon lockharti (Pomel,
1848), which is typical for the MN 4—5 biozones (Hünermann
1999), or to the other species of the genus Listriodon (e.g. L.
splendens Meyer, 1846, MN 6—9). As in the case of the
Zapfe’s fissures locality, some other problematic findings of
terrestrial mammals have been found here too (Table 3). They
belong mainly to the taxa Aceratherium sp. (typical genus for
the MN 9—10 biozones) and Dicerorhinus steinheimensis (MN
7—9). Besides the terrestrial mammals, fossil seals, sirens,
whales and dolphins are also known from this site. However,
these animals are probably younger (MN 7+8?) than the ter-
restrial ones. It is not out of question that the whole problem of
locality age is caused by the redeposition of older osteological
remnants of terrestrial mammals (MN 6) into younger marine
sediments (NN 6, MN 7+8) during the transgressive period. In
this time, the up-grading sea probably attacked the karst forms
(fissures) which could serve as natural traps in the time of the
MN 6 biozone (Table 2).
The marine sandy sediments of the Sandberg Member are
widespread over the larger territory of Devínska Kobyla Hill.
Besides the Wait Quarry and Glavica localities, they also fill
the fissure in the Bonanza locality where terrestrial mammals
and seals together with other vertebrates have been found. The
terrestrial vertebrate assemblage of this site is very similar to
that of Zapfe’s fissures (especially mammals; Table 3). How-
ever, on the basis of the lithological circumstances we can
conclude that the Bonanza fissure is younger than the Zapfe’s
fissures. It is also possible that the site is older than the Sand-
berg locality (Holec et al. 1987). Possibly, the Sandberg and
Bonanza represent two stages of the transgressive event.
Whereas the Sandberg deposits represent probably the top to
final phase of the transgression, the sediments of the Bonanza
can represent the initial phase of this Late Badenian marine
event. From this point of view, the Bonanza site is dated to the
upper part of the MN 6 biozone only.
The osteological remnants of the rhinoceros from the
Dúbravská hlavica site belong to the genus Brachypotherium
(Holec & Sabol 1996). Two species of this genus are known in
the Miocene of Europe – older B. brachypus (Lartet, 1837)
(MN 6 to MN 8) and younger B. goldfussi (Kaup, 1834) from
the MN 9 biozone (Heissig 1999). Because the rhinoceros fos-
sils have been not determined to the species, the age of the lo-
cality can be considered on the basis of the finding circum-
stances only. The overlying clayey sediments, in which
mammal fossils have been found, are younger than the under-
lying sandy deposits of the Late Badenian and their age is
probably the Sarmatian. On account of that, the rhinoceros
TEMPORAL AND SPATIAL DISTRIBUTION OF MIOCENE MAMMALS 277
7+8 (Dúbravská hlavica, Nováky-Mier Mine, and Košice-
Bankov), MN 9—10 (Pezinok and Topo čany-Kalvária), and
MN 10—11? (Borský Svätý Jur).
Altogether more than 100 mammalian taxa have been de-
scribed from the Slovak Miocene sites. Besides the terrestrial
mammals, the marine ones have been found in four sites too
(Bonanza, Sandberg, Wait Quarry, and Glavica). The presence
of all these mammal assemblages in the sites is a result of pa-
leogeographical and paleoenvironmental changes in the terri-
tory of the Central Paratethys in the time from the Middle to
Late Miocene. The fossil mammals are evidence of repeated
migration waves between Eurasia and Africa.
The open problem is the more exact determination of the
age of some localities. This is mainly caused by the absence of
some key fossils and the presence of taxa with a wide bios-
tratigraphical range. The exact age of these problematic sites
probably be found only by the magnetostratigraphy of their
fossiliferous sediments.
Acknowledgment: The authors are indebted to the Grant
Agency for Science, Slovakia (Project No. 1/6192/99; Caino-
zoic Vertebrates of the Vienna Basin and some other localities
in Slovakia) and the European Science Foundation, EU (Eeden
Grant No. 2001/04/0143) for financial support. We also thank
especially O. Fejfar, F.F. Steininger, and J. Michalík for criti-
cal reading of the paper’s first version and for their useful
comments. We also wish to thank to Mrs. Eva Petríková for
some figures.
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estimated age of the Hippotherium primigenium first occur-
rence at the base of the Pannonian C zone in the Vienna Basin
with the age of 11.2 Ma (Rögl & Daxner-Höck 1996). Howev-
er, the oldest specimens of this equid species in Slovak territo-
ry have been found at the Pezinok site in the Danube Basin
where the clayey and sandy sediments have been dated to the
upper part of the E zone (in sense of Papp 1951) (Fordinál
1997; Kováč et al. 1998; Baráth et al. 1999; Pipík 2000). The
age of the Pannonian E zone in the Vienna Basin is determined
at 8.9 to 9.7 Ma (Rögl et al. 1993). It is correlated with the MN
10 biozone (Rögl 1996), whereas the age of the E zone in the
Danube Basin is more than 9.88 Ma (upper part of the Early
Pannonian) (Fordinál et al. 2001) which corresponds to the up-
per part of the MN 9 biozone (Table 1). On the basis of that,
the mammal findings from Pezinok are dated to the upper part
of the MN 9 biozone (Table 2).
On the basis of both invertebrate and vertebrate fossils, the
Borský Svätý Jur locality was dated to the period from the
Pannonian to the Early Pontian (Pipík & Holec 1998). Where-
as the most of the ostracod findings belong to the Early Pan-
nonian E zone (in sense of Jiříček 1985), the mammal fossils
indicate the younger period. Because the total biostratigraphic
range of the mammal taxa from the site is relatively wide
(from MN 9 to MN 13) (Table 3), the finding of the Eomellivo-
ra wimani (Lupták 1995a) is significant for the determination
of their age. This large mustelid species lived in the territory of
Europe during the Early Turolian only (MN 11 to MN 12 bio-
zones; Ginsburg 1999). On this basis, the fossil mammals
from the Borský Svätý Jur locality are preliminarily dated to
the MN 11 biozone (Turolian, Middle Pannonian), but it is not
excluded that they are older (e.g. according to Joniak (pers.
commun.) new unpublished fossils of rodents from this site in-
dicate the MN 10 biozone).
The absence of the mammal elements in the younger periods
of the Late Miocene in Slovak territory could also be caused
by the gradual aridization of the environmental conditions,
which probably caused the Messinian crisis.
Conclusion
So far, findings of Miocene mammals are known from the
sediments of eleven localities in the territory of Slovakia. They
cover a time span from the Middle Badenian to the Pannonian.
The Miocene mammals have been found at sites on the NE
margin of the Vienna Basin, the northern part of the Danube
Basin, and from the northern margin of the Transcarpathian
(East Slovak) Basin. On the basis of new data and correction
of older information, single localities can be dated to the MN-
zones as it follows: MN 6a (Zapfe’s fissures), MN 6b (Bonan-
za), MN 6b—7+8? (Sandberg, Wait Quarry, and Glavica), MN
278 SABOL and HOLEC
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