www.geologicacarpathica.sk
GEOLOGICA CARPATHICA, OCTOBER 2010, 61, 5, 419—436 doi: 10.2478/v10096-010-0025-0
Introduction
Sarmatian continental faunas have rarely been reported from
Austria. Moreover, most of the known localities have only
yielded fossils of large mammals (e.g. Thenius 1960; Mottl
1970). Discovery of a terrestrial fauna from the Sarmatian de-
posits near Gratkorn (clay pit St. Stefan, SE Austria, Styria;
Gross et al. 2007a; Harzhauser et al. 2008; Prieto et al. in print
a and b) is thus essential for our understanding of the evolu-
tion of the late Middle Miocene faunas in the circum-Alpine
and Paratethyan realms: 1) Small and large mammals, as well
as a large spectrum of vertebrate groups (fishes, amphibians,
reptiles; Gross et al. 2009a,b) co-occur in this rich assem-
blage; 2) The deposits are fairly well dated as late Middle
Miocene on the basis of the invertebrate fauna (Gross et al.
2007a; Harzhauser et al. 2008).
Cricetids are often the most useful rodent group regarding
Middle Miocene high-resolution biostratigraphy. Their general-
ly high evolutionary rate allows for the recognition of phyloge-
netic lineages, and thus permits long-distance correlation and
high-resolution biochronological evaluation (e.g. Abdul Aziz et
al. 2008 and in print). This paper describes the cricetid rodents
from Gratkorn and discusses their biostratigraphic significance.
Geographic and geologic settings
The clay pit St. Stefan is situated in the small intramontane
Gratkorn Basin (7 km long and 3 km wide), at the northwest-
The cricetid rodents from Gratkorn (Austria, Styria): a
benchmark locality for the continental Sarmatian sensu stricto
(late Middle Miocene) in the Central Paratethys
JÉRÔME PRIETO
1,2
, MADELAINE BÖHME
1
and MARTIN GROSS
3
1
Senckenberg Center for Human Evolution and Paleoecology (HEP), Eberhard-Karls University Tübingen, Institute for Geoscience,
Sigwartstrasse 10, D-72076 Tübingen, Germany; madelaine.boehme@ifg.uni-tuebingen.de
2
Ludwig-Maximilians-University Munich, Department of Earth- and Environmental Science, Section Paleontology, Richard-Wagnerstrasse 10,
D-80333 Munich, Germany; j.prieto@lrz.uni-muenchen.de.
3
Universalmuseum Joanneum, Department of Geology & Paleontology, Weinzöttlstrasse 16, A-8045 Graz, Austria;
martin.gross@museum-joanneum.at
(Manuscript received September 22, 2009; accepted in revised form June 10, 2010)
Abstract: The recent discovery of a terrestrial vertebrate assemblage in the clay pit St. Stefan at Gratkorn (Austria,
Styrian Basin) is of major importance for our understanding of the evolution of late Middle Miocene mammal assem-
blages in the Paratethys realm. The cricetid rodent assemblage includes four species: Megacricetodon minutus Daxner,
1967, Democricetodon sp. nov. (sensu Kälin & Engesser 2001), Eumyarion sp., and “Cricetodon” fandli sp. nov. The
latter species belongs to the “Cricetodon” fandli-C. klariankae Hír, 2007 lineage, which allows for a long-distance corre-
lation with other late Middle Miocene/earliest Late Miocene European localities. The biostratigraphic conclusions drawn
from the study of the fossils from Gratkorn concurs with the age estimates based on regional geology, paleomagnetic
measurements, and the gastropod-based biostratigraphy at the base of the Late Sarmatian s. str. (late Serravallian, latest
Volhynian), around 12—12.2 Ma (Chron C5An.1n) ago.
Key words: Middle Miocene, Paratethys, Gratkorn Basin, bioprovince, long-distance correlations, Mammalia, Rodentia.
ern margin of the Styrian Basin (Fig. 1). Although outcrops
and biostratigraphic tiepoints are rare, recent studies of the re-
gional geology have produced a detailed correlation with the
stratigraphic framework of the open Styrian Basin.
The poorly sorted silts located at the base of the clay pit
are rich in mammal, amphibian, and reptile remains, as well
as terrestrial molluscs (Harzhauser et al. 2008; Gross et al.
2009a,b). This vertebrate-bearing horizon rests upon the
“Gratkorn Gravel”, which is correlated with the “Mid-”Sar-
matian so-called “Carinthian Phase” (see Harzhauser & Piller
2004) that corresponds to a widely recorded regressional
event around the Early/Late Sarmatian boundary. Biostrati-
graphic data (molluscs, foraminifers, ostracods) obtained
from the under- and overlying strata, together with the gas-
tropod fauna of the vertebrate-bearing horizon, indicate an
early Late Sarmatian age (Upper Ervilia Zone, Porosononion
granosum Zone) for the Gratkorn fauna. The recorded nor-
mal polarity of the mined pelites in the hanging wall of this
bone-bed appears to correlate with Chron C5An.1n (ca.
12.16—12.07 Ma according to Hüsing et al. 2007). This age
estimate is in strong agreement with the astronomically-
tuned age of the Early/Late Sarmatian boundary (ca. 12.2 Ma;
Lirer et al. 2009).
Although the depositional environment is a paleosol, the
vertebrate remains are exceptionally well preserved, some-
times even articulated. The occurrence of almost complete ro-
dent skulls is especially noteworthy.
For detailed information on the regional geological context,
and the section and taphonomy of the deposits exposed in the
420
PRIETO, BÖHME and GROSS
St. Stephan clay pit, refer to Gross et al. (2007a,b), Gross
(2008), and Harzhauser et al. (2008).
Material and methods
The fossils from Gratkorn are housed in the collections of the
Universalmuseum Joanneum in Graz (Geologie & Paläontolo-
gie, abbreviated UMJG) and the Bayerische Staatssammlung
für Paläontologie und Geologie in Munich (abbreviated BSPG).
Measurements were taken with an occular micrometer and are
indicated in mm. Length and width were measured at right an-
gles. The baselines used are the lingual border in lower molars
and labial border in the M1 and M2. The baseline for the M3 is
the anterior border. The terminology used in the description of
the molars follows Freudenthal et al. (1994), while the terminol-
ogy for the molar planation follows Hershkovitz (1967: fig. 4).
The length of the mesoloph(id)s is understood as follows:
‘long’, ‘medium’ and ‘short’ describe lengths either longer,
similar or shorter, respectively, than half the length of the valley
(mesosinusid and mesosinus). All teeth in the illustrations are
shown in left orientation, reversing the right specimens.
Systematic paleontology
Order: Rodentia Bowdich, 1821
Family: Muridae Illiger, 1811
Subfamily: Cricetodontinae Schaub, 1925
Genus: Cricetodon Lartet, 1851
D i a g n o s i s ( e m e n d e d ) : de Bruijn et al. (1993: 177).
D i f f e r e n t i a l d i a g n o s i s : de Bruijn et al. (1993: 178).
T y p e s p e c i e s : Cricetodon sansaniensis Lartet, 1851.
Other species included in Cricetodon: C. caucasicus Argy-
ropulo, 1938; C. meini Freudenthal, 1963; C. aureus Mein &
Freudenthal, 1971; C. jotae Mein & Freudenthal, 1971; C. al-
banensis Mein & Freudental, 1971; C. pasalarensis (Tobien,
1978); C. candirensis (Tobien, 1978); C. cariensis (Sen &
Ünay, 1979) (attributed to Byzantinia by Rummel 1998); C.
hungaricus (Kordos, 1986); C. aliverensis Klein Hofmeijer &
de Bruijn, 1988; C. versteegi de Bruijn et al., 1993; C. tobieni
de Bruijn et al., 1993; C. kasapligili de Bruijn et al., 1993; C.
bolligeri Rummel, 1995; C. jumaensis Rummel, 2001; C. en-
gesseri Rummel & Kälin, 2003; C. soriae Hernandez Fernan-
dez et al., 2006; C. klariankae Hír, 2007.
The generic assignment of the molars from Gratkorn to
Cricetodon remains problematic. The arguments in favour of
assigning the new species from Austria to “Cricetodon” are
discussed in the following sections.
“Cricetodon” fandli sp. nov.
(Fig. 2.A—F, Fig. 3)
E t y m o l o g y : In memory of the late mayor of Gratkorn,
Elmar Fandl (†), for his financial support and interest in the
excavation campaigns.
H o l o t y p e : A fragmentary skull with left and right lower
jaws and postcranial bones of the same individual embedded
in a sedimentary block (UMJG 204.240, Fig. 3).
P a r a t y p e s : See Table 1.
M e a s u r e m e n t s : See Table 1.
T y p e l o c a l i t y : Gratkorn, clay pit St. Stefan (Styria,
Austria). Clay pit of the company Wietersdorfer & Peggauer
Zementwerke AG, 10 km northwest of the city of Graz (E
15°20
’55”, N 47°08’15”).
T y p e h o r i z o n : Basal paleosol at top of the “Gratkorn
Gravel” (Gross et al. 2007a,b), late Middle Miocene, Late
Sarmatian s. str. (Upper Ervilia Zone; Porononion granosum
Zone).
D i a g n o s i s : Medium-sized Cricetodontinae species with
moderately high cusps(ids). A very narrow anteroconid and
reduced antero- and protosinusid of the m1 are characteristic.
