N. Jb. Geol. Paläont. Abh. 258/1, 107– 119
published online August 2010
Article
Insectivores and bat (Mammalia) from the late Middle Miocene
of Gratkorn (Austria): biostratigraphic and ecologic implications
Jérôme Prieto, Munich, Martin Gross, Graz, Christine Böhmer, Munich and
Madelaine Böhme, Tübingen
With 5 figures and 1 table
PRIETO, J., GROß, M., BÖHMER, C. & BÖHME, M. (2010): Insectivores and bat (Mammalia) from the
late Middle Miocene of Gratkorn (Austria): biostratigraphic and ecologic implications. – N. Jb. Geol.
Paläont. Abh., 258: 107–119; Stuttgart.
Abstract: The discovery of a late Sarmatian (latest Middle Miocene) small mammal fauna at the base
of the clay pit St. Stefan near Gratkorn is of primary importance for the biostratigraphic and ecologic
comprehension of the faunal evolution during the late Middle Miocene of Central Europe. In this
paper five species of insectivores and one bat are described. The gymnure Schizogalerix voesendorfensis is the most abundant species of the small mammal fauna. The Gratkorn sample represents
the earliest occurrence of the species in the fossil record. A single molar of a second, large-sized
Galericinae cannot be attributed to any known species. The new talpid Desmanodon fluegeli n. sp.
shares similarities with the Anatolian lineage D. minor / D. major. An exceptionally well preserved
mandible of the large shrew Dinosorex cannot be determinate at the species level. In addition, some
rare remains of the bat cf. Myotis sp. are also reported. From biogeographical point of view the
insectivore fauna can be related to eastern Central European / Anatolian assemblages. Although
taphonomic biases are suspected, the environmental conditions proposed on the basis of sedimentary
analyzes (open forest with moist soil) can contribute to the low alpha diversity of insectivores in this
locality.
Key words: Sarmatian, faunal relationships, Mammal, palaeoecology, insectivores, bats.
1. Introduction
The fossil insectivore and chiropteran assemblages
from Austria were recently intensively studied and
revised (ZIEGLER & DAXNER-HÖCK 2005; ZIEGLER
2006a, b). As a result, the evolution of these mammalian groups is detailed for the Lower and early Middle
Miocene and especially for the Late Miocene. The
period in between, however, yielded few data. In
this context the find of a late Middle Miocene (late
Sarmatian, HARZHAUSER et al. 2008) small mammal
fauna at the base of the clay pit St. Stefan by Gratkorn
(GROSS et al. 2007; GROSS 2008, PRIETO submitted) is
of primary importance for the biostratigraphic and
ecologic comprehension of the faunal evolution
during this period, especially as it is critical in terms
© 2010 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany
DOI: 10.1127/0077-7749/2010/0088
of climatic changes (BÖHME 2003; BÖHME et al.
2008). The insectivores are particularly interesting,
as ZIEGLER (2006b) found surprising differences
between German and Austrian assemblages, in spite of
the geographic vicinity. Furthermore, Sarmatian small
mammal samples are still extremely rare in Central
and Eastern Europe, making the finds from Gratkorn
an important piece in the puzzle of the faunal interchanges at the end of the Middle Miocene.
2. Material and methods
The measurement method of the erinaceids is presented in
Fig. 1, and the nomenclature follows ENGESSER (1980).
The measurements and nomenclature of the talpids follow
ZIEGLER (2003 a) based on HUTCHISON (1968, 1974); the
www.schweizerbart.de
0077-7749/2010/0088 $ 3.25
108
J. Prieto et al.
Fig. 1. Measurement method of the erinaceids (after PRIETO & RUMMEL 2009a). Dotted lines: baselines. In case of
T-shaped teeth (f. e. part of the P3, D3) the width will be measured perpendicular to the length. In case of reduced talonid
(f. e. lower premolars) then just W1 is measured. L: length; L1: labial length; L2: shortest antero-posterior length;
W: width; W1: anterior width; W2: posterior width.
soricids follow REUMER (1984). All measurements are given
in mm and the specimens in the figures are presented in left
orientation. Hence, right specimens are reversed. The fossils
from Gratkorn are stored in the Landesmuseum Joanneum
of Graz (Geologie & Paläontologie, abbreviated LMJG) and
in the Bayerische Staatssammlung für Paläontologie und
Geologie in Munich (abbreviated BSPG).
1980
Schizogalerix voesendorfensis (RABEDER). – ENGESSER, figs. 3d, 4b.
Part. 1993 Galerix aff. socialis (RABEDER). – KÄLIN, p. 5456, fig. 23a.
2001 Schizogalerix voesendorfensis (RABEDER 1973). –
KÄLIN & ENGESSER, p. 13-15, fig. 8.
Type locality: Vösendorf (Austria).
3. Systematic paleontology
Order Erinaceomorpha GREGORY, 1910
Family Erinaceidae FISCHER, 1814
Subfamily Galericinae POMEL, 1848
Genus Schizogalerix ENGESSER, 1980
D ia g n o s is (emended): VAN
686).
DEN
HOEK OSTENDE (2001:
D iff e r e n tia l di a gnosi s: ENGESSER (1980: 63) and
comments in VAN DEN HOEK OSTENDE (2001).
Ty p e s p e c ie s : Schizogalerix anatolica ENGESSER, 1980.
O t h e r s p e c i e s i n c l u d e d : S. zapfei (BACHMAYR &
WILSON, 1970) (= Galerix atticus RÜMKE, 1976), S. moedlingensis (RAEDER, 1973), S. voesendorfensis (RABEDER,
1973), S. pasalarensis ENGESSER, 1980, S. anatolica
ENGESSER, 1980, S. sinapensis SEN, 1990, S. macedonica
DOUKAS, 1995, S. sarmatica (LUNGU, 1981), S. duolebulejinensis BI et al. 1999, S. intermedia SELÄNNE, 2003, S.
evae DE BRUIJN et al. 2006.
‘Schizogalerix’ iliensis KORDIKOVA, 2000 is transferred to
the genus Galerix by DOUKAS & VAN DEN HOEK OSTENDE
(2006). DE BRUIJN et al. (2006) and DOUKAS & VAN DEN
HOEK OSTENDE (2006) consider S. pristina ZIEGLER, 2003
as an ancestor of Parasorex and not as Schizogalerix.
Schizogalerix voesendorfensis (RABEDER, 1973)
Fig. 2A-J
1973
Galerix voesendorfensis n. sp. – RABEDER, p. 430433, figs. 1-10.
S tratig raphic cor relation: early Late Miocene
(MN9), Middle Pannonian “zone” D/E (“zonation” after
PAPP 1951).
M aterial und meas urements : 67 isolated teeth, 22
mandibles, 7 maxillaries and 1 fragmentary skull (BSPG
2008 IV 1-27; LMJG 203.740, 984-999, 204.022, 024,
026-30, 043, 048, 104, 105, 138-143, 146, 208.732-737,
739-742). (Table 1).
