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Sandelzhausen field photos 1963–1975 (all photos by V.F., except where noted). a Occasional find of vertebrate remains during an institutional trip in July 1963. Richard Dehm with his graduate student Volker Fahlbusch. Photo by Herbert Hagn. b View of the north side of the gravel pit in May 1964. Afterwards, the pit was abandoned and partly refilled. The fossiliferous marl layer is situated in the upper third of a gravel package termed ‘‘N ̈rdlicher Vollsch- otter.’’ c First systematic investigation of the profile for its fossil distribution and sedimentology in May 1966. Definition of the layers A , B , C , D with sublayers B1 , B2 , C1 , C2 , and C3 (see Fahlbusch and Gall 1970). d Nearly complete vertical section with layers A – E . August 1972. e An E-W section exposed at the end of the excavation period in September 1973 showing continuously developed layers. This is typical for most of the western part of the Sandelzhausen site, but not for the eastern part excavated during the 1990s, where the thickness of layers is more variable. f Preserving a vertical section by gluing and transferring the sediment in situ onto fabric in August 1970. Several of these sections are housed in the BSPG and one is on exhibit in the City Museum of Mainburg. g First complete skull find: the rhinocerotid ‘‘Karl-Otto.’’ October 1970. Photo by M. Kindl. h Systematic excavation in September 1970. The area is divided by a grid of north-oriented square meters, each of which was worked on one by one down to the base. Photo by Wolf-Dieter Grimm. i Systematic excavation in August 1972. The upper parts of the marl layer had been removed, as they were thought to be barren of vertebrate remains at that time. j A frequently recurring problem during all excavation periods was the complete watering of the excavation site by strong rain events. August 1970
… 
Sandelzhausen field photos 1997–2001 (all photos by M.M.). a The fieldwork was conducted mostly with the help of volunteers, some of whom were active for several years, such as F. Reinfelder, N. Ballerstaedt, H. Hinle, and J. Herrlen ( left to right , 1997). b Rewarding finds: well-preserved molar of Gomphotherium subtapiroideum (1997). c – e In a line just a few meters off the eastern border of the fossil site, a higher concentration of fossils was observed. Sometimes, plaster near plaster had to be made, and the next fossils were encountered even before the plasters could be removed (lower jaws of a juvenile Gomphotherium in c , close-ups of later excavation stages d , e ; 1997). f R. Liebreich measuring length and position of a Gomphotherium tusk. Inset: plaster package number 5,000. More than 6,000 larger fossils and plaster packages were recovered up to the end of the campaign in 2001 (picture 1997). g Old site west of the Prof.- Dehm-Straße, with well-exposed layers A – D . The weakly coalified ‘‘coal-layer’’ is visible as dark band in the middle of the profile (1997). h With the mobile drilling equipment of the Bayerisches Geologisches Landesamt (Bavarian Geological Survey, now part of the Bayerisches Landesamt f ̈r Umwelt) ten cores (e.g., inset figure) were taken to find the extent of the fossiliferous layer towards the north and east of the new digging site (January 1998). i For the University of Munich, 1 week each year was reserved for geology students for an educational digging course. Twice, digging courses were also arranged for students from Mainz and Vienna (this picture, 2000). k The high layers E – F are exposed here on the left (southern) side. Selected material was put into sacks for screen washing (2001)
… 