Ectolophs are complete in the upper molars. The anterior
metalophulid is interrupted or missing in m1. The mesoloph is
missing in the upper molars, while the mesolophid is absent to
reduced in m1 and m2.
D i f f e r e n t i a l d i a g n o s i s : The Cricetodon species lack-
ing complete ectolophs and/or having the mesoloph on the
upper molars are easily distinguished from “C.” fandli. The
earliest Hispanomys species (see discussion section for more
information), with the exception of H. bijugatus, usually pos-
sess a less well developed posterior metalophulid in m1. H. bi-
jugatus typically displays an anterior metalophulid in m1. The
overall morphology of the molars from Gratkorn is reminis-
cent of what is observed in C. klariankae from Hungary. Be-
sides the slightly larger size, the latter species differs from
“C.” fandli in the presence of a double metalophid in m1, and
the spur of the anterocone that is directed lingually and thus
not closing the anterosinus.
D e s c r i p t i o n o f t h e h o l o t y p e : The skull is free from
sediment on the anterior left side. The anterior part is pre-
served and the left molar row observable. The left mandible is
more or less in its original position, and covers the right man-
dible, the posterior part of which is still buried in the sedi-
ment. Below the horizontal ramus of the left mandible a
fragmentary bulla tympanica is preserved. Limb materials
(hand bones) are preserved above and below the skull.
A detailed description and comparison of the cranial and
postcranial material of “Cricetodon” fandli will be provided in
a forthcoming paper. Here we focus on the dental material,
which usually displays the most reliable diagnostic characters.
The tip of the incisor ends more or less on the level of the
margo alveolaris of the mandible. The lower incisors have
double parallel ridges on their ventral side; the left m1 and m2
are observable, but the m3 is almost completely covered by
sediment.
The small anteroconid in the m1 is round, the broad labial
anterolophid does not close the protosinusid, which is strongly
reduced; the lingual anterolophid is absent; the anteroconid
and protoconid are interconnected by a strong anterolophulid;
a small crest is developed on the posterior wall of the antero-
conid, but does not reach the metaconid; the protoconid hind
arm and posterior metalophulid are connected with the ectol-
ophid, forming a Y-shaped structure; the mesolophid is ab-
sent; the hypolophulid is directed forwards; the posterior part
421
MIDDLE MIOCENE CRICETID RODENTS FROM GRATKORN (AUSTRIA)
of the ectolophid is curved; the sinusid is directed backwards
and is closed by a cingulid, which extends on to the labial wall
of the hypoconid; the molar is bileveled.
In the m2 protoconid and metaconid are connected to the an-
teroconid; the labial anterolophid reaches the base of the proto-
conid, delimiting a deep and round protosinusid; the lingual
anterolophid is missing; the mesolophid is very short; the me-
sosinusid is closed by a cingulid extending to the labial wall of
the hypoconid; this cingulid is more pronounced than in the m1.
The upper incisors have a narrow longitudinal groove on
their dorsal part.
M1: The anterocone is bifid; its anterior wall is fissured
(reaching about half of the high of the cusp); the lingual anter-
oloph is absent and the massive protostyl closes the protocone
platform; the posterior spur departing from the labial lobe of
the anterocone merges with the anterior spur of the paracone,
giving an irregular trace to the ectoloph; it has the same orien-
tation as the paracone spur, which reaches the metacone; the
two crests form complete ectolophs; the ento- and mesostyls
are present; the mesoloph is reduced to a slight thickening of
the entoloph (mesocone?); the metalophule is directed back-
wards and fused with the extremely short posteroloph.
M2: The ectolophs are complete and the lingual anteroloph
is well developed; the overall morphology of the remaining
teeth is similar to that of the M1; the lingual part of the tooth is
hardly observable.
The M3 is covered by sediment on its posterior part; the
transverse labial anteroloph does not close the anterosinus.
D e s c r i p t i o n o f t h e p a r a t y p e s : m1: The teeth do
not differ in basic structure from the holotype; the connection
anteroconid-metaconid is interrupted (3 out of 5 m1) or ab-
sent; the mesolophid is very short or absent; the hypolophulid
is directed forwards, joining the ectolophid near the mesol-
ophid, if this structure is present; the broad posterolophid does
not reach the base of the entoconid; a cingulid closes the space
resulting from the constriction of the posterolophid near the
hypoconid, and delimits a small posteriolabial valley; the me-
sosinusid is directed backwards and closed by a cingulid; the
molars each possess two roots.
m2: The teeth are very similar to those seen in the type
specimen; the mesolophid is either missing or very short, but
in the latter case always directed towards the metaconid; on
some molars, the sinusid is more transverse than in the m1; the
curved posterolophid does not reach the base of the entoconid;
the small posterior valley resulting from constriction of the
posterolophid, as described for the m1, may be developed; the
molars each possess three roots.
m3: The overall arrangement of the crests and cuspids is
similar to that seen in m2; the main differences include a nar-
rower talonid, a usually longer mesolophid, and a narrow and
transverse sinusid; in one molar (Fig. 2E), the protosinusid is
strongly reduced and incorporated into the base of the labial
Fig. 1. Geographical setting and regional geology of the study area. a – Postion of the Styrian Basin (S.B.) within the Pannonian Basin
System; b – Geological sketch of the Styrian Basin; c – Simplified geological map of the north-western margin of the Styrian Basin
(from Gross et al. 2007a).
422
PRIETO, BÖHME and GROSS
wall of the protoconid, moreover, a well-developed crest di-
vides this valley; the molars each possess three roots.
M1: The molars are very similar to those of the holotype;
the mesostyl may be present or absent; the mesoloph is most
often absent but a faint thickening of the entoloph (as in the
holotype) may be present; the molars each possess 4 roots, and
a fifth small root is developed on the posterior basis of the an-
tero-labial root.
Reference Specimen
Orientation
Length
(mm)
Width
(mm)
Figure
BSPG 2008 IV 56
Maxillary with M1
left
3.20
1.97
BSPG 2008 IV 57
M1
left
2.98
1.93
BSPG 2008 IV 61
m1
left
*
1.72
UMJG 204.225
m1
right
*
1.77
UMJG 204.226
Mandible with m1
left
2.35
1.55
UMJG 204.227
m1
left
2.55
1.68
2C
BSPG 2008 IV 59
M3
right
1.88
1.82
UMJG 204.228
M3
right
*
*
BSPG 2008 IV 58
Maxillary with M3
left
1.95
1.67
BSPG 2008 IV 60
M3
right
1.93
1.63
UMJG 204.229
Maxillary with M3
right
1.87
1.73
2B
UMJG 204.230
m2
right
*
*
UMJG 204.231
m2
right
*
*
BSPG 2008 IV 62
m2
left
2.48
1.87
UMJG 204.232
m2
right
2.53
*
UMJG 204.233
m2
right
*
1.87
UMJG 204.234
m2
left
2.43
1.73
2D
BSPG 2008 IV 63
m3
right
2.2
*
BSPG 2008 IV 64
m3
right
2.32
1.73
UMJG 204.235
m3
right
2.27
1.78
UMJG 204.236
m3
right
2.25
1.62
BSPG 2008 IV 65
m3
right
*
1.68
UMJG 204.237
m3
left
2.30
1.70
2E
UMJG 204.003
Mandible with m2
right
*
*
UMJG 204. 016
Fragmentary skull with:
M1
right
3.05
*
M2
right
2.27
*
M3
right
1.73
*
M2
left
2.27
1.80
M3
left
*
1.57
UMJG 204.020
Fragmentary skull with:
M1
left
3.03
*
M2
left
2.22
*
M3
left
*
*
UMJG 204.025
Maxillary with
2A, F
M1
right
3.08
1.90
M2
right
2.43
1.78
M3
right
1.88
1.70
UMJG 204.049/2
m2
left
2.43
1.87
UMJG 204.019
Mandible with
m2
left
2.43
2.03
m3
left
2.17
1.60
UMJG 203.738
Mandible with:
m1
right
*
*
m2
right
*
*
UMJG 204.018
Maxillary with:
M1
right
2.90
1.88
M2
right
*
*
UMJG 204.029
Fragmentary skull with:
M1
left
*
*
M1
left
*
*
M2
left
*
*
M3
left
*
*
UMJG 204.050
M3
left
*
*
UMJG 204.238
Mandible with
m1
left
2.40
1.57
m2
left
2.27
1.85
m3
left
2.08
1.72
Table 1: “Cricetodon” fandli nov. sp. Material and measurements.
M2: The anterocone is triangular; the ecto-
lophs are complete and the lingual anteroloph
is well developed; the overall morphology of
the remaining teeth is similar to that seen in
M1; notable differences include a stronger cin-
gulum closing the sinus and a narrower contact
zone between the paracone spur and metacone.
M3: The transverse labial anteroloph does
not close the anterosinus; the lingual antero-
loph is short and incorporated into the anterior
wall of the protocone; protocone, paracone and
entoloph are interconnected in a Y-like struc-
ture; the spur of the paracone may be curved;
the hypocone is highly reduced; a narrow pos-
terosinus is usually present and completely
closed, otherwise the posteroloph is missing
and the metalophule very short; the cingulum
closing the sinus may be as strong as in the
M2; the metacone is indistinct; the molars each
possess four roots.