D iagnos is : Since RABEDER (1973) did not give any diagnosis for his new species (only a differential diagnosis) and
considered it part of the genus Galerix, a diagnosis is
proposed herein: Medium sized Schizogalerix species
having a continuous to slightly interrupted and clearly
S-shaped mesostyle in the first two upper molars. The
species does not fully develop the posterior cingulid/
entoconid connection in the two first lower molars. The
hypolophid is continuous.
D es cription and dis cus s ion. – Dental remains: the p2
is smaller than the p3; the well-developed paralophid of
the p 4 is continuous; the hypolophid of the m1 and m2 is
continuous; the connection posterior cingulid/entoconid is
variable, most often not fully achieved: in some specimens
this connection is absent, as found in the German and Swiss
Parasorex socialis, whereas in some teeth this cingulid
reaches the top of the entoconid. The mesostyle of the first
two upper molars is clearly S-shaped (‘Schlaufenbildung’
in German, ENGESSER 1980) and most often continuous
with tapering of its middle part; in some specimens, this
structure is interrupted, but without the development of a
clear valley; there is no connection between metaconule and
protocone; the posterior arm of the metaconule ends to the
postero-labial corner of the M1 and M2 and the P3 has two
Insectivores and bat from the late Middle Miocene of Gratkorn (Austria)
109
Fig. 2. The erinaceids, soricid and bat from Gratkorn. All teeth are shown in left orientation and in occlusal view (except
M: labial view). A-J: Schizogalerix voesendorfensis (RABEDER, 1973): A: right P3 (reversed, LMJG 208.732); B: right
P4 (reversed, LMJG 208.733); C: left M1 (LMJG 208.734); D: left M2 (LMJG 208.735); E: left M3 (LMJG 208.736);
F: right d4 (reversed, LMJG 204.159); G: left p4 (LMJG 208.742); H: left m1 (LMJG 208.740); I: left mandible with
m2-m3 (LMJG 203.986); J: left m3 (LMJG 208.741); K: Galericinae gen. et sp. indet.: right M1 (reversed, LMJG
208.738); L-M: Dinosorex sp.: L: left P4 (LMJG 204.161); M: left upper incisor (LMJG 204.160); N-O: cf. Myotys sp.: N:
left m1 (LMJG 204.162); O: left M2 (LMJG 204.163).
lingual cusps. All these morphologic characters permit to
attribute the teeth to the genus Schizogalerix.
The dental remains from Gratkorn fit best, in size
and morphology, with the species S. voesendorfensis and
S. anatolica. Following the differential diagnosis given by
ENGESSER (1980: 70), the specimens from Gratkorn can be
attributed to the Austrian species. This is especially clear in
the trigonid basin of the lower molar, which is broader than
in S. anatolica, and the absence of diagonal enlargement in
the P4. The fossil record of S. voesendorfensis is, at present,
Table 1. Sample statistics of the Schizogalerix voesendorfensis (RABEDER, 1973) molars. n: number of specimens; min:
minimum; max.: maximum; mean: arithmetic mean. For method and additional legend see Fig. 1.
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J. Prieto et al.
Fig. 3. Mandible fragments of
Schizogalerix voesendorfensis
(RABEDER, 1973) and
Parasorex socialis (VON MEYER, 1865):
A: Parasorex socialis, labial view
(Petersbuch 6, BSPG 1993 IV 653);
B-C: Schizogalerix voesendorfensis
(Gratkorn): B. labial view (reversed,
LMJG 204.239); C1: occlusal view;
C2: labial view (both reversed,
LMJG 204.104).
restricted to Austria and Switzerland. In addition to the
Austrian type locality Vösendorf (RABEDER 1973), the species is also reported in Nebelbergweg (KÄLIN & ENGESSER
2001). Metrically these assemblages are very similar,
though the teeth from Gratkorn are slightly smaller.
Mandible: the mental foramen is found under the p3; in
general, the mandibles from Gratkorn resemble Parasorex
socialis mandibles (figures in ENGESSER 1980, NOVACEK
et al. 1985, and ZIEGLER 2005a). Nevertheless, some significant differences can be observed (Fig. 3): the p1 and the
preceding anterior teeth are clearly deeply slanted forwards
in P. socialis; furthermore the anterior part of the mandible
tapers from the p3 onwards in P. socialis, but more anteriorly (p2) in Gratkorn. Based on the large size of the premolars
from Vösendorf, KÄLIN & ENGESSER (2001: 15) hypothesize a low reduction level of the mandible of the Swiss
species, which is a primitive character state within the
genus. This is confirmed by the fossils from Gratkorn.
galerix voesendorfensis molar sample due to its larger
size and the presence of a straight and undivided mesostyl.
Furthermore, the posterior arm of the metaconule does not
join the postero-labial margin of the tooth and the hypocone
is very conical, due to the absence of an anterior arm. A
small crest is developed between the two lingual cusps in
the direction of the metaconule.
The overall morphology of the M1 fits well with Parasorex socialis even though the molar is too large to be
attributed to this species. In addition to S. voesendorfensis,
KÄLIN & ENGESSER (2001) report a second and larger
Schizogalerix species in Nebelbergweg. Unfortunately this
species is a rare element of the Swiss fauna (four teeth) and
the M1 is not represented in the sample. The M2 displays a
divided mesostyle, and such a character should be present
in the M1 or at least the mesostyle has to be S-shaped. This
is not the case in Gratkorn. Therefore, due to the lack of
diagnostic characteristics the molar cannot be determined.
Galericinae gen. et sp. indet.
Fig. 2K
Order Soricomorpha GREGORY, 1910
Family Talpidae F ISCHER, 1814
Subfamily Talpidae incertae sedis
Genus Desmanodon ENGESSER, 1980
M a te r ia l: Partly damaged right M1 (LMJG 208.738).
D e s c r ip tio n a nd di sc ussi on: The single M1 is corroded but can be clearly differentiated from the Schizo-
D iagnos is : See ENGESSER (1980: 116), differential diagnosis in ENGESSER (1980: 116-117).
Insectivores and bat from the late Middle Miocene of Gratkorn (Austria)
111
Fig. 4. Desmanodon fluegeli n. sp. All specimens are shown in left orientation; A2-E: occlusal view. A: Holotype, left lower
mandible with p3-m3 (LMJG 204.164, A1: labial view, A2: occlusal view); B: right m2 (reversed, LMJG 204.165);
C: right m3 (reversed LMJG 204.166); D: left M2 (LMJG 204.167); E: right M3 (reversed, LMJG 204.168); F: left
humerus (F1: cranial view, F2: caudal view, LMJG 204.056).
Ty p e s p e c ie s : Desmanodon major ENGESSER, 1980.