The Sandelzhausen Meeting 2005 in Mainburg (all photos by M.M.). a Participants of the Sandelzhausen meeting: 1 Gertrud E. R ̈ssner (Munich), 2 Volker Fahlbusch (Munich), 3 Daria Petruso (Palermo, Italy), 4 Wilma Wessels (Utrecht, The Netherlands), 5 Ursula B. G ̈hlich (Munich, now Vienna, Austria), 6 Chiara Angelone (Roma, Italy), 7 Margery C. Coombs (Amherst, Massachusetts, USA), 8 Josef Egger (Mayor of Mainburg), 9 Henriette Jechorek (G ̈rlitz), 10 Thorsten Kowalke (Munich), 11 Dietmar Jung (Munich), 12 Reinhard Ziegler (Stuttgart), 13 Gerhard Doppler (Munich), 14 Wolfgang Witt (G ̈ndlkofen), 15 Pierre Mein (Lyon), 16 John Damuth (Santa Barbara, CA, USA), 17 Norbert Schmidt-Kittler (Mainz), 18 Jan van der Made (Madrid, Spain), 19 Ralph Annau (Munich), and 20 Kurt Heissig (Munich). Participants not shown in the picture: Madelaine B ̈hme (Munich), Hans Hinle (Kelheim), Thomas Kaiser (Hamburg), Markus Moser (Munich), Doris Nagel (Vienna), Jerome Prieto (Munich), Bettina Reichenbacher (Munich), Zbigniew Szyndlar (Krakow), and Clara Stefen (Dresden). b During session in the City Hall of Mainburg. c Opening ceremony of the information panels at the filled-in Sandelzhausen pit. d Reconstruc- tion of the landscape of Sandelzhausen 16 million years ago by artist Wenzel Balat. Shown are the rhinos Plesiaceratherium ( foreground ) and Lartetotherium ( background ), and the chalicothere Metaschizotherium ( right , with hind legs that are too short; M. Coombs, oral communication, at symposium 2005). The painting is also shown on the left panel in Fig. 4c. e Volker Fahlbusch with Josef Egger, first Mayor of Mainburg and generous supporter of the field campaigns of 1994–2001 and the meeting in 2005, with his Sandelzhausen digging shirt (cartoon painting by Bettina Schenk, formerly Munich, now Vienna)
… 
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INTRODUCTION
The fossil lagersta
¨tte Sandelzhausen (Miocene; southern
Germany): history of investigation, geology, fauna,
and age
Markus Moser ÆGertrud E. Ro
¨ssner Æ
Ursula B. Go
¨hlich ÆMadelaine Bo
¨hme Æ
Volker Fahlbusch
Received: 6 May 2008 / Accepted: 29 August 2008 / Published online: 7 February 2009
Springer-Verlag 2009
Introduction
The Miocene fossil lagersta
¨tte Sandelzhausen in southern
Germany ranks among the most important Neogene fossil
terrestrial localities in Europe, due to its exceptional record
of both macro- and microvertebrates as well as inverte-
brates. From its discovery in 1959 until its final closure in
2001, the site has been the subject of two long-term, sys-
tematic digging campaigns (1969–1975 and 1994–2001) as
well as occasional collecting by the Ludwig-Maximilians-
Universita
¨t (LMU) together with the Bayerische Staats-
sammlung fu
¨r Pala
¨ontologie und Geologie (BSPG; both
Munich). In addition to a few plant remains a fossil fauna
comprising more than 200 taxa and 50,000 identifiable
specimens (Fahlbusch 2003, and this paper Table 1) has
been collected including ostracods, molluscs, and, espe-
cially, vertebrates of all groups, with an emphasis on
mammals (all material housed in BSPG, Munich). Though
the vertebrate remains are almost exclusively disarticulated,
they represent a variety of both autochthonous and allo-
chthonous environments and they document extensive
series of ontogenetic stages and within-species variability.
Scientific results derived from Sandelzhausen have been
published since 1970 and, ultimately, 26 specialists
involved in Sandelzhausen research came together in Sep-
tember 2005 at the Sandelzhausen Symposium in Mainburg
to discuss the latest results on all faunistic, ecological, and
genetic aspects of the fossil site (Fig. 5a, b; Anonymous
2005,Ro
¨ssner 2006a,Ro
¨ssner and Fahlbusch 2006). The
present volume resulted from that meeting and presents new
contributions to the fossil fauna of Sandelzhausen. The
purpose of this paper is to give a short overview and a
summary of previous work on the Sandelzhausen fossil site,
to document the fieldwork and the geological context, and
to discuss the age of the fauna. A German-language sum-
mary article including the most recent faunal list, details of
the excavation, and the complete publication list up to 2002,
including popular accounts (e.g., Fahlbusch and Liebreich
1996), can be found in Fahlbusch (2003). A partly synthetic
paper on the genesis and ecology of the site compiling all
available research results and new evidence about the fossil
lagersta
¨tte will be published elsewhere (in preparation).