D i s c u s s i o n : The molars from Gratkorn
cannot be attributed to the following Cricet-
odontinae genera based on distinct differences
in morphology and size: Deperetomys Mein &
Freudenthal, 1971; Ruscinomys Depéret, 1890;
Plesiodipus Young, 1927; Tsaganocricetus To-
pachevsky & Skorik, 1988; Gobiocricetodon
Qiu, 1996; Lartetomys Mein & Freudenthal,
1971 (= (pars) Mixocricetodon Rummel, 1997
in Mein 2003: p. 411; Rummel pers. comm.).
Furthermore, we propose that:
1) The following morphological characters
seen in the Gratkorn material preclude assign-
ment of the material to the genus Byzantinia de
Bruijn, 1976:
– the cusp(id)s are less deeply incorporat-
ed into the loph(id)s (de Bruijn & Ünay 1996:
p. 233);
– all check teeth possess terraced to bi-lev-
eled wear surfaces (diagnostic character: de
Bruijn 1976; Sen & Ünay 1979, surface rela-
tively concave in Byzantinia);
– the molars are less distinctly hypsodont;
– the anterocone of the M1 is less deeply
fissured;
– the posterior part of the M2 is not nar-
rower than its anterior part (diagnostic charac-
ter: Rummel 1998);
– a well developed anterior metalophulid is
absent in the m1 (diagnostic character: Ünay &
de Bruijn 1984; Rummel 1998);
– secondarily the molars do not show the longitudinally
“stretched” appearance (sensu de Bruijn & Ünay 1996) of the
evolved Byzantinia species.
2) The youngest species of Hispanomys (Late Miocene; in-
cluding the type species H. aragonensis) clearly differ from
the molars from Gratkorn because of:
– the absence of the posterior metalophulid in most m1
specimens. This character is variable and double metalo-
423
MIDDLE MIOCENE CRICETID RODENTS FROM GRATKORN (AUSTRIA)
Fig. 2. Cricetid rodents from Gratkorn. All teeth shown in left orientation. A—F – “Cricetodon” fandli nov. sp.: A – right Maxillary
with M1—M3 (UMJG 204.025); B – right M3 (reversed, UMJG 204.229); C – left m1 (UMJG 204.227); D – left m2 (UMJG 204.234);
E – left m3 (UMJG 204.237); F – same specimen as in A, lingual view of M1. G—I – Democricetodon sp. nov. (sensu Kälin & Engesser
2001): G – left m1 (UMJG 204.191); H – left M1 (UMJG 204.192); I – left M3 (UMJG 204.193). J—L – Megacricetodon minutus
Daxner, 1967: J – right m1 (reversed, UMJG 204.215); K – left maxillary with M1—M2 (UMJG 204.214); L – right M3 (reversed,
UMJG 204.216). M—P – Eumyarion sp.: M – right m2 (reversed, UMJG 204.223); N – left M1 (UMJG 204.221); O – right M2 (re-
versed, UMJG 204.222); P – right m1 (reversed, UMJG 204.224).
424
PRIETO, BÖHME and GROSS
phulids may be observed in some rare Hispanomys specimens
(e.g. 1 out of 9 specimens in H. mediterraneus from Montre-
don; Aguilar 1981);
– the less bulbous cusp(id)s;
– the flat wear surface;
– the shorter posterolophid of the m2 and m3.
As a result, two genera are morphologically similar to the
Gratkorn specimens: Hispanomys (the older, Middle to earli-
est Late Miocene species) and Cricetodon. Cricetodon, a ge-
nus with a long taxonomic history, initially accomodated most
of the European fossil cricetid rodents. In the present sense,
Cricetodon regroups a wide array of large cricetids. Differ-
ences between genera therefore may not be clear, which may
lead to differences in taxonomic interpretation.
In the following analysis, and in order to simplify the com-
parisons, six groups of Middle to Late Miocene Cricetodon-
Hispanomys species are considered (the Early Miocene
Anatolian Cricetodon species, de Bruijn et al. 1993, and the
Greek C. aliverensis Klein Hofmeijer & de Bruijn, 1988, are
not taken into account because of their stratigraphic distance):
Group 1: The Middle Miocene Anatolian Cricetodon
group. Originally, these Cricetodon species were described as
Turkomys (Tobien 1978). Boon (1991) placed this taxon into
the synonymy of Cricetodon, arguing that a short posterior
Fig. 3. “Cricetodon” fandli nov. sp. Holotype (UMJG 204.240): A – specimen overview; B – Details of the fossils. 1 – Skull; 2 – Left
and right mandibles; 3 – Bulla tympanica; 4—7 – Forelimb, wrist and hand bones; C – Detailed view of dental elements. C1 – left lower
tooth row (m1—m2); C2 – left upper tooth row (M1—M3).
425
MIDDLE MIOCENE CRICETID RODENTS FROM GRATKORN (AUSTRIA)
ectoloph, a posterior or double metalophid, and the enlarged
M3/m3 also occur in some western European Cricetodon
species. The species considered by this author include C.
candirensis and C. pasalarenis. Moreover, Rummel (1998)
included the species cariensis in Byzantinia. Regarding the
clear morphological resemblance with the two other species,
the species cariensis is here included in Cricetodon and in
this group.
Group 2: C. hungaricus group from Hasznos, Sámsonháza
and Mátraszölös (Hungary, Kordos 1986; Hír & Mézáros
2002; Middle Miocene, Badenian). De Bruijn et al. (1993: p.
210) recognized a close resemblance between the Anatolian
species (especially C. candirensis) and C. hungaricus (main
difference: presence/absence of mesolophid in the m1). Based
on the figured material of C. candirensis (Tobien 1978:
figs. 7—9; Sen & Ünay 1979: figs. 7—11; de Bruijn et al. 1993:
plate 5), C. pasalarensis (Tobien 1978: figs. 1—6; Ünay 1990:
plates I—II) and C. cariensis (Sen & Ünay 1979: figs. 1—6) on
the one side, and C. hungaricus on the other (Kordos 1986:
plates I—III), additional differences can be seen: especially in
the M1 of the Anatolian Cricetodon group the labial cusps
typically are clearly standing out from the outline, whereas
the labial border is almost straight in the Hungarian species;
there is also a trend towards development of an anterior ec-
toloph in the M2 of the C. hungaricus group. Furthermore,
the Anatolian Cricetodon species (group 1) either have a
short labial anterolophid in the m1 (C. pasalarensis) or this
structure is absent.
Group 3: Cricetodon klariankae from Felsötarkany-Fel-
német (Hungary, Late Sarmatian; Hír 2007). This species dif-
fers from the other Cricetodon groups in having a very narrow
anteroconid and a reduced antero- and protosinusid of the m1
(Hír 2007: plate I, fig. 16). In addition, this group differs from
the C. hungaricus group by the relatively longer M2 and
shorter M3.
Group 4: Early Hispanomys species (Middle Miocene to
earliest Late Miocene) from Spain and France. This group re-
fers to H. aguirrei, H. dispectus, H. castelnovi, H. daamsi, H.
sp. from Can Vila (Casanovas-Vilar 2007; Casanovas-Vilar et
al. 2008, now assigned to H. cf. aguirrei, Casanovas-Vilar
pers. comm.), H. bijugatus, and H. decedens. The species
lavocati, which has long remained uncertain with regard to
its generic status, is today included in Hispanomys (Casano-
vas-Vilar 2007; López Guerrero et al. 2008: p. 260; López-
Anton
~anzas & Mein 2009). The members of group 4 differ
from the late Hispanomys species by having less hypsodont
and reduced longitudinally “stretched” molars.
Group 5: Brachiodont Cricetodons characterized by ectol-
ophs that are very short or absent (Middle Miocene). This
group refers to the western European C. meini, C. aureus, C.
jotae, C. albanensis, C. sansaniensis, and C. soriae, the cen-
tral European C. bolligeri, C. jumaensis, and C. engesseri (ex-
cept Cricetodon sp. from Nebelberg TGL in Rummel & Kälin
2003 (Hispanomys sp. in Kälin & Kempf 2009); Boon 1991;
Bolliger 1992, 1994; Kälin 1993; Rummel 1995, 2000, 2001;
Rummel & Kälin 2003; Prieto 2007), and C. meini from Aus-
tria (Daxner-Höck 2003) and Greece (de Bruijn et al. 1993).
The rare findings of Cricetodon sp. from Anwil correspond
taxonomically to a new Cricetodon from the German faunas
Giggenhausen and Kleineisenbach, which also contain Dep-
eretomys hagni (Prieto 2007).
Group 6: Cricetodon caucasicus from Belometchetskaya
(Middle Miocene, Tshokrakian respectively/or more precisely
Badenian, Langhian; Argyropulo 1938; Sen & Ünay 1978;
Pickford et al. 2000). The species is characterized by M1 with
undivided anterocone and no ectolophs.