O th e r s p e c ie s i nc l ude d: D. minor ENGESSER, 1980, D.
antiquus ZIEGLER, 1985 (= D. meuleni DOUKAS, 1986 in
DOUKAS & VAN DEN HOEK OSTENDE 2006), D. daamsi VAN
DEN HOEK OSTENDE, 1997, D. ziegleri, VAN DEN HOEK
OSTENDE 1997, D burkarti VAN DEN HOEK OSTENDE, 1997,
D. crocheti PRIETO, in press.
Paratypes : 2 fragmentary lower molars (trigonid), 2 m1,
3 m2, 4 m3, 2 M1, 2 M2, 2 M3, left humerus (LMJG
204.056, 165-180). Measurements of teeth: m2: 2.13 x
(1.33 x1.35); m3: 1.80x(0.92 x0,72); 1.60 x(0.97x0.80);
1.60x(0.95 x0.80); M2: 1.90x2.15; 2.02x2.13; M3:
1.55x(1.10 x1.72); 1.70x(1.10 x1.73). Measurements of
humerus: Diameter of the shaft: 3.00; distal width: 5,65;
distal with without epicondyles: 4.65.
Type locality: Gratkorn (Austria).
Desmanodon fluegeli n. sp.
Fig. 4
In press Desmanodon sp. from Gratkorn. – PRIETO.
E t y m o l og y : In honour of HELMUT W. FLÜGEL, for his
intensive geologic work in the Graz area.
Ho l o t y p e: Right lower jaw fragment with c, p3-m3
(LMJG 204.164, Fig. 3A); measurements: c: 0.90 x0.60;
p3: 0.83x 0.55; p4: 1.23x0.63; m1: 1.93 x(1.10 x1.28); m2:
1.97 x (1.28 x 1.32); m3: 1.63 x1.13.
Type hor i zon: Clay pit of the company Wietersdorfer &
Peggauer Zementwerke AG, 10 Km NW from Graz
(15° 20’55’’E / 47°08’15’’N). Mottled and intensively bioturbated paleosoil at the base of the profile (GROSS et al.
2007: 180); late Middle Miocene, late Sarmatian (upper
Ervilla Zone; Porononion granosum Zone).
D iagnos is : Medium-sized Desmanodon species morphologically close to D. minor; single-rooted p3; lower molars
with a short entoconid and continuous labial cingulid.
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J. Prieto et al.
D iff e r e n tia l di a gnosi s: Desmanodon fluegeli n. sp.
differs from:
- Desmanodon antiquus (= D. meuleni) in the more divided
mesostyl of the upper molars and in the broader pectoral
crest of the humerus (compared with the specimen from
Rauscheröd in ZIEGLER 1985: fig. 16).
- Desmanodon crocheti in the broader trigonid of the lower
molars (PRIETO in press).
- Desmanodon daamsi in being larger, having less inflated
cusps in the lower molars and the complete division of the
mesostyle of the upper molars.
- Desmanodon ziegleri in being larger, having a more
divided mesostyle and reduced / absent metaconule / paraconule in the upper molars and in the longer and straighter
teres tubercle of the humerus.
- Desmanodon burkarti in the position of the mental foramen (under the p3, under p4 in D. burkarti), the less developed paraconule of the M2.
The new species is morphologically close to the Anatolian lineage D. minor / D. major. It differs from:
- Desmanodon major in its smaller size, the position of
the foramen mentale (under the p3), the more lingually
positioned paraconid of the m1 and m2, the continuous
labial cingulid of the m1 and m2, the interrupted metacingulum of the M2 and the more divided mesostyle of the
upper molars.
- Desmanodon minor in the single-rooted p3, the shorter
entoconid of the lower molars, the continuous labial cingulid of the m1 and m2, the presence of narrow labial
cinguli in the M1 and M2, the relatively longer teres
tubercle of the humerus and the less strong and more curved
mandible.
D e s c r ip tio n o f t he hol ot ype : The anterior part of the
mandible is broken somewhat 0.8 mm before the mental
foramen; the ascending ramus preserves only its lowermost
part; the thinner part of the horizontal ramus is found under
the contact point of p4 and m1; the mental foramen lies
under the p3; the p3, p4, m1, m2 and m3 are conserved in
situ, whereas the anteriorste tooth fell out of the mandible
and is conserved as isolated element; this tooth is difficult
to interpret. ENGESSER (1980: 123) described a mandible
fragment of D. minor in which the alveole of a small singlerooted p 2 and a double-rooted tooth are found before the
p 3. He cannot specify the nature of this tooth, p1 or canine.
In the mandible from Gratkorn a small alveole can be
observed before the p3; it has originally supported a small
single-rooted premolar; the ? p1/c lays originally before this
alveole.
The two-rooted ? p1/c is larger than the p 3; the high
protoconid is directed forwards; a very low crest descends
along its posterior wall; the tooth is surrounded by a cingulum, which becomes stronger on the posterior border of the
canine.
The overall morphology of the single-rooted p3 does not
differ from the ? p1/c; the protoconid is clearly lower and
the cingulid surrounding the tooth is relatively stronger; an
extremely reduced parastylid is present.
The protoconid of the two-rooted p4 is situated in the
anterior part of the tooth; a cingulum is present all around
the p4, forming a small parastylid at its anterior border; the
height of the protoconid of the p4 corresponds to the height
of the hypoconid of the m1.
The talonid of the m1 is broader than its trigonid; the thin
paracristid is higher than the postcristid; the thin precingulid
extends on the lingual wall of the paraconid; labially the
cingulids are continuous; the postcingulid ends lingually in
a strong entostylid; the crista oblica strongly descends from
the hypoconid to the base of the posterior wall of the
protocristid.
The m2 mainly differs from the m1 in trigonid and
talonid having about the same width; the trigonid basin is
also antero-posteriorly narrower than in m1 and the precingulid is stronger.
In the m3, the paraconid is not aligned with the meta- and
entoconid like in m1 and m2, but positioned somewhat
more centrally; the precingulid is as strong as in m2; the
labial walls of the proto- and hypoconid have no cingulid
(in the Figure 4 A1, the ectocingulid seems to be complete,
but this is not correct); the crista oblica descends strongly
from the hypoconid to the base of the posterior wall of the
protocristid, almost parallel to the entocristid.
Description of the paratypes: Humerus: The humerus
is broken around the head but preserved in full length; the
teres tubercle forms a long straight crest; the small pectoral
tubercle is situated in the middle of the shaft; the scalopine
ridge extends from the head to the medial side of the lesser
tubercle.
The lower molars basically do not differ from the holotype.
M1: The two M1 are either broken or corroded; the labial
border of the molar is strongly oblique; the mesostyle is
strongly divided; the metaconule is present; a cingulum is
present on both anterior and posterior walls of the protocone; three roots.