Geological and geographical setting
The fossil site of Sandelzhausen near Mainburg, 60 km
north of Munich (Mu
¨nchen) (Fig. 1)(R
44
85 540–610, H
53
87 680–720 Gradabteilungsblatt 7336 Mainburg, 1:25 000,
Bayerisches Landesvermessungsamt Mu
¨nchen 1959; GPS
4837036.900,1148011.600; 493 m height above sea level) is
located within the Molasse Basin, the subsiding North
V. Fahlbusch: deceased.
M. Moser (&)G. E. Ro
¨ssner
Bayerische Staatssammlung fu
¨r Pala
¨ontologie und Geologie,
Richard-Wagner-Strasse 10, 80333 Munich, Germany
e-mail: dml_moser@web.de
G. E. Ro
¨ssner
e-mail: g.roessner@lrz.uni-muenchen.de
U. B. Go
¨hlich
Geological–Paleontological Department,
Natural History Museum of Vienna, Burgring 7,
1010 Vienna, Austria
e-mail: ursula.goehlich@nhm-wien.ac.at
M. Bo
¨hme V. Fahlbusch
Sektion Pala
¨ontologie, Department fu
¨r Geo- und
Umweltwissenschaften, Ludwig Maximilians Universita
¨t
Mu
¨nchen, Richard-Wagner-Strasse 10, 80333 Munich, Germany
e-mail: m.boehme@lrz.uni-muenchen.de
123
Pala
¨ontol Z (2009) 83:7–23
DOI 10.1007/s12542-009-0012-x
Alpine Foreland Basin which received sediment load
mainly from the Alps (Lemcke 1988, Schwerd et al. 1996:
142 ff). At the beginning of the Miocene, this originally
marine marginal basin of the Paratethys, itself a Cenozoic
side arm of the Tethys (Kova
´c
ˇet al. 2007), became a relic
basin as the Paratethys continuously retreated eastwards.
The nonmarine younger Molasse deposits from late Early
Miocene (Early Karpatian) to early Late Miocene (Early
Pannonian), the so-called Upper Freshwater Molasse (Ob-
ere Su
¨ßwassermolasse, or OSM), document the final filling
up of the basin under freshwater conditions (Doppler 1989;
Doppler et al. 2000). During that time, the retreating sea
level gave rise to what became predominantly a wetland
with a mosaic of marshes, fens, peatlands, forests, and with
permanent and temporary water bodies such as ponds and
braided rivers, all draining westwards. This palaeoenvi-
ronmental setting is shown conclusively by the
sedimentological record (Lemcke et al. 1953; Blissenbach
1957; Unger 1989; Schmid 2002; Seehuber 2002) as well
as by ecological inference derived from the fossil record of
plants, invertebrates, and vertebrates (e.g., Fahlbusch et al.
1974; Gregor 1982a; Schwarz and Reichenbacher 1989;
Sach 1999;Go
¨hlich 2002;Bo
¨hme 2003;Bo
¨hme and Rei-
chenbacher 2003; Sach et al. 2003; Reichenbacher et al.
Table 1 Number of taxa and
references for Sandelzhausen
fossil groups
Group Minimum number
of taxa
References
Plantae 3 Jung in Fahlbusch and Gall (1970); Gregor (1982a,b); Gregor
et al. (1989); Jung (1996)
Ostracoda 14 Witt (1998)
Mollusca 69 Gall (1972,1973); Moser et al. (2009 this volume)
Teleostei 4 Bo
¨hme (1999b;2009)
Amphibia 12 Bo
¨hme (1999b;2009)
Reptilia 29 Scherer (1973,1981); Schleich (1981,1983,1984); Bo
¨hme
(1999a,b,2009); Szyndlar (2009 this volume)
Aves 14 Go
¨hlich (2002,2003)
Marsupialia 1 Ziegler (2000)
Lipotyphla 17 Ziegler (1999,2000)
Chiroptera 2 Ziegler (2000)
Carnivora 9 Morlo et al. (2004); Nagel et al. (2009 this volume)
Rodentia 18 Fahlbusch (1964,1975); Mayr (1979); Heissig (1997); Daams
et al. (1999); Hugueney (1999); Ka
¨lin (1999); Rummel
(1999); Ziegler (2005); Bruijn (2009 this volume); Stefen
(2009 this volume); Wessels and Reumer (2009 this volume)
Lagomorpha 3 Fahlbusch et al. (1974), Angelone (2009 this volume)
Pholidota 1 Fahlbusch (1998), Koenigswald (1999)
Perissodactyla 5 Heissig (1972,1999); Abusch-Siewert (1983); Yan and Heissig
(1986); Peter (1999,2002); Coombs (2009 this volume);
Kaiser (2009 this volume); Schulz and Fahlke (2009 this
volume); Tu
¨tken and Vennemann (2009 this volume)
Artiodactyla 7 Schmidt-Kittler (1971); Fahlbusch (1977); Ko
¨hler (1993);
Ro
¨ssner (1997,2002,2004,2009); Gentry et al. (1999);
Kaiser and Ro
¨ssner (2007); Kaiser (2009 this volume); Made
(2009 this volume); Tu
¨tken and Vennemann (2009 this
volume)
Proboscidea 2 Schmidt-Kittler (1972); Go
¨hlich (1998,1999,2009); Tu
¨tken
and Vennemann (2009 this volume); Calandra et al. (2009)
Total fauna 210 Fahlbusch and Gall (1970); Fahlbusch et al. (1974); Fahlbusch
(1976b); Fahlbusch (2003); Moser et al. (this paper)
Fig. 1 Location and lithology of the Sandelzhausen fossil lagersta
¨tte.
In the aerial photo, the position of the areas of systematic excavations
west and east of the Prof.-Dehm-Straße are indicated by black outlines.