Comparison of the molars from Gratkorn with these groups
The groups with very short or missing ectolophs are dis-
tinctly different from the species from Gratkorn (groups 5, 6,
both attributed to Cricetodon). In the other groups, the ecto-
lophs are well developed. The transition from Cricetodon to
Hispanomys in Western Europe, and likewise from Crice-
todon to Byzantinia in the eastern Mediterranean, is probably
complicated by parallel evolution (de Bruijn 1976; Sen &
Ünay 1979; Ünay 1980; Ünay & de Bruijn 1984: p. 125;
Casanovas-Vilar 2007: p. 226). Moreover, the generic attri-
bution of the species also depends on whether a morphologi-
cal or phylogenetic approach to taxonomy is used (van de
Weerd 1976: p. 104). Thus, the generic assignment of the
early members of these three Cricetodontinae is not well-de-
fined (de Bruijn et al. 1993: 177). De Bruijn et al. (1993)
emended the diagnosis of Cricetodon, but did not consider
Hispanomys in their differential diagnosis. Consequently,
the differences between Hispanomys and Cricetodon rely ex-
clusively on the differential diagnosis of Hispanomys in van
de Weerd (1976).
One special feature of the m1 from Gratkorn is the very
short or missing anterior metalophulid, while the posterior
metalophuid is well developed. In the early Hispanomys
(group 4), the posterior metalophulid is usually absent in the
m1, while it is well developed in the members of group 1.
This character does not allow for the unambiguous separation
of the two genera due to the morphological variability of this
structure. In fact, 13 % of the m1 from Pa alar (C. pasalare-
nis) possess a complete anterior metalophulid, and in 34 % of
the molars this crest is incomplete, whereas in the rest of the
sample only the posterior metalophulid is present (Ünay
1990). Similarly the posterior metalophulid is present in 25 %
of the H. aguirrei teeth from Escobosa (Sese 1977), and dou-
ble metalophulids occur in most of the molars of H. bijugatus
(Mein & Freudenthal 1971; López-Anton
~anzas & Mein 2009;
Casanovas-Vilar pers. comm.).
The morphology of the m1 from Gratkorn is characterized
by the extremely narrow anteroconid and reduced antero- and
protosinusids. However, this character is also found in C.
klariankae (see Hír 2007: plate 1). Hír (2007) excluded this
species from Hispanomys based on the following characters:
the presence of a well-developed anterolophulid in m1; the
presence of a posterior metalophulid; the absence of reduction
of the hypoconid and posterolophid in m3. A well-developed
anterolophulid is actually present in the m1 of H. daamsi, H.
castelnovi, H. aguirrei, H. decedens and H. bijugatus. More-
over, the hypoconid and posterolophid are only weakly re-
duced in the species of the group 4. Finally, the anterior and
posterior metalophulids are equally well developed in all C.
klariankae (group 3) m1, while this feature is rather rarely
426
PRIETO, BÖHME and GROSS
seen in most of the early Hispanomys. However, this differ-
ence is not sufficient to clearly exclude the Western European
genera because it is also common in H. bijugatus (see above
and etymology of the species).
C. candirensis and C. cariensis (group 1) both possess a
very simple anteroconid and lack anterolophids.
Based on the formal diagnoses for Cricetodon and Hispano-
mys (Mein & Freudenthal 1971: p. 19; van de Weerd 1976:
106; de Bruijn et al. 1993), the Gratkorn material could be as-
signed to either of these genera, even though the teeth clearly
differ from the type species of both genera. In the light of
these problems, we prefer to include the new Austrian species
in the genus “Cricetodon” in quotation marks because de
Bruijn et al. (1993) pointed out that the majority of the
Cricetodon m1 have a double metalophulid or posterior me-
tatophulid, and the lower incisors have two ridges on their
ventral part (see Flynn et al. 1985). However, the co-occur-
rence of a complete ectoloph in the upper molars and groove
in the upper incisors argues against assignment to Crice-
todon in the restricted sense ( = Cricetodon (Cricetodon) of
Mein & Freudenthal 1971: p. 17, as group 5 in this paper).
Phylogenic relationships
The record of Cricetodon from the North Alpine Foreland
Basin
As mentioned above, the Western European and Swiss/Ger-
man Cricetodon species are morphologically clearly different
from “C.” fandli sp. nov. However, the unnamed high-
crowned Cricetodon from Nebelberg TGL III, and probably
also documented in Petersbuch 14 (Rummel 2000; Rummel &
Kälin 2003 (assigned to Hispanomys by Kälin & Kempf 2009,
without comments); material not seen), possesses lower mo-
lars that, with regard to size, range between the molars of “C.”
fandli and C. klariankae, whereas the upper molars are as big
as those of the Hungarian species. These molars probably do
not belong to either “C.” fandli or C. klariankae based on the
absence of complete ectolophs.
The Austrian record of Cricetodon
Cricetodon has rarely been reported from Austria; the
records to date include Mühlbach, Grund (C. meini, 15.1 Ma
Daxner-Höck 2003; Harzhauser et al. 2003) and Bullendorf
(Pannonian “Zone B/C”, Daxner-Höck 1996, 2004). Gudrun
Daxner-Höck (Vienna) kindly provided images of the speci-
mens from Bullendorf. A single M3 is similar to “C.” fandli,
whereas the other molars cannot be attributed to the new spe-
cies, and probably do not belong to Cricetodon. As a result,
we cannot rule out that two large cricetid species occur in the
Bullendorf site.
The Central and Eastern European and Anatolian record of
Cricetodon
Pannonian Basin. The oldest representative of the genus is
C. hungaricus, reported from the Badenian of Hasznos and
Szentendre (Kordos 1986). A descendant of C. hungaricus oc-
curs in the Middle Badenian of Sámsonháza (Hír et al. 1998;
Hír & Meszáros 2002). From the Late Badenian of
Mátrazölös, a small sample of Cricetodon sp. has been de-
scribed (Hír & Kókay 2004). The upper molars of this fossil
have a funnel structure and fit within the upper segment of the
size range of C. hungaricus. A few teeth have been reported
from the Early Sarmatian (Mohrensternia Zone, Early Vol-
hynian) of Tă ad (Hír et al. 2001). The identification of Crice-
todon sp. from this locality remains problematic. The molars
are distinctly larger than those of “C.” fandli, in spite of the
fact that the locality is slightly older than Gratkorn. The ab-
sence of m1 in the specimen from Tă ad does help in assign-
ing Cricetodon sp. to either “C.” fandli sp. nov., C. klariankae
or the C. hungaricus lineages. Cricetodontinae have not been
recorded for the early Late Sarmatian (Late Volhynian) locali-
ty Tau (Feru et al. 1979), wheras the slightly older earliest
Late Sarmatian locality Comăne ti 1 contains Hispanomys cf.
lavocati and H. cf. bijugatus, which are briefly described but
not illustrated (Feru et al. 1980; material not seen). These two
taxa are distinguished from one another by the presence of a
double metalophulid in the m1 of H. cf. bijugatus. However,
this feature may be variable as demonstrated in the molar pop-
ulation of Cricetodon pasalarensis (Ünay 1990) and H. bijug-
atus from La Grive (Casanovas-Vilar pers. comm.), which
suggests that all specimens from Comăne ti 1 belong to one
species. The m1 from Comăne ti 1 are somewhat larger than
the specimens from Gratkorn (L m1 H. cf. lavocati: 2.72 mm,
n = 1; L m1 H. cf. bijugatus: 2.69 mm, n = 1; Feru et al. 1980),
and thus are within the size range of C. klariankae. A final de-
cision, however, cannot be reached at present. The Late Sar-
matian record of Cricetodon (C. klariankae) is better from
Felsötárkány-Felnémet (Hír 2006, 2007). The abundance of
this genus decreases in the upward direction of the profile, and
the taxon is absent from the younger strata (Hír 2004, 2006;
Hír & Kókay 2010). As a consequence, Cricetodon has not yet
been recorded for the Pannonian localities of this basin (e.g.
Comăne ti 2 – Feru et al. 1980, and Rudabánya – Kretzoi &
Fejfar 2005). “C.” fandli is morphologically close to C. klari-
ankae from Felsötárkány-Felnémet. We suggest an ancestor/
descendant relationship between the two species. If this hy-
pothesis is accurate, then the “C.” fandli—C. klariankae lin-
eage pricipally characterized by: 1) an increase in size of the
molars, 2) development of the anterior metalophulid in m1,
3) loss of the complete anterior ectoloph.
Despotovac and Kolubara Basins (Serbia). Cricetodon
meini has been reported from Mala Miliva (Despotovac Basin,
Marković 2003), Lazarevac and Bele Vode (Kolubara Basin,
Marković 2008). Moreover, Cricetodon sp. has been men-
tioned from the Sarmatian of Vracevici (Marković 2003),
from the small Kolubara Basin SSW of Belgrade, a locality
originally correlated with Anwil based on the presence of De-
peretomys hagni. Marković in fact revised his determinations
in his unpublished dissertation, and instead listed Byzantinia
bayraktepensis from Vracevici.
Southern Bohemian Massif (České Budějovice Basin).
Fejfar (1974) reported two Cricetodon cf. meini molars from
the Early Badenian of Strakonice (Abdul Aziz et al. in print).
However, the sample is too small to speculate the phylogenetic
relationships of these fossils.
427
MIDDLE MIOCENE CRICETID RODENTS FROM GRATKORN (AUSTRIA)
Turkey and Greece. The Miocene record of Cricetodontin-
ae from Anatolia is well documented (e.g. de Bruijn & Ünay
1996). We thus restrict our discussion to the Middle Miocene
forms. The Cricetodon fossils from Anatolia are morphologi-
cally close to “C.” fandli. Nevertheless, it is not possible to af-
filiate the Gratkorn species with the Anatolian lineage because
dissimilarities exist with regard to size and morphology. Fol-
lowing Ünay & de Bruijn (1984: p. 125) the transition from
Cricetodon to Byzantinia was completed when most of the m1
had developed an anterior or double metalophulid. A transi-
tional form has been described from Kalami (Ünay 1990).