M2: The M2 differs from the M1 in its labial border,
which is not oblique; the parastyle is slightly more labially
positioned than the others labial styles; the mesostyle is
strongly divided; the metaconule is present; the paracingulum and metacingulum are missing; reduced cingula
are found on both the anterior and posterior sides of the
protocone; a very weak paraconule is present in unworn
specimens.
M3: The labial border of the molar is oblique backwards;
the metacrista is missing; the mesostyle is clearly divided;
the metacingulum is missing; an extremely reduced cingulum is present on the posterior wall of the protocone of one
M3 (out of four).
Di scussi on: The differences between Desmanodon and
Paratalpa were extensively discussed by VAN DEN HOEK
OSTENDE (1997). The teeth and humerus from Gratkorn
clearly belong to Desmanodon. Furthermore, only one mole
species is present in the locality and the attribution of the
humerus to the tooth sample is confident.
The overall morphology of the specimens clearly links D.
fluegeli to the lineage D. minor / D. major as defined by
Insectivores and bat from the late Middle Miocene of Gratkorn (Austria)
ENGESSER (1980). The new species shares characteristics
found either in D. minor (specifically the size, the entoconid
length) or D. major (especially the humerus, the singlerooted p3, the form of the mandible). Nevertheless, it is
not possible to interpret D. fluegeli as intermediate stadium
between the two Anatolian species. Finally, it cannot be
excluded that the differences observed in the new species
are correlated to geographical differentiation.
Family Soricidae FISCHER, 1814
Subfamily Heterosoricinae VIRET & ZAPFE, 1952
Genus Dinosorex ENGESSER, 1972
D ia g n o s is : ENGESSER (1972: 80-81), Differential diagnosis: ENGESSER (1972: 78-80), comments in ENGESSER
(1975: 662-663).
Ty p e s p e c ie s : Sorex sansaniensis LARTET, 1851.
O th e r s p e c ie s i nc l ude d: D. pachygnathus ENGESSER,
1972, D. zapfei ENGESSER, 1975, D. anatolicus VAN DEN
HOEK OSTENDE, 1995, D. grycivensis RZEBIK-KOWALSKA &
TOPACHEVSKY, 1997, D. engesseri RABEDER, 1998.
Dinosorex sp.
Figs. 2L-M, 5
M a te r ia l u n d m e a sure m e nt s: Right mandible with
incisor, a(?), m1- m3, left upper incisor, left P4 (LMJG
204.004, 160, 161): lower incisor: 8x2.1; a(?): 1.45x1.22;
m1: 2.5x (1.47 x 1.73); m2: 2.07x(1.67 x1.63); m3: 1.68x
(1.58 x 1.05); upper incisivor: 4.9 x2.78x2.35; P 4: 2.01 x2.
D e s c r ip tio n and di sc ussi on: Heterosoricinae are not
well represented in the Gratkorn sample but a very wellpreserved mandible was discovered. This mandible has a
divided massetic fossa and an acuspulate incisor. Therefore,
the lower jaw belongs to Dinosorex. The two other isolated
teeth are also attributed to the genus. In the jaw, the lower
incisor, the anteriormost antemolar and all the molars are in
situ. The posterololophid extends to the postero-labial (m1)
or labial (m2) wall of the entoconid. The teeth are coloured,
suggesting an original pigmentation. The posterior margin
of the incisor extends below the rear of the m1 trigonid. The
mental foramen is situated under the protoconid of the m2,
in the upper half of the horizontal ramus. A small postsymphyseal foramen is preserved below the rear of the m3,
slightly above the ventral margin of the jaw. The condyl is
divided into two facets, the ventral one being clearly larger.
The facets are connected by a narrow ridge.
The differences with Dinosorex sansaniensis, D. grycivensis, D. anatolicus and D. zapfei are clear:
- D. sansaniensis has a less robust mandible with four antemolars (see ENGESSER 1972: fig. 18b, 1975: pl. 3, fig. 3,
1980: pl. 8, fig. a) and isolated condylar facettes. The lower
molars are without entocristid. Furthermore, the posterolingual valley of the P4 is more developed.
113
- D. zapfei has 4 antemolars (including p 4, ENGESSER 1975:
pl. 5, fig. 3 b), a smaller m3 and lower molars without
entocristid.
- D. grycivensis has four antemolars and lower molars without entocristid (RZEBIK-KOWALSKA & TOPACHEVSKY 1997).
- D.anatolicus has a foramen mentale lying under the
anterior root of the m2 and the first two lower molars have a
large valley separating the posterolophid from the entoconid.
The specimens from Gratkorn differ from Dinosorex
pachygnathus in the clearly longer and narrower lower
incisor; the lingual borders of the lower molars from Gratkorn are convex, whereas they are almost straight in D.
pachygnathus; the horizontal ramus of the mandible is also
narrower; the m2 does not cover the postcingulid of the m1,
as is usually the case in Dinosorex; the ascending ramus
shows clear differences with the specimen figured by
ENGESSER (1972: fig. 18 a), especially in its anterior border
which is less convex. It is not possible to estimate precisely
the original number of antemolars, probably two with the
p4. The P4 differs clearly from the premolar of D. pachygnathus in the postero-lingual part of the tooth, with
extends more lingually. This characteristic is also noticed by
KÄLIN & ENGESSER (2001) for D. aff. pachygnathus of
Nebelbergweg (see below).
The main characteristic of D. engesseri from the Upper
Miocene of Austria is the crest developed between hypocone and metacone in the M1 and M2 (RABEDER 1998). These
elements are not preserved in Gratkorn. RABEDER (1998:
fig. 5) figures a fragmentary mandible from Götzendorf
which shares some morphologic similarities with the
mandible from Gratkorn although the processus angularis
is better developed. ZIEGLER (2005 a, 2006 a) reports the
presence of the species in the Upper Miocene localities
Schernham, Richardhof-Wald, Neusiedl am See, Richardhof-Golfplatz (all Austria) and Rudabánya (Hungary). The
incisor from Gratkorn is larger than in these localities,
whereas the molars are somewhat smaller. Furthermore, the
hypolophid of the m1 and m2 joins the entoconid on its
postero-labial wall, not clearly posteriorly like in D. engesseri. Indeed, this character can be slightly variable in the
Dinosorex species.
KÄLIN & ENGESSER (2001) report Dinosorex in Nebelbergweg (Switzerland). They notice some differences with
D. engesseri but refrain from describing a new species. The
incisor from Gratkorn, though also long, is nevertheless
shorter than those from Nebelbergweg. The molars are also
smaller, but, in the overall morphology, the specimens from
Gratkorn share great similarities with the material from
Nebelbergweg.
In size and morphology the Austrian lower molars are
not basically different from Dinosorex sp. from Sari Çay
(Turkey, ENGESSER 1980), but the tooth sample of this
locality is also very small.