The eastern area was exploited 1996–1998, while the western part was
exploited both in the campaigns of 1969–1975 and 1994–2001 (for a
detailed plan of grid squares with the years of excavation see Fahlbusch
2003). Not indicated are sporadic diggings from 1959 until 1968, and
in 1996 (see text for history). Triangles and circles show the position
of selected profiles and logs, respectively, which are represented
schematically below by small lithological columns (horizontally
normalized to the ‘‘coal layer’’). The larger column (right) shows
lithological details and the subdivision into layers ADfor the profile at
grid square meter 10 G, which was sampled in 5-cm intervals in 1969
(see also Moser et al. 2009 this volume). The subdivision into layers
had been coined already in 1966 and was retained as being useful ever
since (photo of test profile in lower left corner, photo by V.F.)
c
8 M. Moser et al.
123
The fossil lagersta
¨tte Sandelzhausen (Miocene; southern Germany) 9
123
10 M. Moser et al.
123
2004;Ro
¨ssner 2004; Heissig 2006; Reichenbacher and
Prieto 2006;Ro
¨ssner 2006b; Eronen and Ro
¨ssner 2007).
Plants are rarely preserved in most vertebrate-rich
localities (e.g., Sandelzhausen), however, a rich palaeobo-
tanical fossil record in favorable localities shows that,
besides an azonal vegetation surrounding the water areas,
the hinterland was composed of evergreen to deciduous
zonal forests and woodlands with legumes, indicating humid
subtropical to warm temperate climatic conditions contain-
ing forests (Schmidt 1976,1980; Gregor 1982a,b; Schmid
and Gregor 1983; Gregor et al. 1989; Spitzlberger 1989;
Selmeier 1989; Riederle and Gregor 1997; Schweigert 1992;
Jechorek and Kovar-Eder 2004;Bo
¨hme et al. 2007).
The Sandelzhausen fossil section is embedded within
gravel beds termed ‘‘No
¨rdlicher Vollschotter,’’ which
constitutes a member of the OSM deposits (Doppler 1989:
Table 1; Abdul Aziz et al. 2008: Fig. 2). The section is up
to 3 m thick and built up mainly by marls with pebbles (for
details see Fahlbusch et al. 1972, and below).
History of investigation
The fossil-rich 2–3-m-thick marl layer of Sandelzhausen
was discovered by Richard Dehm in 1959, during a
weekend family trip to a gravel pit, at that time still in use.
On the spot he found gastropod shells, turtle bones, and a
rhinocerotid jaw, as well as micromammals. Occasional
collecting in the following years and campaigns for
microfossil bulk sampling in 1962 and 1963 yielded sedi-
ment containing molluscs and micromammals well suited
for screen washing. Further investigations in 1964 found
that disarticulated remains of larger mammals were also
rather abundantly present and this discovery resulted in
several smaller digging campaigns in 1964–1968 and
finally in a larger long-term digging campaign from 1969
till 1975. During the latter campaign systematic excava-
tions based on a square meter grid (‘‘Planquadrate’’, PQ)
(Fig. 2h, i; see Fahlbusch et al. 1974; Fahlbusch 2003) and
distinct lithological horizons (see Fahlbusch and Gall 1970;
Fahlbusch et al. 1974; this paper below and Fig. 1) allowed
recording of the exact position within the fossil site for a
large portion of the recovered fossils and even the orien-
tation for the larger fossils. The sections derived during
these early years were fairly uniform with respect to sed-
imentology and distribution of fossil content, showing only
a small amount of lateral variation, though already a con-
centration of fossils in the eastern area of the site along a
NW–SE axis was noted.
After 1975, the remaining site was used agronomically
until 1994, when the City of Mainburg bought the ground
and offered the opportunity for further investigations.
During the second extensive field campaign period from
1994 to 2001 (Figs. 3and 4), the eastern natural border of
the fossil site was found and an area continuous with the
earlier observed NW–SE axis was found to be extraordi-
narily rich in larger mammal fossils (Fig. 4a–f).
Additionally, a core drilling program with the help of the
Bayerisches Geologisches Landesamt (Bavarian Geologi-
cal Survey) was initiated to find the distance to which the
fossil site extended northeast and east (Fig. 4h; see log
positions in Fig. 1). Westwards, the marly layer can be
traced for at least 1 km, but diminishes both in thickness
and grain size as well as in fossil richness. In 2002, the last
remaining excavation cavity of the fossil site was filled in
and is planned to be covered with buildings in the near
future. However, towards the north and west, the fossilif-
erous layer is still available, though covered by several
meters of younger deposits, and the area is still agronom-
ically used. The street Prof.-Dehm-Straße and two
information panels (Fig. 5c, d) for the public recall the
fossil site and its contribution to the knowledge of the
fauna and the palaeoenvironment 16 million years ago.