The genus Byzantinia has been recorded for the latest Middle
Miocene of Turkey and Greece (e.g. de Bruijn & Ünay 1996;
Koufos 2006). If a similar evolutionary pattern is accepted for
the lineage “C.” fandli—C. klariankae, then the double metalo-
phulid-level is recognizable in Felsötárkány-Felnémet, while
the specimens from Gratkorn are more primitive.
Spain and France. The overall resemblance between “C.”
fandli and Hispanomys from the late Middle Miocene of
Spain is impressive. The transition from Cricetodon to His-
panomys is characterized by the presence of an anterior met-
alophulid in the m1 of Hispanomys. In the Valles-Penedes
Basin, H. decedens occurs at the base of the “Megacricetodon
ibericus + Democricetodon larteti” Zone, which correlates to
the lower part of the Zone G3 of the Catalayud-Daroca Basin
(Casanovas-Vilar 2007). H. daamsi probably appeared at
some time during the top of this zone, and extended into the
following M. ibericus + D. crusafonti Zone, which correlates
with the upper part of the G3. H. dispectus and H. lavocati
have been reported from this zone. Hispanomys appeared later
in the Catalayud-Daroca Basin where the association H.
aguirrei + H. lavocati has been recorded for Nombrevilla 2
(Top of the G3, Garcés et al. 2003; López Guerrero et al.
2008). All these species are characterized by a well-developed
anterior metalophulid, and thus a direct relationship with “C.”
fandli is unlikely. The double metalophulids are present in
most of the m1, but do not reach 100 % as observed in
Felsötárkány-Felnémet. One of the French species is perhaps
related to C. klariankae: H. bijugatus from the fissures infill-
ings of La Grive L3 and L5 (Mein & Ginsburg 2002).
Subfamily: Megacricetodontinae Mein & Freudenthal, 1971
Genus: Megacricetodon Fahlbusch, 1964
D i a g n o s i s : Fahlbusch (1964).
T y p e s p e c i e s : Cricetodon gregarius Schaub, 1925.
Other species included in Megacricetodon: M. minor
(Lartet, 1851); M. bourgeoisi (Schaub, 1925); M. gregarius
(Schaub, 1925); M. ibericus (Schaub, 1944); M. collongensis
(Mein, 1958); M. lappi (Mein, 1958); M. crusafonti
(Freudenthal, 1963); M. primitivus (Freudenthal, 1963); M.
bavaricus Fahlbusch, 1964; M. similis Fahlbusch, 1964; M.
minutus Daxner, 1967; M. bezianensis Bulot, 1980; M. ger-
Upper molars
M1
M2
M3
References
n Length Width n Length Width n Length Width
M. minor
FF 2/3
19
1.44
0.9
7
1.14
0.93
5 0.73 0.75 Hír
2006
M. minor
FF 2/7
1
1.43
0.94
2
1.05
0.84
Hír 2006
M. minutus
F 3/2
10
1.38
0.86 5 1.08 0.97 3
0.74
0.76 Hír 2003
M. minor
Sámsonháza 19/18
1.55
0.98 9 1.16 0.95 2
0.79
0.77 Hír & Mészáros 2002
M. minor
Matraszölös 1+2
9
1.42
0.9
3
1.09
0.87
3
0.79
0.76 Hír & Kókay 2004
M. minor
Taşad
2
1.52
0.96
Hír, Kókay & Venczel 2001
M. minutus
Inzersdorf
1
1.39
0.86
1
1.09
0.88
Daxner 1967
M. minutus
Brunn-Vösendorf
1
1.47
0.92
Daxner 1967; Daxner-Höck 2004
M. minutus
Mataschen
1 1.1
0.95
Daxner-Höck 2004
M. minutus
Richardhof-Golfplatz
2 1
0.85
1
0.65
0.65 Daxner-Höck 2004
M. minutus
Bullendorf
1
1.45
0.85
Daxner-Höck 2004
M. minor
Nebelbergweg 29
1.44
0.89 32
1.09
0.92
20
0.78
0.78 Kälin & Engesser 2001
M. crisiensis Comăneşti 1
3
1.51
0.97 5
1.16
0.99
Radulescu & Samson 1988
M. minutus
Gratkorn
1
1.44
0.88 6
1.08
0.94
2
0.82
0.80 this paper
m1 m2 m3
Lower molars
n Length Width n Length Width n Length Width
References
M. minor
FF 2/3
17
1.35
0.82 13 1.09 0.86 9 0.92 0.73 Hír
2006
M. minor
FF 2/7
1
1.41
0.87 1
1.11
0.94
Hír 2006
M. minutus
F 3/2
7
1.34
0.77 5
1.06 0.8
9 0.92 0.71 Hír
2003
M. minor
Sámsonháza 16
1.43
0.85 19 1.15 0.92 5
1.03
0.84 Hír & Mészáros 2002
M. minor
Matraszölös 1+2
10
1.37
0.83 4
1.09
0.82
1
0.87
0.76 Hír & Kókay 2004
M. minor
Taşad
2
1.45
0.84 1
1.15
0.88
Hír, Kókay & Venczel 2001
M. minutus
Inzersdorf
Daxner 1967
M. minutus
Brunn-Vösendorf
1 1.5
0.9
Daxner 1967; Daxner-Höck 2004
M. minutus
Mataschen
1
1.45
0.8
1
1.1 0.9
2
0.95
0.78 Daxner-Höck 2004
M. minutus
Richardhof-Golfplatz
2
1.33
0.75 3
1.03
0.78
1
0.75 0.6
Daxner-Höck 2004
M. minutus
Bullendorf
Daxner-Höck 2004
M. minor
Nebelbergweg 22
1.36
0.8
33
1.09
0.86
14
0.98
0.75 Kälin & Engesser 2001
M. crisiensis Comăneşti 1
1
1.46
0.93 1
1.27
1.05
2/1
1.14 0.8
Radulescu & Samson 1988
M. minutus
Gratkorn
8 1.4
0.86 4
1.15
0.90
2
0.91
0.74 this paper
Table 2: Measurements of Megacricetodon minutus Daxner, 1967 from Gratkorn, and small Megacricetodon populations from the Parat-
ethys late Middle Miocene/lower Upper Miocene (F – Felsötárkány, FF – Felsötárkány-Felnémet).
`
`
428
PRIETO, BÖHME and GROSS
manicus Aguilar, 1980; M. gersii Aguilar, 1980; M. sinensis
Qiu et al., 1981; M. lopezae Garcia Moreno, 1986; M. roussi-
lonensis Aguilar et al., 1986; M. crisiensis Radulescu & Sam-
son, 1988; M. rafaeli Daams & Freudenthal, 1988; M.
fournasi Aguilar, 1995; M. lemartinelli Aguilar, 1995; M. pu-
sillus Qiu, 1996; M. fahlbuschi Aguilar et al., 1999; M. wuae
Aguilar et al., 1999; M. lalai Aguilar et al., 1999; M. andrewsi
Pelaez-Campomanez & Daams, 2002; M. tautavelensis Laz-
zari & Aguilar, 2007; M. aunayi Lazzari & Aguilar, 2007; M.
yei Bi et al., 2008.
M. aguilari Lindsay, 1988; M. sivalensis Lindsay, 1988; M.
daamsi Lindsay, 1988 and M. mythikos Lindsay, 1988 are ex-
cluded from Megacricetodon by Wessels (1996).
M. debruijni Freudenthal, 1968 is considered as junior syn-
onym of M. minutus by Joniak (2005).
Megacricetodon minutus Daxner, 1967
(Fig. 2J—L)
D i a g n o s i s : Daxner (1967).
T y p e l o c a l i t y : Inzersdorf (Austria).
A g e : Late Miocene, Pannonian “Zone E” (sensu Papp 1951).
M a t e r i a l : 1 mandible with m1—m3, 1 maxillary bone
with M1/—M2/ and 21 isolated molars (UMJG 204.194—216).
M e a s u r e m e n t s : See Table 2.
D e s c r i p t i o n : m1: The anteroconid is most often round,
with a long labial anterolophid extending to the protoconid as
a cingulum; one molar shows a broad anteroconid; the labial
anterolophid sensu stricto is then missing, but a strong cin-
gulid closes the protosinusid; the anterolophulid runs in more
or less longitudinal direction, but always connects to the an-
teroconid on its postero-labial wall; meta- and hypolophulid
are directed forwardly; the mesolophid more frequently is of
medium length, but may also be shorter or reach the lingual
border of the molar; the molars each possess two roots.
m2: Two crests connect the protoconid and the metaconid to
the anteroconid; the labial anterolophid reaches the base of the
protoconid; the mesolophid is short to medium-sized; the mo-
lars each possess two roots.
m3: The two m3 are heavily worn, so the morphology is dif-
ficult to assess; the molars each possess two roots.
M1: The anterocone is subdivided; the lingual anteroloph
extends to the base of the protocone; the anterolophule is dou-
ble, with each of the branches connecting the protocone to the
two cusps of the anterocone; the anterior protolophule is short
and does not reach the paracone; the short posterior protolo-
phule and the metalophule are directed backwardly; the me-
soloph is short; the sinus is closed by a cingulum; the
posterosinus is strongly reduced.