The specimens from Gratkorn, in spite of the presence
of the especially well-preserved mandible, do not allow one
to reach confident conclusions regarding their specific
attribution.
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J. Prieto et al.
Fig. 5. Dinosorex sp. from Gratkorn. Right mandible (reversed, LMJG 204.004). A: lingual view; B: occlusal view;
C: labial view; D: distal view.
Order Chiroptera BLUMENBACH, 1779
Family Vespertilionidae G RAY, 1821
Genus Myotis KAUP, 1829
HORÁČEK (2001) for emended diagnosis and discussion on
this subject.
R e ma r k s : The systematic rank of Leuconoe BOIE, 1830,
as genus or subgenus of Myotis, is a subject of debate. Thus
the diagnosis/differential diagnosis of Myotis is dependent
of the generic attribution of Leuconoe. The reader is referred to BAUDELOT (1972), MENU (1987), ZIEGLER (2000) and
Ot her speci es i ncl uded (Oligocene-Pliocene species
only):
M. insignis (VON MEYER, 1845) (nomen dubium in ZIEGLER
2003b), M. salodorensis REVILLIOD, 1922, M. wuesti
KORMOS, 1934, M. schaubi KORMOS, 1934, M. steiningeri
Type s pecies : Vespertilio myotis BORKHAUSEN, 1797.
Insectivores and bat from the late Middle Miocene of Gratkorn (Austria)
KORMOS, 1934, M. kormosi HELLER, 1936, M. rapax
HELLER, 1936, M. aemulus HELLER, 1936, M. gunderheimensis HELLER, 1936, M. praevius HELLER, 1936, M. delicatus HELLER, 1936 (?= M. dasycneme subtilis KOWALSKI,
1956), M. exilis HELLER, 1936, M. sanctialbani VIRET,
1951, M. danutae KOWALSKI, 1956, M. helleri KOWALSKI,
1962, M. boyeri MEIN, 1964, M. ziegleri (BAUDELOT, 1972),
M. lavocati SIGÉ & MENU, 1992, M. minor Ziegler, 2000,
M. intermedius ZIEGLER, 2000, M. major ZIEGLER, 2000,
M. reductus ZIEGLER, 2003, M. bavaricus ZIEGLER,
2003.
cf. Myotis sp.
Fig. 2N-O
M a te r ia l a n d m e a sure m e nt s: 1 m1 (LMJG 204.162):
1.10 x (0.63 x 0,71); 1 M2 (LMJG 204.163): ~1,4x 1,7.
D e s c r ip tio n and di sc ussi on: Based on their size, the
two teeth are considered to belong to a single species. The
identification of chiropterian material based on isolated
teeth is always extremely difficult and the two molars from
Gratkorn are clearly insufficient to expect confident taxonomical conclusions. Nevertheless, based on the overall
morphology of the upper molar, it should be possible to
conclude that the two specimens are related to Myotis.
4. Biostratigraphic considerations
The insectivores are usually of minor interest regarding fine biostratigraphic correlations. However,
at a larger scale the use of well defined insectivore
lineages may be very useful. Such a lineage is reported for Schizogalerix in the Middle to Late Miocene from Turkey (ENGESSER 1980; SELÄNNE 2003).
Schizogalerix voesendorfensis is not directly integrated in this lineage, but in the morphologically close
S. voesendorfensis-S. zapfei-S. moedlingensis lineage
(in which SELÄNNE (2003) includes the Schizogalerix
species of the MN 12 and 13 from Greece).
The locality Vösendorf (type locality of S. voesendorfensis) belongs to Middle Pannonian sediments of
the mollusc Mytilopsis czjzeki Zone (= Congeria
suglobosa-“Zone” in RÖGL & DAXNER-HÖCK 1996)
with an age between 10.3 and ~10.5 Ma (HARZHAUSER et al. 2004).
The Swiss locality Nebelbergweg, also containing
this Schizogalerix species, cannot be at present dated
with absolute accuracy. Based on the evolutionary
level of the rodent Collimys longidens, this locality
may be younger than Hammerschmiede in Germany
(MAYR & FAHLBUSCH 1975, containing C. hiri), but
slightly older than the Hungarian Felsötarkany 3/2
(containing C. dobosi PRIETO & RUMMEL 2009), and
115
may be similar in age as the fauna from FelsötarkanyFelnémet (Hír 2006, 2007).
PRIETO et al. (in prep.) describe from Gratkorn a
new species of Cricetodontinae which is interpreted as
the ancestor of Cricetodon klariankae HÍR, 2007 from
Felsötarkany-Felnémet. Following this proposal, Gratkorn has to be older than Nebelbergweg. Based on
terrestrial gastropod and geologic evidences, HARZHAUSER et al. (2008) date Gratkorn at around 12 Ma.
Thus the Gratkorn specimens can be interpreted as the
first occurrence of Schizogalerix voesendorfensis.
This will imply that Schizogalerix voesendorfensis
shows no major evolutionary changes between Gratkorn and Vösendorf, in a time-span ranging from
~10.5 to 12 Ma.
Desmanodon fluegeli is a new species which
cannot be certainly integrated in the Anatolian lineage
D. minor / D. major and its stratigraphic value is therefore limited.
Dinosorex sp. cannot be directly related to a previously described species, although some similarities
with D. engesseri from the Upper Miocene (MN 9-10)
of Austria and Hungary are evident. However, the
material from Gratkorn also seems to be similar to the
species from Nebelbergweg. The correlation of the
Swiss locality is difficult but it is certainly older than
the reported sites containing D. engesseri.
The bat cf. Myotys sp. does not provide any stratigraphic information.
5. Geographical variations
The insectivore fauna from Gratkorn seems to be
related to eastern European / Anatolian faunas. The
most impressive species illustrating this observation is
Desmanodon fluegeli, which shares several morphologic features with D. minor / D. major from the
Turkey, although it cannot be directly integrated in this
lineage. This can be interpreted as parallel evolution
between lineages of the two regions, maybe as allo-or
parapatric speciation modes. Indeed ENGESSER (1980)
interprets the resemblance between Turkish and Austrian Schizogalerix as such a parallel evolution. In the
NAFB, the first unequivocal occurrence of Schizogalerix is found in Nebelbergweg, a locality which can
be confidently considered younger than Gratkorn. In
the locality Hammerschmiede, also older than Nebelbergweg, Schizogalerix is absent. In the much older
deposits of Anwil, Kleineisenbach and Giggenhausen
(~13.2 My), the genus is missing, but another galericine, Parasorex socialis is present. At present, it can
116
J. Prieto et al.
be hypothesised that Schizogalerix is an immigrant in
the NAFB, from eastern origin, which might explain
the resemblances of the Nebelbergweg specimen with
the Austrian lineage.
6. Environmental and ecological
implications
With four species, the α-diversity of the Gratkorn
insectivores is low. The difference with Anwil, for
example, is impressive (13 species, ENGESSER 1972).