Lithology (Fig. 1)
The initial subdivision from 1966 of the marly horizon into
layers A–D, later supplemented by layers E–F (Fahlbusch
and Gall 1970; Fahlbusch et al. 1972,1974; Fahlbusch
2003), has been applied during all investigations of the site.
However, several marker horizons used initially proved
Fig. 2 Sandelzhausen field photos 1963–1975 (all photos by V.F.,
except where noted). aOccasional find of vertebrate remains during
an institutional trip in July 1963. Richard Dehm with his graduate
student Volker Fahlbusch. Photo by Herbert Hagn. bView of the
north side of the gravel pit in May 1964. Afterwards, the pit was
abandoned and partly refilled. The fossiliferous marl layer is situated
in the upper third of a gravel package termed ‘‘No
¨rdlicher Vollsch-
otter.’cFirst systematic investigation of the profile for its fossil
distribution and sedimentology in May 1966. Definition of the layers
A,B,C,Dwith sublayers B1,B2,C1,C2, and C3 (see Fahlbusch and
Gall 1970). dNearly complete vertical section with layers AE.
August 1972. eAn E-W section exposed at the end of the excavation
period in September 1973 showing continuously developed layers.
This is typical for most of the western part of the Sandelzhausen site,
but not for the eastern part excavated during the 1990s, where the
thickness of layers is more variable. fPreserving a vertical section by
gluing and transferring the sediment in situ onto fabric in August
1970. Several of these sections are housed in the BSPG and one is on
exhibit in the City Museum of Mainburg. gFirst complete skull find:
the rhinocerotid ‘‘Karl-Otto.’’ October 1970. Photo by M. Kindl. h
Systematic excavation in September 1970. The area is divided by a
grid of north-oriented square meters, each of which was worked on
one by one down to the base. Photo by Wolf-Dieter Grimm. i
Systematic excavation in August 1972. The upper parts of the marl
layer had been removed, as they were thought to be barren of
vertebrate remains at that time. jA frequently recurring problem
during all excavation periods was the complete watering of the
excavation site by strong rain events. August 1970
b
The fossil lagersta
¨tte Sandelzhausen (Miocene; southern Germany) 11
123
12 M. Moser et al.
123
either not to hold throughout all sections (e.g., the ‘‘coal
layer’’ and several sandy or pebbly layers) or to be other-
wise unreliable (pedogenetic carbonate nodules and
cements, diagenetic colouration, partial decarbonatization,
and minor cryoturbation). Detailed investigations of the
sedimentology and taphonomy were conducted by Fa-
hlbusch and Gall (1970); Herold and Ibrahim (1972);
Fahlbusch et al. (1972,1974), and finally Schmid (2002),
who also dealt with the pedogenesis. Additional details will
be published elsewhere (in preparation). The fossil-rich
layers B and C have been exploited consistently, while
higher parts of layer D and above could be examined only
during the second digging period (1994–2001).
Layer A (thickness 0–25 cm, usually 15 cm)
The basal layer A consists of marly, coarse gravels,
overall brownish, in some areas heavily cemented by ped-
ogenetic carbonates, resulting in a solid conglomerate which
rather hampers proper preservation. The rare fossil content
shows signs of transport and is mostly limited to robust
fragments such as proboscidean tusks and cheek teeth.
Layer B (thickness 40–120 cm, accelerating northeast-
wards)
The mostly greenish (in places yellowish) layer B (ini-
tially subdivided using various bands of pedogenetic
carbonate nodules and colour differences), consists roughly
of a fining upward sequence of gravel-rich marls, in which
both size and number of pebbles diminish upwards. In
detailed investigation, intercalated sand horizons and
repeated fining upward subsequences provide evidence for
nonuniform, discontinuous sedimentation. Layer B yielded
most of the macrovertebrate remains and is rich in car-
bonate nodules (*30%).
Layer C (thickness 0–40 cm)
The dark-brownish, almost black, organic rich layer
(‘‘coal layer’’ or C2) in the middle of the fossiliferous
horizon is the most easily recognizable marker (up to
*10 cm thick) in the section (e.g., Fig. 3g). Above (C3)
and below (C1), the sediment is coloured dark-brownish or
violet. This color pattern is present even in areas where the
dark organic-rich horizon is lacking. The dark organic-rich
layer consists of weakly coalified plant debris (recogniz-
able as wood fragments and twigs), abundant mollusc
shells (some even preserving color pattern), and the marly
sediment. Towards the marker horizon, gravel size dimin-
ishes to very fine gravel and coarse sand. Above the marker
bed, gravel content is present, but is greatly reduced.
Layer C is further characterized by the complete lack of
any pedogenetic carbonate nodules or cementations and by
the highest fossil concentration, though the fossil material
is strongly compacted.