M2: The two anterolophs are either well developed or the
lingual one is reduced; protolophule and metalophule are
transverse in some molars but the protolophule may also be
directed forwards and the metalophule backwards; a double
protolophule is observed in one specimen; the broad sinus is
directed backwardly; the mesoloph is long and may reach the
labial border of the M2, or it is connected to the posterior spur
of the paracone; the molars each possess three roots.
M3: Only the proto- and paracone can be recognized as
cusps; the labial anteroloph extends to the base of the paracone,
while the lingual anteroloph is reduced; the protolophule is di-
rected forwardly; the posterosinus is present; the mesoloph
reaches the labial border of the M3 in one tooth, but is missing
on the other molar; the molars each possess three roots.
D i s c u s s i o n : The molars from Gratkorn are characterized
by their small size, the anterolophulid of the m1 connects to
the anteroconid on the labial side, and the double anterolo-
phule in most of the M1.
These traits are characteristic of M. minutus, a species insti-
tuted on the basis of a small molar sample from Inzersdorf and
Brunn-Vösendorf (Daxner 1967). Based on the rich sample
from Borský Svätý Jur (Slovakia), Joniak (2005) confirmed
the validity of the species. M. minutus has also been reported
from Austria in Mataschen, Bullendorf, Richardhof-Golfplatz
(Daxner-Höck 2004) and St. Margarethen/Zollhaus (G. Dax-
ner-Höck pers. comm.).
In Germany the small Megacricetodon from Hammer-
schmiede 1 (initially described as M. aff. debruijni (Mayr &
Fahlbusch, 1975) and Hillenloh corresponds well in size and
morphology to the teeth from Gratkorn; the same species is
present in Nebelbergweg, although Kälin & Engesser (2001)
ascribe those fossils to M. minor. Comparison of the teeth
from Gratkorn with these molar populations allows for the fol-
lowing preliminary observations:
– In the M1, the anterocone is broader in Gratkorn and the
metacone stands out from the outline of the labial border in
the Hammerschmiede molars.
– In the M2, the posterosinus is better developed due to a
more rounded posteroloph in Hammerschmiede. These molar
samples are tentatively interpreted as belonging to M. cf. minutus.
The molars from Gratkorn, although sharing some morpho-
logical features (i.e. the double anterolophule) with M. crisien-
sis from Comăne ti 1 (Radulescu & Samson 1988), cannot be
attributed to this species because of the somewhat larger mo-
lars and the presence of a lingual spur of the anterolophulid in
the m1 of M. crisiensis. In any case, the limited number of
specimens from Comăne ti 1 render comparisons with other
Megacricetodon molars difficult. Hír (2004, 2006) assigns the
small-sized Megacricetodon from Hungary and Romania ei-
ther to the species M. minor or M. minutus based on size (Ta-
ble 2). We hold the opinion that a detailed comparison of the
morphology of these forms is required in order to attribute the
samples to any species.
Small-sized Megacricetodon species also occur in the Mid-
dle Miocene of Anatolia, for example in Çandir (de Bruijn et
al. 2003). These teeth are identified as M. collongensis and
correspond well in size with the Gratkorn molars. Several
morphological characters can be used to distinguish these two
populations (e.g. length of the mesoloph in the M1).
The few specimens from Gratkorn are here assigned to M.
minutus.
Subfamily: Copemyinae Jacobs & Lindsay, 1984
Genus: Democricetodon Fahlbusch, 1964
D i a g n o s i s : Fahlbusch (1964).
T y p e s p e c i e s : Democricetodon crassus Freudenthal,
1969 ( = D. minor (Lartet, 1851) sensu Fahlbusch 1964).
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MIDDLE MIOCENE CRICETID RODENTS FROM GRATKORN (AUSTRIA)
Other species included in Democricetodon (we limit the
list to Freudenthal (2006)’s concept): Democricetodon affi-
nis (Schaub, 1925); D. brevis (Schaub, 1925); D. gaillardi
(Schaub, 1925); D. vindobonensis (Schaub & Zapfe, 1953);
D. romieviensis (Freudenthal, 1963); D. freisingensis Fahl-
busch, 1964 (considered by de Bruijn as junior synonym of
D. gaillardi in de Bruijn et al. 2003); D. gracilis Fahlbusch,
1964; D. mutilus Fahlbusch, 1964; D. hispanicus
Freudenthal, 1964; D. sulcatus Freudenthal, 1964; D. fran-
conicus Fahlbusch, 1966; D. cretensis de Bruijn & Meulen-
kamp, 1972; D. nemoralis Agustí, 1981; D. kohatensis
Wessels et al., 1982; D. hasznosensis Kordos, 1986; D. iazy-
gum Radulescu & Samson, 1988; D. zarandicus Radulescu
& Samson, 1988; D. walkeri Tong & Jaeger, 1993; D. lind-
sayi Qiu, 1996; D. tongi Qiu, 1996; D. hanniae Aguilar et
al., 1999; D. sudrei Aguilar et al., 1999; D. fourensis Mari-
det et al., 2000; D. doukasi Theocharopoulos, 2000; D. ana-
tolicus Theocharopoulos, 2000; D. moralesi van der Meulen
et al., 2004.
Democricetodon sp. nov. (sensu Kälin & Engesser, 2001)
(Fig. 2.G—I)
M a t e r i a l a n d m e a s u r e m e n t s : 1 fragmentary lower
jaw with m3, 1 fragmentary upper jaw with M2, 10 isolated
molars (UMJG 204.181 to 193). M1: 1.85 1.26, 1.84 1.28,
1.88 1.28; M2: 1.40 1.30, 1.44 1.30; M3: 1.06 1.13; m1:
1.65 1.13, 1.68 1.21, 1.59 1.11; m3: 1.28 1.03.
D e s c r i p t i o n : m1: The anteroconid is round in young in-
dividuals but becomes ovaloid in worn molars by integration
of the long convex labial anterolophid; the lingual antero-
lophid is absent or very short; the anterolophulid is very short;
the metalophulid connects to the anterolophulid; the meso-
lophid is long and reaches the border of the molars or the cin-
gulid closing the mesosinusid; the hypolophulid is proverse;
the molars each possess two roots.
m2: The single tooth is broken on the labial side; the low
mesolophid reaches the border of the molar.
m3: The entoconid is indistinct; the labial anterolophid ex-
tends to the labial wall of the protoconid; mesolophid and hy-
polophulid are of about the same length.
M1: The anterocone has two adjoining cusps with a superfi-
cial groove on its anterior wall; the two anterolophs are
present, forming a bow-shaped structure with the anterocone;
the anterolophule is broad and transverse; the labial spur of the
anterolophule is most often absent, but may also be present
and low and long; the protolophule is directed forewardly or
double, the mesoloph is of medium length or somewhat long-
er; the sinus and the mesosinus are closed; the metalophule is
directed backwardly; the posterosinus is reduced to a narrow
valley; the molars each possess three roots.
M2: The two M2 are either damaged or partially covered by
hard sediment; the two anterolophs are present; the protolo-
phule is double; the metalophule is directed backwardly in one
molar, directed forwardly in the second; the molars each pos-
sess three roots.
M3: The hypocone is highly reduced, the metacone absent;
the two anterolophs are well developed; double protolophule,
the posterior protolophule extends into the axioloph; the me-
soloph is of medium length; sinus and mesosinus are closed
by a cingulum; three roots.
D i s c u s s i o n : The molars from Gratkorn are clearly related
to the Democricetodon species found in Nebelbergweg. Al-
though that material shows a number of resemblances with
Democricetodon brevis, Kälin & Engesser (2001) assigned the
fossils to a new species, but refrained from naming it. The
same species occurs in the German locality Hammerschmiede
(Mayr & Fahlbusch 1975). The mesoloph of the three M1
from Gratkorn are somewhat smaller than those from the two
other localities.
Democricetodon from Felsötárkány-Felnémet (Hír 2006:
plate 2, figs. 5 and 10) is similar in size and morphology to the
Gratkorn specimens.
Casanovas-Vilar (2007) reports Democricetodon sp. (= D.
cf. sulcatus in Casanovas-Vilar et al. 2006) from Creu Conill
(Spain, Valles-Penedes Basin). The single M1 has a slightly
subdivided anterocone, but the labial spur of the anterolophule
is missing and the corresponding m2 has no mesolophid. D.
brevis brevis from the Barranc de Can Vila 1 is characterized
by long mesoloph(id)s and M1 with a slightly divided antero-
cone and long labial spur of the anterolophule (Casanovas-
Vilar 2007). This subspecies differs from D. brevis nemoralis
by the presence of a double protophule in the M1 and the un-
divided anteroconid of the m1. The fact that the teeth from
Gratkorn and Barranc de Can Vila 1 may belong to the same
(sub)species cannot be ruled out.
D. zarandicus from Tau (early Late Sarmatian, Romania) is
characterized by M1 with fissured anterocone and complete la-
bial spur of the anterolophule (Radulescu & Samson 1988). The
mesolophid of the m1 is long and reaches the labial border. The
specimens from Gratkorn differ from this species in their shorter
transversale crests of the M1 and the somewhat larger size. A
phylogenetic relationship between these species is possible.