A bias due to insufficient collected material can be
excluded since the chiropterans, which are generally
extremely rare in the stratified deposits, are recorded
in Gratkorn. This low diversity may be better explained by taphonomic bias. Indeed the small mammal
remains in this locality were largely accumulated by
owl, as proved by the find of fossilized pellet remains.
Furthermore, these finds indicate that the pellets were
not transported before fossilisation because even
transport of small distance can rapidly disintegrated
them and lead to the dispersal of the animal remains
(for example TERRY 2004). It may indicate that the
prey accumulation illustrates local conditions, in
which the alimentary preferences of the owls, size and
abundance of the prey for example, are extremely
reflected.
The ecological preferences of the galericines are
still a subject of debate. Based on the ecology of
extent gymnures from South-east Asia, ZIEGLER
(2005b: 150) proposed that the galericines have preferred moist forested areas and explain their wide
geographic distribution in the Miocene by a wide
distribution of their habitat. By this, he challenges the
interpretation of VAN DEN HOEK OSTENDE (2001) who
proposes a larger panel of ecologic adaptations for the
Galericini.
The presence of large samples of Parasorex socialis
in Middle Miocene fissure fillings of the Franconian
Alb, fossil sites which are reputed to be dryer than the
contemporary sites of the NAFB, is explained by
ZIEGLER (2005b) by the hunting facilities of the owls
in the basin. This seems somewhat exaggerated.
Indeed the deposit place of isolated pellets can be
extremely distant from the capture place of the prey
(KOWALSKI 1990), but the accumulation of a large
number of pellets is mostly found around the bird
nests. MICHELAT & GIRAUDOUX (1991) studied the
hunting patch of Tyto alba during the nesting using
radio senders. They conclude that the owl never
exceed the distance 5.6 km for its hunt (1.5 km on
average).
The northern limit of the NAFB deposit in Germany is found in Adelschlag (MN 5, REICHENBACHER
et al. 2004). If this limit is assumed to be the same at
the time of fossil accumulation in Petersbuch, the
minimal distance to attain the basin from Petersbuch
would be 10 km, which seems to be too far for the
owls compared with modern conditions. Alternative
hypothesizes can be proposed to explain the presence
of Parasorex in the Petersbuch fissures:
- Parasorex socialis could have been adapted to the
environmental conditions of the Frankonian Alb (environmental differences between plateau and basin
were not significant or the species was an ubiquiste).
- Temporary migrations of Parasorex in the Franconian Alb can also be hypothesized (dispersal phase
period of the young for example, PRIETO 2007).
VAN DEN HOEK OSTENDE (2001) listed a series of
dental characters in which Parasorex and Schizogalerix differ from Galerix. Since these characters
lead in the enlargement of the grinding surface, he
interprets them as an adaptation to a more herbivorous
diet. Furthermore, he remarks on the high position of
the condyle of the lower jaws of Parasorex ibericus.
The high position of the condyle is in extant desman
moles a character of more herbivorous diet (RÜMKE
1985). Indeed VAN DEN HOEK OSTENDE uses for his
observation two species of Galerix (G. exilis and G.
stehlini) and one species of Parasorex. The ascending
ramus is preserved in P. socialis mandibles of Petersbuch 6 (Fig. 5 and ZIEGLER 2005 b: fig. 1B). In this
species the condyle is higher positioned than in G.
exilis but the height difference is clearly smaller than
in P. ibericus. The resemblance between the P. socialis
and S. voesendorfensis mandibles are impressive
(Figure 3), which permits one to hypothesize that the
two species could have had a comparable diet.
Schizogalerix is by far the most common small
mammal in the Gratkorns sample, followed by a new
Cricetodontinae species (PRIETO et al. in prep.). Both
species are relatively large small mammals and correspond in their size. From this observation we can
conclude that they also corresponded in their ecology
and abundance within the optimal prey type for the
owls, which avoid close forested area during their hunt
(MICHELAT & GIRAUDOT 1993).
Desmanodon fluegeli n. sp. is the only mole recorded in Gratkorn. Although taphonomical biases
have to be taken in account (see above), it is interesting to notice that Desmanodon is also the only
talpid found in the Early and lower Middle Miocene of
Spain (VAN DEN HOEK OSTENDE 2003: 300). During
Insectivores and bat from the late Middle Miocene of Gratkorn (Austria)
this time (Spanish zone B to Da) relatively humid
conditions are proposed (VAN DER MEULEN & DAAMS
1992; VAN DEN HOEK OSTENDE 2003). PRIETO (in
press) describes a new Desmanodon species from the
late Middle Miocene from Germany. Based on faunal
differences with the contemporaneous Swiss locality
Anwil, where this species is missing, he concludes
that the mole avoids lake environments. Furthermore,
the morphology of the Desmanodon humerus indicates that the animal was not extremely adapted to
fossorial life-style. The vertebrate producing layer at
the base of the clay pit of Gratkorn corresponds to a
paleosol showing root traces. GROSS et al. (2007)
interpreted these strata to be deposits of a vegetated
alluvial plain with a moist soil cover and open areas.
This reconstruction agrees with the hypothesized
ecological preferences of Desmanodon.
7. Conclusions
The insectivores from Gratkorn illustrate a new faunal
evolutionary level in Austria and are therefore of
primary importance for the comprehension of the
faunal evolution and exchanges in the late Middle
Miocene of Central Europe. Furthermore, the excellent preservation of the bony and dental remains
in this locality allows one to approach determining
the ecology of these extant species. The fauna from
Gratkorn contains a very large panel of vertebrates
(small and big mammals, reptilians, amphibians,
birds, GROSS et al. 2009) and will, in the near future,
permit greater understanding of environmental and
climatic conditions in the Late Sarmatian of Central
Europe.
Acknowledgements
LARS W. VAN DEN HOEK OSTENDE (Leiden) is especially
thanked for his comments and fruitful discussions. REINHARD ZIEGLER (Stuttgart) provided an excellent review. We
thank the “Wietersdorfer & Peggauer Zementwerke” (JOSEF
PLANK) for permitting studying the clay pit and the Marktgemeide Gratkorn for providing the facilities. We are also
thankful to NORBERT WINKLER (Graz, preparation), GERNOT
TONAUER and GERALD PHILIP (students at the University of
Graz, both field work). CHRISTOPHER “M. H.” MURGATROYD
(Munich) improved the English style. This project was supported by DFG grant BO 1550/7-2.
References
BAUDELOT, S. (1972): Etude des Chiroptères, Insectivores et
Rongeurs du Miocène de Sansan (Gers). – Thèse de
doctorat Université Toulouse, 496: 1-364.
117
BÖHME, M. (2003): Miocene Climatic Optimum: evidence
from Lower Vertebrates of Central Europe. – Palaeogeography, Palaeoclimatology, Palaeoecology, 195 (3-4):
389-401.