Layer D (thickness *100 cm)
The light-greenish to grey marls (mainly silt) contain
few pebbles, but a significant proportion of diffuse car-
bonate and carbonate nodules (subhydrous pedogenesis)
especially in the lower part. Larger fossils often show signs
of reworking or long transport. However, though rare,
some of the best preserved specimens come from this
horizon due to a less intensive compaction.
Layer E (thickness *10 cm)
Dark-greenish silty clays. These were originally thought
to be free of fossils, but are now known to contain mi-
crovertebrates (Bo
¨hme 2009).
Layer F (thickness *30 cm)
The base consists of a few-centimetres-thick laminite
containing up to 17 bands with alternating light and dark
color (Bo
¨hme 2009 this volume), locally containing whit-
ish carbonate concretions and polygonal patterns of
desiccation cracks. Above this follows greenish–yellowish
clays. No fossils are found.
Fauna
Summarizing previous investigations from 1959 until
1969, including the first systematic digging season in 1969,
Fahlbusch and Gall (1970) already noted that Sandelz-
hausen ‘is the richest fossil vertebrate collecting site of
the OSM of Bavaria found in decades—if not at all’
(translated). The fossil list provided by Fahlbusch and Gall
(1970) contained, besides three plant forms, about 65 taxa
of animals including ostracods (*3 taxa), molluscs (*20),
and undifferentiated lower vertebrates (*7) as well as
mammals (*31). As scientific research progressed and
with additional material added during the following years,
the number of mollusc taxa grew to 51 species (Gall 1972)
Fig. 3 Sandelzhausen field photos 1994–1997 (all photos by M.M.).
aW. Werner, V. Fahlbusch, and R. Liebreich at the western digging
site (end of season 1994). bTypical arrangement of work: the
fossiliferous horizon was worked on with small picks and knives
while the cemented base of some areas had to be removed with a large
pick-axe. Fossils were referred to the field bureau (1995). cWorkers
of the City of Mainburg frequently removed the ever-growing
excavation debris (picture 1996). d,eRepeatedly, heavy rainfalls
flooded the pit with mud (picture 1996 with W. Werner and
K. Heissig). Therefore, the City of Mainburg built a drainage system
and kept it clean (picture 1996). fhIn 1996, during work for the new
street (later called Prof.-Dehm-Straße) fossils had to be picked at
great speed without position data. fEarly morning. gLate evening. h
A rhino skull in plaster package ready for transport (W. Werner,
K. Heissig, R. Liebreich). iScreen washing for microfossils was
undertaken usually once a week (picture 1997). kNew site east of the
newly built Prof.-Dehm-Straße in 1997
b
The fossil lagersta
¨tte Sandelzhausen (Miocene; southern Germany) 13
123
14 M. Moser et al.
123
and that for mammals to 57 species (Fahlbusch et al.
1974). Summarizing results of the first continuous digging
period (1969–1975), Fahlbusch (1976b) thus estimated the
fauna contained in the site to be clearly in excess of 100
species. After the second digging period (1994–2001) and
additional investigations by specialists, Fahlbusch (2003)
was able to list 185?taxa from Sandelzhausen, thus doc-
umenting the exceptional richness of this fossil lagersta
¨tte
(see Table 1for details). Regarding the future potential of
the site, it is noteworthy that even in most recent years
every digging season resulted in the addition of new taxa.
We thus conclude that further digging would still add
significantly to our knowledge of rare species, complete
our knowledge of the general morphology and variability
of known species, as well as augment our detailed
knowledge of the palaeoecology, environment, and genesis
of the site. A summary of the fossil content listing taxon
numbers and key references is presented in Table 1.
In comparison with other localities in the OSM, the so-
called lagersta
¨tten effect known from other fossil-rich
localities (Messel, Solnhofen, Burgess Shale, etc.) has
recently been recognized for Sandelzhausen: earlier
investigations indicated that the faunas older than *15 Ma
(including Sandelzhausen) seemed to be richer, especially
in the ecologically demanding ectothermic vertebrates,
than the immediately subsequent species-poor faunas. This
had been attributed to a catastrophic regional extinction
during the Ries impact event (Schleich 1985). However,
this pattern is now known to be the result of the smaller
fossil sample sizes recovered from all localities except
Sandelzhausen, partly amended in recent years (Heissig
1986;Bo
¨hme et al. 2002).