The limited sample of molars of D. iazygum from
Comăne ti 1 (earliest Late Sarmatian, Romania) also contains
M1 with fissured anterocone but with mesolophs and labial
spur of the anterolophule shorter than in D. zarandicus. With
regard to these characters, the form from Gratkorn does not
differ substantially. However, the sample from Comăne ti is
too small to depict the intraspecific variability of D. iazygum.
For this reason, the Gratkorn material should not be attributed
to this species.
The Democricetodon molars from Çandir and Gratkorn are
of almost equal size and similar in basic morphology. De
Bruijn et al. (2006) attributed the Turkish specimens to D. aff.
gaillardi (where D. freisingensis is a junior synonym of D.
gaillardi). Moreover, they noted the resemblance of this form
to the poorly documented D. kohatensis from Pakistan.
Democricetodon lindsayi from Moergen (Nei Mongol, Chi-
na; Qiu 1996) has a well-fissured anterocone in the M1. How-
ever, it differs from the species from Gratkorn in the presence
of a lingual anterolophid and shorter mesolophid in the m1.
We conclude that the teeth from Gratkorn are related to a
suite of European and Asiatic species characterized by a fis-
sured anterocone in the M1. The earliest representatives come
from Çandir (Anatolia, middle part of Middle Miocene),
which allows for the hypothesis to be advanced that the evolu-
tionary origin of this type of Democricetodon was located
`
`
430
PRIETO, BÖHME and GROSS
somewhere in the East. The molars from Gratkorn are provi-
sionally assigned to the species from Nebelbergweg.
Subfamily: Eumyarioninae Ünay, 1989
Genus: Eumyarion Thaler, 1966
D i a g n o s i s ( e m e n d e d ) : Mein & Freudenthal (1971).
D i f f e r e n t i a l d i a g n o s i s : Ünay (1989).
T y p e s p e c i e s : Cricetodon medium Lartet, 1851 ( = Cric-
etodon helveticum Schaub, 1925).
Other species included in Eumyarion: E. latior (Schaub &
Zapfe, 1953); E. weinfurteri (Schaub & Zapfe, 1953); E. bifidus
(Fahlbusch, 1964); E. leemanni (Hartenberger, 1965); E. valen-
cianum Daams & Freudenthal, 1974 (nomen dubium in de
Bruijn & Saraç 1991: 14 and de Bruijn 2009: 78); Eumyarion
montanus de Bruijn & Saraç, 1991; E. intercentralis de Bruijn
& Saraç, 1991; E. microps de Bruijn & Saraç, 1991; E. carboni-
cus de Bruijn & Saraç, 1991; E. orhani de Bruijn et al., 2006.
Eumyarion sp.
(Fig. 2.M—P, Fig. 4)
M a t e r i a l a n d m e a s u r e m e n t s : Fragmentary skull
with two complete molar rows (UMJG 204.017), 2 fragmen-
tary upper jaws (one with M1, the other with M3), 6 isolated
teeth (UMJG 2004.217-224). M1: 2.16 1.50, 2.15 1.63,
2.06 1.41; M2: 1.56 1.54, 1.54 1.48; M3: 1.15 1.33,
1.26 1.29; m1: 2.11 1.38; m2: 1.75 1.39.
D e s c r i p t i o n : Skull: The fragmentary skull (Fig. 4) is
prepared so that it can be viewed from the ventral side; the
maximal length of the specimen is 15 mm; it includes the two
molar rows; the left row shows slight corrosion marks, where-
as the left row is clearly more corroded, especially on the labi-
al side; the palatine is anteriorely broken; the posterior
foramen palatinum is located lingual to the M2, and is about
1.2 mm long; with the exception of the basisphenoid, which is
isolated, the posterior part of the skull is missing.
m1: The single m1 is hardly worn; the ovaloid anteroconid
is positioned in the centre of the width of the molar and isolat-
ed from the metaconid, although a small longitudinal crest is
present on the postero-lingual part of the anteroconid; the
strong labial anterolophid begins at about half of the high of
the anteroconid and ends at the base of the protoconid; the an-
terolophulid is oblique, reaching the posterior wall of the an-
teroconid centrally (the anterolophulid appears to be
interrupted in the specimen illustrated in figure 4P, but this is
not the case); a short labial anterolophulid spur extends to the
metalophulid; this crest is double close to the metaconid,
whereas a single crest reaches the anterolophulid; protoconid
hind arm, mesolophid and ectolophid together form a triangu-
lar structure that lingually merges with the posterior wall of
the metaconid; the strong metaconid ridge closes the mesosi-
nusid; a small ectomesolophid is present along the labial wall
of the ectolophid; the broad posterosinusid contains a well-de-
veloped posterior hypolophulid; two roots.
m2: The metaconid and entoconid are high, worn on their
labial wall, whereas the other cuspids and crests are worn on
the occlusal plan of the molar, forming a somewhat horizontal
surface; the anteroconid is triangular with the labial antero-
lophid extending to the base of the protoconid; the small lin-
gual anterolophid connects to the base of the metaconid; the
metalophulid is directed forwardly and extends to the antero-
conid; the protoconid hind arm connects to the medium-sized
mesolophid; the two fused crests reach the metaconid; small
ectomesolophid present; hypolophulid transverse; the postero-
lophid connects to the posterior wall of the entoconid and
well-developed posterior hypolophulid; two roots.
m3: The m3 is heavily worn and broken; two roots.
M1: The anterocone is subdivided; anterolophs are missing
but the anterior valleys are closed by massive cinguli; a dou-
ble anterolophule connects to both cusps of the anterocone;
the protolophule is transverse; the long mesoloph does not
reach the mesostyl closing the mesosinus; metalophule and
posteroloph are parallel; three roots.
M2: The para- and metacone are high, worn on their labial
walls, whereas the other cusps and crests are worn in the oc-
clusal plan of the molar, forming a somewhat horizontal sur-
face; the anterocone is indistinct; the lingual anteroloph is
very short; all transversal crests are parallel; the long meso-
loph does not reach the mesostyl closing the mesosinus; en-
tomesoloph absent or very short; a very short “anterior”
mesoloph can be distinguished on the entoloph of one molar;
three roots.
M3: Anterocone and protocone abut; the lingual anteroloph
missing; anterolophule and protolophule joining the axioloph;
a posterior crest occurs on the protolophule; in one specimen
it is very short, while it is developed as a “second axioloph” in
the second molar; the neo-entoloph is absent as the two lingual
cusps are abut; the sinus is delimited by a cingulum or absent;
metalophule, posteroloph and mesoloph are of equal length.
D i s c u s s i o n : Due to the high intraspecific variability of the
Eumyarion species (e.g. de Bruijn 2009), and the fact that more
than one species may be present at one locality (e.g. Wu 1982;
de Bruijn et al. 2006), the attribution of small molar samples is
mostly difficult. The molars are morphologically and metrically
similar to E. latior, a species defined in the Middle Miocene of
Neudorf (Slovakia; Schaub & Zapfe 1953; Fejfar 1974). The
stratigraphically younger E. leemanni, reported from latest Mid-
dle Miocene and Late Miocene deposits of the Valles-Penedes
Basin, is characterized by cusps integrated into the ridges that
join them. A different taxonomic approach has been proposed
by Kretzoi & Fejfar (2005: p. 127), who recognized a latior lin-
eage in Central Europe. Based on taxonomic ambiguities (early
E. leemani vs. E. latior) and the limited number of specimens at
hand, the molars from Gratkorn cannot be identified with confi-
dence at species level.
Biostratigraphic discussion
Stratigraphic range of the cricetid species from Gratkorn
Four cricetid rodents have been recorded in the fauna from
Gratkorn:
– Megacricetodon minutus was discovered in St. Margare-
then/Zollhaus, within the Late Sarmatian s. str. part of the
`
`
431
MIDDLE MIOCENE CRICETID RODENTS FROM GRATKORN (AUSTRIA)
succession (G. Daxner-Höck pers. comm.), stratigraphically
close to Gratkorn. This species is the only cricetid found in
Mataschen (Styrian Basin, Daxner-Höck 2004), and it has also
been reported from Bullendorf (Vienna Basin, Daxner-Höck
1996). Both localities correspond to the Pannonian “Zone B”,
respectively “B/C” (sensu Papp 1951). The species also oc-
curs in localities dated as Pannonian “Zone E” (e.g. Vösen-
dorf, Inzersdorf and Richardhof-Golfplatz; Daxner-Höck 1996
and pers. comm.). Moreover, M. minutus has been reported
from the lacustrine deposits of Borský Svätý Jur (Slovakia,
Pannonian “Zone E”; Joniak 2005) and the Hungarian locality
Felsötárkány 3/2 (Hír 2004). The genus Megacricetodon dis-
Fig. 4. Eumyarion sp. Fragmentary skull in ventral view (UMJG 204.017). 1 – Molar rows; 2 – Os palatinum; 3 – Foramen palatinum
posterior; 4 – Basisphenoidale.
appears from the record in Austria some time during the Late
Pannonian (“Zone F”; Daxner-Höck 1996).
– Democricetodon sp. nov. (sensu Kälin & Engesser 2001)
is documented in Gratkorn by a small sample of teeth, but the
intraspecific variability remains incompletely documented.