BÖHME, M., ILG, A. & WINKELHOFER, M. (2008): Late
Miocene “washhouse” climate in Europe. – Earth and
Planetary Science Letters, 275 (3-4): 393-401, doi:
10.1016/j.epsl.2008.09.011.
BRUIJN, H. DE, MAYDA, S., HOEK OSTENDE, L. W. VAN DEN,
KAYA, T. & SARAÇ, G. (2006): Small mammals from the
Early miocene of Sabuncubeli (Masina, S. W. Anatolia,
Turkey). – Beiträge zur Paläontologie, 30: 57-87.
DOUKAS, C. S. & HOEK OSTENDE, L. VAN DEN (2006):
Insectivores (Erinaceomorpha, Soricomorpha; Mammalia) from Karydia and Komotini (Thrace, Greece;
MN 4/5). – Beiträge zur Paläontologie, 30: 109-131.
ENGESSER, B. (1972): Die obermiozäne Säugertierefauna
von Anwil (Baselland). – Tätigkeitsberichte der Naturforschenden Gesellschaft Baselland, 28: 37-363.
– (1975): Revision der europäischen Heterosoricinae
(Insectivora, Mammalia). – Eclogae Geologicae Helvetiae, 68 (3): 649-671.
– (1980): Insectivora and Chiroptera (Mammalia) aus dem
Neogen der Türkei. – Schweizerische Paläontologische
Abhandlungen, 102: 45-149.
GROSS, M. (2008): A limnic ostracod fauna from the surroundings of the Central Paratethys (Late Middle Miocene / Early Late Miocene; Styrian Basin; Austria). –
Palaeogeography, Palaeoclimatology, Palaeoecology,
264: 263-276.
GROSS, M., BÖHME, M. & PRIETO, J. (2009): Gratkorn – A
new late Middle Miocene vertebrate fauna from Styria
(Late Sarmatian, Austria). – Geophysical Research
Abstracts, vol. 11, EGU2009-7091, EGU General
Assembly 2009.
GROSS, M., FRITZ, I., PILLER, W. E., SOLIMAN, A., HARZHAUSER, M., HUBMANN, B., MOSER, B., SCHLOGER, R.
& SUTTNER, T. J. (2007): The Neogene of the Styrian
Basin – Guide to Excursion. – Jaannea – Geologie und
Paläontologie, 9: 117-193.
HARZHAUSER, M., GROSS, M. & BINDER, H. (2008): Biostratigraphy of Middle Miocene (Sarmatian) wetland
systems in an Eastern Alpine intramontane basin (Gratkorn Basin, Austria): the terrestrial gatropod approach. –
Geologica Carpathica, 59 (1): 45-58.
HARZHAUSER, M., DAXNER-HÖCK, G. & PILLER, W. E.
(2004): An integrated stratigraphy of the Pannonian
(Late Miocene) in the Vienna Basin. – Austrian Journal
of Earth Sciences, 95/96: 6-19.
HÍR, J. (2006): Late Astaracian (Late Sarmatian) Lagomorphs and Rodents from Felsötárkány-Felnémet
(Northern Hungary). – Beiträge zur Paläontologie, 30:
155-173.
– (2007): Cricetodon klariankae n. sp. (Cricetodontini,
Rodentia) from Felsötárkány-Felnémet (Northern Hungary). – Fragmenta Paleontologica Hungarica, 24-25:
16-24.
HOEK OSTENDE, L.W. VAN DEN (1997): Insectivore faunas
from the Lower Miocene of Anatolia. Part 4: The genus
Desmanodon (Talpidae) with the description of a new
species from the Lower Miocene of Spain. – Pro-
118
J. Prieto et al.
ceedings of the Koninklijke Nederlandse Akademie van
Wetenschappen, 100 (1-2): 27-65.
– (2001): A revised generic classification of the Galericini
(Insectivora, Mammalia) with somes remarks on their
palaeogeography and phylogeny. – Géobios, 34: 681695.
– (2003): Insectivores (Erinaceomorpha, Soricomorpha,
Mammalia) from the Ramblian of the Daroca-Calamocha area. – In: LÓPEZ-MARTINEZ, N., PELÁEZCAMPOMANES, P. & HERNÁNDEZ FERNÁNDEZ, M. (Eds.):
En torno a fósiles de mamiferos: datación, evolución y
paleoambiente. – Coloquios de Paleontología, Volumen.
Extraordinario, 1: 281-310.
HORÁ ČEK, I. (2001): On the early history of vespertilionid
bats in Europe: the Lower Miocene record from the
Bohemian Massif. – Lynx, 32: 123-154.
HUTCHINSON, J. H. (1968): Fossil Talpidae (Insectivora,
Mammalia) from the later Tertiary of Oregon. – Bulletin
of the Museum of Natural History of the University of
Oregon, 11: 1-117.
– (1974): Notes on the type species of European Miocene
Talpidae and tentative classification of Old World
Tertiary Talpidae. – Géobios, 7: 211-256.
KÄLIN, D. (1993): Stratigraphie und Säugetierfaunen der
oberen Süsswassermolasse der Nordwestschweiz. –
Dissertation an der Eidgenössischen Technischen Hochschule Zürich. – 238 pp.
KÄLIN, D. & ENGESSER, B. (2001): Die jungmiozäne Säugertierfauna vom Nebelbergweg bei Nunningen (Kanton
Solothurn, Schweiz). – Schweizerische Paläontologische
Abhandlungen, 121: 1-61.
KOWALSKI, K. (1990): Some problems of the taphonomy of
small mammals. – In: FEJFAR, O. & HEINRICH, W.-D.
(Eds.): International Symposium Evolution, Phylogeny
and Biostratigraphy of Arvicolids, 285-296; Praha
(Czech Geological Survey).
MAYR, H. & FAHLBUSCH, V. (1975): Eine unterpliozäne
Kleinsäugerfauna aus der Oberen Süßwasser-Molasse
Bayerns. – Mitteilungen der Bayerischen Staatssammlung für Paläontologie und historische Geologie, 15:
91-111.
MEULEN, A. J. VAN DER & DAAMS, R. (1992): Evolution of
Early-Middle Miocene rodent faunas in relation to longterm paleoenvironmental changes. – Paleogeography,
Paleoclimatology, Paleoecology, 93: 227-253.
MENU, H. (1987): Morphotypes dentaires actuels et fossiles
des chiroptères vespertilioninés. 2ème partie: implications systématiques et phylogéniques. – Palaeovertebrata, 17 (3): 77-150.
MICHELAT, D. & GIRAUDOUX, P. (1991): Dimension du
domaine vital de la chouette effraie Tyto alba pendant la
nidification. – Alauda, 59 (3): 137-142.