Age of the Sandelzhausen fossil site (Fig. 6)
In earlier publications, the age of Sandelzhausen had been
cited as Upper Miocene (e.g., Fahlbusch and Gall 1970;
Fahlbusch et al. 1974; Fahlbusch 1977). A shift in
Tethyan–Paratethyan marine correlations as well as conti-
nental–marine correlations (Fahlbusch 1976a,1981)
resulted in a new age estimate as old as the (middle)
Middle Miocene for Sandelzhausen (cited, e.g., by Schle-
ich 1983; Heissig 1989a,b). Finally, based on a change in
mammal biochronological correlation (Heissig 1997),
Sandelzhausen is now regarded as close to the Lower/
Middle Miocene boundary in age (see below).
Lithostratigraphy
The regional lithostratigraphical setting was recently rein-
vestigated by Abdul Aziz et al. (2008: Fig. 2). According
to regional mapping and lithostratigraphic correlations, the
Sandelzhausen section belongs to the upper part of the
No
¨rdlicher Vollschotter lithostratigraphic unit. This upper
part can be subdivided by the 7-m-thick Zwischenmergel
into two members and Sandelzhausen belongs to the lower
one. Within this up to 35-m-thick member, the fossil
locality is situated near the top and can thus lithostrati-
graphically correlated to other nearby fossil localities such
as Gisseltshausen 1b, Unterempfenbach 1b ?1d, Putten-
hausen E, and Maßendorf (the latter might be only slightly
older).
Biochronology
Initially, the age of the Sandelzhausen locality was bio-
stratigraphically dated as belonging to the regional
‘Mittlere Serie’’ of the OSM, a subdivision proposed using
proboscideans as index fossils (Dehm 1951,1955;
Fahlbusch and Gall 1970). According to Dehm (1955), the
‘Mittlere Serie’’ is thus characterized by the presence of
Gomphotherium angustidens and ‘Deinotherium’’ bavari-
cum; the lack in Sandelzhausen of the latter species was
initially attributed to the then incompletely known fossil
record of the site (Fahlbusch and Gall 1970). However,
Deinotherium (actually Prodeinotherium) fossils have
never been encountered in Sandelzhausen (Schmidt-Kittler
1972;Go
¨hlich 1999,2009), therefore the site must be
assigned to the ‘‘A
¨ltere Serie’’ of the Bavarian OSM, from
which Prodeinotherium, an African immigrant to Europe
during the Lower Miocene (Tassy 1989; Antunes 1989), is
unknown regionally, possibly for ecological reasons.
Fig. 4 Sandelzhausen field photos 1997–2001 (all photos by M.M.).
aThe fieldwork was conducted mostly with the help of volunteers,
some of whom were active for several years, such as F. Reinfelder,
N. Ballerstaedt, H. Hinle, and J. Herrlen (left to right, 1997). b
Rewarding finds: well-preserved molar of Gomphotherium subtapi-
roideum (1997). ceIn a line just a few meters off the eastern border
of the fossil site, a higher concentration of fossils was observed.
Sometimes, plaster near plaster had to be made, and the next fossils
were encountered even before the plasters could be removed (lower
jaws of a juvenile Gomphotherium in c, close-ups of later excavation
stages d,e; 1997). fR. Liebreich measuring length and position of a
Gomphotherium tusk. Inset: plaster package number 5,000. More than
6,000 larger fossils and plaster packages were recovered up to the end
of the campaign in 2001 (picture 1997). gOld site west of the Prof.-
Dehm-Straße, with well-exposed layers AD. The weakly coalified
‘coal-layer’’ is visible as dark band in the middle of the profile
(1997). hWith the mobile drilling equipment of the Bayerisches
Geologisches Landesamt (Bavarian Geological Survey, now part of
the Bayerisches Landesamt fu
¨r Umwelt) ten cores (e.g., inset figure)
were taken to find the extent of the fossiliferous layer towards the
north and east of the new digging site (January 1998). iFor the
University of Munich, 1 week each year was reserved for geology
students for an educational digging course. Twice, digging courses
were also arranged for students from Mainz and Vienna (this picture,
2000). kThe high layers EFare exposed here on the left (southern)
side. Selected material was put into sacks for screen washing (2001)
b
The fossil lagersta
¨tte Sandelzhausen (Miocene; southern Germany) 15
123
16 M. Moser et al.
123
The first investigations on mammals and gastropods
from Sandelzhausen (Fahlbusch 1964; Fahlbusch et al.