Nevertheless, its close relationships to the medium-sized
Democricetodon populations characterized by a fissured an-
terocone in the M1 are obvious. These forms have been re-
corded from Spain in the West to China in the East, and are
abundant in the late Middle Miocene and earliest Late Mio-
cene. The genus does not persist to the end of the MN10 in
Central Europe (Daxner-Höck 1996).
432
PRIETO, BÖHME and GROSS
– Eumyarion has been reported in Austria from the Early
Miocene of Oberdorf (E. cf. weinfurteri, Daxner-Höck
1998a). The species E. weinfurteri is also known from
Obergänserndorf and Teiritzberg (Early Miocene, Late Karpa-
tian; Daxner-Höck 1998b) and the Middle Miocene (Early
Badenian) of Mühlbach and Grund (Daxner-Höck 2003).
However, these samples are taxonomically not directly relat-
ed to Eumyarion sp. from Gratkorn. Eumyarion sp. from
Götzendorf (Pannonian “Zone F”; Daxner-Höck 1996) is
stratigraphically younger. As a result, the molars from Grat-
korn increase our knowledge of the fossil history of this ge-
nus in Austria. The youngest Eumyarion representatives
have been recorded from the base of the MN10 (Sucho-
masty; Daxner-Höck 1996).
– The fourth cricetid, “Cricetodon” fandli, represents a
new species. Cricetodon meini from Mühlbach and Grund
(Daxner-Höck 2003) is not directly related to the form from
Gratkorn. The small Cricetodontini sample from Bullendorf
(Daxner-Höck 1996) has not yet been precisely determined.
Biostratigraphical implications and correlation of the small
mammal fauna
A comparison of the Gratkorn fauna with faunas from local-
ities of the North Alpine Foreland Basin (NAFB) suggests that
the small mammals from Gratkorn generally are more closely
related to faunas from NAFB-localities, which are traditionally
regarded as corresponding to the base of the Late Miocene
(MN9, occurrence of Microtocricetus molassicus), than to
faunas from Middle Miocene sites (e.g. Anwil, Kleineisen-
bach, Giggenhausen; Prieto 2007).
For example, until recently the gymnure Schizogalerix voe-
sendorfensis was believed to have appeared not before the
Late Miocene (Kälin & Engesser 2001; Ziegler 2006). Its dis-
covery in Gratkorn now shows that the species was already es-
tablished in the Late Sarmatian (Prieto et al. in print a).
Following the biostratigraphic concept proposed in Prieto &
Rummel (2009), the NAFB-localities traditionally correlated
with MN9 (Hammerschmiede, Hillenloh, Nebelbergweg) are
older than, or at least of about the same age as, Felsötárkány
3/2, a locality correlated with the Sarmatian (Hír 2004). This
implies that most of the MN9 cricetid rodents of the NAFB are
Middle Miocene in age, or close to the Middle/Late Miocene
boundary. This has fundamental biostratigraphic consequenc-
es, as Hammerschmiede is the type locality of Microtocricetus
molassicus. The first occurrence of the genus Microtocricetus
defines the beginning of MN9 (de Bruijn et al. 1992; Fejfar
1999; Mein 1999). Microtocricetus was unknown from Aus-
tria before the Late Pannonian “Zone F” of Götzendorf (Dax-
ner-Höck 1996), and its absence in Gratkorn may argue for a
first occurrence of the genus during the Late Miocene. How-
ever, taphonomic and ecological bias may alter the assem-
blage, because Microtocricetus is usually regarded as an
inhabitant of moist, riparian environments (Fejfar 1999) that
are different from the paleoenvironment represented by the
Gratkorn fossiliferous beds (Harzhauser et al. 2008; Gross et
al. 2009a,b and subm.). Humidity data from Microcricetus-
bearing localities (e.g. Hammerschmiede 1—3, Rudabánya,
Götzendorf) provide mean annual precipitation estimates
(Böhme et al. 2006, 2008) ranging from 974 mm (Hammer-
schmiede 1) to 1303 mm (Götzendorf), which is significantly
more humid than the estimate for Gratkorn (486 mm; Gross et
al. subm.).
In Hungary, Microtocricetus is first recorded at the top of
the Felsötárkány sequence (Felsötárkány 3/8 and 3/10). The
co-occurring mollusc fauna is suggestive of a Sarmatian age
(Hír 2006 and Hír & Kókay 2010). Microtocricetus has also
been reported from the Ukrainian locality Grytsiv (Topachev-
sky & Skorik 1988; Kowalski 1993). The fissure fillings con-
taining the fossil vertebrates share Sarmatimactra vitaliana
(Topachevsky & Scorik 1992; Topachevsky et al. 1996; Rze-
bik-Kowalska & Topachevsky 1997; Vangengeim et al.
2006), a characteristic bivalve of the Late Sarmatian s. str. in
the Central Paratethys (Papp 1954; Piller et al. 2007).
The value of Microtocricetus as an index fossil for the onset
of the Late Miocene remains questionable and its first occur-
rence traces back into the Sarmatian s. str.
We interprete the new Gratkorn species “C.” fandli as an
ancestor of C. klariankae from Felsötárkány-Felnémet (Sar-
matian; Hír 2006), a form that may represent the youngest
Cricetodon species from Central Europe. The proposed lin-
eage fandli—klariankae cannot be directly related to other
cricetodontine rodents. Nevertheless, parallel evolutionary
patterns have been noted between the faunas in Europe and
Anatolia. This allows the comparison of geographically dis-
tant fossil localities. For instance, Casanovas-Vilar et al.
(2008) date Barranc de Can Vila 1 at between 12.0 and
12.5 Ma, more probably 12 Ma (Moya-Sola
et al. 2009). The
faunal composition at this locality (Casanovas-Vilar 2007) is,
with the exception of Democricetodon larteti ( = Fahlbuschia
larteti), very similar to that from Gratkorn (12.0—12.2 Ma):
both localities have yielded a large cricetid rodent with moder-
ate hypsodonty and of comparable size (Hispanomys sp./
“Cricetodon” fandli), a medium-sized Democricetodon with
fissured anterocone in the M1 (D. brevis brevis/D. sp. nov.), a
small Megacricetodon (M. minor minor/M. minutus), and a
form attributable to Eumyarion (E. leemani/E. sp.).
These parallel evolutionary patterns may pose an argument
for the existence of different paleobioprovinces around the
Middle to Late Miocene transition. The bioprovinciality is
documented by the heterochrone last occurrences of taxa at
the subfamily level, including the Cricetodontinae. While the
Cricetodontini disappeared from the fossil record of Central
Europe close to the Middle/Late Miocene boundary, Hispano-
mys/Ruscinomys persisted into the Pliocene in Western Eu-
rope, and Byzantinia survived until the end of the Miocene in
Anatolia and Southeastern Europe. The insectivore fauna from
Gratkorn in fact seems to be related to faunas from Eastern
Europe/Anatolia (Prieto et al. in press a). The mole Desmano-
don fluegeli shares several features with D. minor/D. major
from Turkey. Similarly, Schizogalerix voesendorfensis has
been assigned to an Austrian lineage that shares similarities
with Schizogalerix from Turkey (Engesser 1980). The cricetid
rodents from Gratkorn are generally more closely related to
assemblages from the Pannonian Basin (Hungary in particu-
lar), and to a lesser degree also Anatolian assemblages, than to
late Middle Miocene faunas from the NAFB (e.g. Anwil,
Kleineisenbach, Giggenhausen). On the other hand, the lack
`
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433
MIDDLE MIOCENE CRICETID RODENTS FROM GRATKORN (AUSTRIA)
of sufficient material and studies of Middle to Late Miocene
fossil localities in Austria and especially Germany currently
precludes a more decisive assessment of the suggested pres-
ence of two independent bioprovinces at least at the time of
the accumulation of the Gratkorn fauna.
Conclusions
In spite of the low species diversity (four species), the crice-
tid rodent assemblage from Gratkorn increases our knowledge
about the Austrian rodent succession. Well-dated as Late Sar-
matian s. str., the lineage “Cricetodon” fandli—Cricetodon
klariankae may become a key to the correlation of fossil local-
ities from Central Europe. In a broader context, the fauna
Gratkorn is of major importance for the understanding of the
continental biostratigraphy of the late Middle Miocene of
Austria and in a general view of the Paratethyan realm. More-
over, the geographical position of the locality, relatively cen-
tral in Europe, makes it important for the understanding the
faunal interchanges and bioprovincialism at the end of the
Middle Miocene.
Acknowledgments: The authors express gratitude to Isaac
Casanovas-Vilar and Gudrun Daxner-Höck for fruitful discus-
sion and information on taxonomy. We are grateful to Hans de
Bruijn (Utrecht), Peter Joniak (Bratislava), Mathias Harzhauser
(Vienna) and János Hír (Paszto) for their constructive com-
ments on the text. Special thanks go to the community of
Gratkorn, especially to mayor Elmar Fandl (†), for providing
financial support during the 2008 field season. For generously
granting access to the clay pit, we are indebted to the Wieters-
dorfer & Peggauer Zementwerke AG (especially, Josef
Plank). Norbert Winkler (Joanneum) has meticulously pre-
pared the specimens discovered during excavations conducted
by the staff of the Landesmuseum Joanneum (financed in part
by the Land Steiermark/Wissenschaftsreferat). Christopher
Murgatroyd and Michael Krings (both Munich) are thanked
for the linguistic improvements.
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