– (1993): Relation proies-prédateur-paysage chez la
chouette effraie Tyto alba pendant l’élevage des jeunes. –
Alauda, 61 (2): 65-72.
NOVACEK, M. J., BOWN, T. M. & SCHLNAKER, D. (1985): On
the Classification of the Early Tertiary Erinaceomorpha
(Insectivora, Mammalia). – American Museum Novitates, 2813: 1-22.
PAPP, A. (1951): Das Pannon des Wiener Beckens. – Mitteilungen der Geologischen Gesellschaft in Wien,
39-41: 99-193.
PRIETO, J. (2007): Kleinsäuger-Biostratigraphie und Paläoökologie des höheren Mittelmiozäns (MN 8) Bayerns:
Spaltenfüllungen der Fränkischen Alb und Lokalitäten
der Oberen Süßwassermolasse im Vergleich. – Unpublished Dissertation LMU München. – 213 pp.
– (in press): The Middle Miocene mole Desmanodon
crocheti sp. nov. (Talpidae, Mammalia): the last representative of the genus in the North Alpine Foreland
Basin. – Paläontologische Zeitschrift. DOI: 10.1007/
51254-009-003-0
PRIETO, J. & RUMMEL, M. (2009a): Erinaceidae (Mammalia,
Erinaceomorpha)
from
the
Middle
Miocene
fissure filling Petersbuch 68 (Southern Germany). –
Zitteliana. A 48/49: 75-88.
– (2009b): Evolution of the genus Collimys DAXNERHÖCK, 1972. (Rodentia, Cricetidae) – A key to Middle
to Late Miocene biostratigraphy in Central Europe. –
Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 252 (2): 237-247.
RABEDER, G. (1973): Galerix und Lanthanotherium (Erinaceidae, Insectivora) aus dem Pannon des Wiener
Beckens. – Neues Jahrbuch für Geologie und Paläontologie, Monatshefte, 1973: 429-446.
– (1998): Dinosorex (Insectivora, Mammalia) aus dem
Miozän von Österreich. – Geologisch-Paläontologische
Mitteilungen Innsbruck, 23: 117-126.
REICHENBACHER, B., BÖHME, M., HEISSIG, K., PRIETO, J. &
KOSSLER, A. (2004): New approach to assess biostratigraphy, palaeoecology and past climate in the South
German Molasse Basin during the Early Miocene (Ottnangian, Karpatian). – Courier Forschungsinstitut
Senckenberg, 249: 71-89.
REUMER, J. W. F. (1984): Ruscinian to Early Pleistocene
Soricidae (Insectivora, Mammalia) from Tegelen and
Hungary. – Scripta Geologica 73: 1-173.
RÖGL, F. & DAXNER-HÖCK, G. (1996): Late Miocene Paratethys correlations. – In: BERNOR, R. L., V. FAHLBUSCH
& MITTMANN, W. (Eds.): The Evolution of Western
Eurasian Neogene Mammal Faunas, 47-55; New York
(Columbia University Press).
RÜMKE, C. G. (1985): A review of fossil and recent Desmaninae (Talpidae, Insectivora). – Utrecht Micropaleontological Bulletin, special publication 4: 1-241
RZEBIK-KOWALSKA, B. & TOPACHEVSKY, V. O. (1997):
Insectivora (Mammalia) from the Miocene of Grytsiv in
Ukraine. 1. Heterosoricidae VIRET & ZAPFE 1951. –
Acta Zoologica Cracoviensia, 40 (2): 237-247.
SELÄNNE, L. (2003): Genus Schizogalerix (Insectivora). –
In: FORTELIUS, M., KAPPELMAN, J., SEN, S. & BERNOR,
R. L. (Eds.): Geology and Paleontology of the Miocene
Sinap Formation, Turkey, 69-89; New York (Columbia
University Press).
TERRY, R. C. (2004): Owl Pellet Taphonomy: A Preliminary
Study of the Post-Regurgitation Taphonomic History of
Pellets in a Temperate Forest. – Palaios, 19 (5): 497-506.
ZIEGLER, R. (1985): Talpiden (Mammalia, Insectivora) aus
dem Orleanium und Astaracium Bayerns. – Mitteilungen
der Bayerischen Staatssammlung für Paläontologie und
historische Geologie, 25: 131-175.
Insectivores and bat from the late Middle Miocene of Gratkorn (Austria)
– (2000): The bats (Chiroptera, Mammalia) from the Late
Oligocene Fissure Fillings Herrlingen 8 and Herrlingen
9 near Ulm (Baden-Württenberg). – Senckenbergiana
lethaea, 80 (2): 647-683.
– (2003 a): Moles from the late Middle Miocene of South
Germany. – Acta Palaeontologica Polonica, 48: 617-648.
– (2003 b): Bats (Chiroptera, Mammalia) from Middle
Miocene karstic fissure fillings of Petersbuch near Eichstätt, Southern Franconian Alb (Bavaria). – Géobios, 36:
447-490.
– (2005 a): The insectivores (Erinaceomorpha and Soricomorpha, Mammalia) from the Late Miocene hominoid
locality Rudabánya. – Palaeontographia Italica, 90:
53-81.
– (2005 b): Erinaceidae and Dimylidae (Lipotyphla) from
the Upper Middle Miocene of South Germany. –
Senckenbergiana lethaea, 85: 131-152.
– (2006 a): Insectivores (Lipotyphla) and Bats (Chiroptera) from the Late Miocene of Austria. – Annalen des
Naturhistorischen Museums Wien, 107A: 93-106.
– (2006 b): Miocene Insectivores from Austria and Germany – An Overview. – Beiträge zur Paläontologie, 30:
481-494.
119
ZIEGLER, R. & DAXNER-HÖCK, G. (2005): Austria. – In:
HOEK OSTENDE, L.W. VAN DEN, DOUKAS, C. S. &
REUMER, J. W. F. (Eds.): The fossil record of the Eurasion Neogene Insectivores (Erinaceomorpha, Soricomorpha, Mammalia), Part I. – Scripta Geologica,
Special Issue, 5: 11-29.
Manuscript received: May 12th, 2009.
Revised version accepted by the Stuttgart editor: July 23rd,
2009.
Addresses of the authors:
JÉRÔME PRIETO, CHRISTINE BÖHMER, Department of Earthand Environmental Science, Section Palaeontology, Ludwig-Maximilians-University Munich, Richard-Wagner-Str.
10, D-80333 Munich, Germany;
e-mails: j.prieto@lrz.uni-muenchen.de;
MARTIN GROSS, Landesmuseum Joanneum, Raubergasse
10, A-8010 Graz, Austria;
e-mail: martin.gross@museum-joanneum.at
MADELAINE BÖHME, Eberhard-Karls-University Tübingen,
Institute for Geoscience, Sigwartstr. 10, D-72076 Tübingen,
Germany;
e-mail: m.boehme@ifg.uni-tuebingen.de