1974; Gall 1972) refined the age determination as being
contemporaneous either with Sansan (based on mammals,
Fahlbusch et al. 1974; Heissig 1989b), which later became
the reference locality for the European Neogene Land
Mammal Zone/Age/Reference-Fauna MN6 (Mein 1975,
1989,1999; Bruijn et al. 1992), or even Steinheim (‘‘Sil-
vana-Schichten’’, based on gastropods; Fahlbusch and Gall
1970; Gall 1972), the later reference locality for MN7
(Mein, op. cit.). Successive advances in the biochronology
of the ‘‘A
¨ltere’’ and ‘‘Mittlere Serie’’ of the OSM in
Bavaria (Heissig 1989a,b,1997;Bo
¨hme et al. 2002)
resulted in a regional subdivision of the OSM into bio-
chronologically discernible units intermediate between
MN4 and MN6, of which OSM C is represented by San-
delzhausen as standard fauna reference locality (Heissig
1997). Heissig correlated this unit to the middle (or even
early) MN5 and a middle position within the ‘‘A
¨ltere
Serie’’ of Dehm (1955), using the evolutionary stages of
the rodents Megacricetodon aff. bavaricus and Miodyro-
mys aegercii and the last occurrence of Anomalomys minor
(Heissig 1997,2005). Later, Abdul Aziz et al. (2008)
reinvestigated the local biozonation and concluded that the
units OSM C and OSM D are not distinctive enough and
should be unified into the unit OSM C ?D. Mein (2009
this volume) further corroborates the MN correlation by the
presence of the eomyid Keramidomys thaleri, which is
known from MN5 localities in France, Switzerland, and
Austria, and which is replaced by Keramidomys carpathicus
in MN6. In conclusion, considering all biochronological
evidence, the fauna of Sandelzhausen belongs to the early
middle MN5.
Absolute age and correlation to standard time scales
A correlation to the standard geological time scale is possible
by magnetostratigraphy and via cross-correlation of locali-
ties in France and Austria, which are intercalated into marine
strata dated biostratigraphically as Burdigalian (La Romieu:
MN4) and Langhian (Beaugency-Tavers, Pontlevoy-The-
nay: MN5) as well as upper Karpatian (Oberga
¨nserndorf,
Teiritzberg: old MN5) and lower Badenian (Mu
¨hlbach,
Grund: young MN5) (Fig. 5) (Heissig 1997; Daxner-Ho
¨ck
1998,2003;C
´oric
´et al. 2004; Harzhauser et al. 2002,2003).
Absolute ages for the OSM are derived from intercalated
bentonites found in Bavaria and Switzerland and from
debris related to the Ries impact event, which also forms a
regionally important marker horizon (‘‘Brockhorizont’’) in
Bavaria (Bolliger 1994; Heissig 1997;Bo
¨hme et al. 2002;
Abdul Aziz et al. 2008). In Bavaria, only the main bentonite
(late part of OSM F) is dated so far with an Ar–Ar age of
14.55 ±0.19 Ma, and new measurements of Ries glasses
(corresponding to the ‘Brockhorizont’, base of OSM F)
result in an age estimate of 14.88 ±0.11 Ma (Abdul Aziz
et al. 2008). However, according to magnetostratigraphical
results of sections containing the Brockhorizont, this age
may be—although within error—slightly too old (Abdul
Aziz et al. 2008). Combining both methods, the age of the
Ries impact can be estimated to 14.78 ±0.1 Ma. Thus, the
biochronostratigraphical units OSM E–F of Heissig (1997,
updated by Bo
¨hme et al. 2002), can be correlated indepen-
dently to the Langhian stage (Middle Miocene) (Fig. 6). For
the older units OSM A (MN4) and B -C?D (MN5), no
absolute ages are available yet from southern Germany, but
unpublished Ar-Ar data of a bentonite within the upper part
of OSM C ?D indicate an age close to the Burdigalian-
Langhian boundary (in preparation). Absolute ages for MN4
faunas obtained in France, Poland, and Hungary have been
recently published and summarized by Pa
´lfy et al. (2007),
resulting in age estimates for MN4 faunas ranging upwards
to at least 17.0 Ma. Thus OSM units B, C ?D are certainly
bracketed by the absolute dates of 17.0 Ma and 15.3 Ma.
In Spain, large terrestrial sections including faunas
belonging to the MN4 and MN5 zones have been repeatedly
dated by means of magnetostratigraphical correlation (e.g.,
Sen 1997; Daams et al. 1998,1999; Agustı
´et al. 2001;
Larrasoan
˜a et al. 2006; van Dam et al. 2006). The faunas,
however, are biochronologically only roughly correlatable
to the middle European faunas, due to the large geographical
distance and faunal differences. Because of this, the several
Fig. 5 The Sandelzhausen Meeting 2005 in Mainburg (all photos by
M.M.). aParticipants of the Sandelzhausen meeting: 1Gertrud
E. Ro
¨ssner (Munich), 2Volker Fahlbusch (Munich), 3Daria Petruso
(Palermo, Italy), 4Wilma Wessels (Utrecht, The Netherlands), 5
Ursula B. Go
¨hlich (Munich, now Vienna, Austria), 6Chiara Ange-
lone (Roma, Italy), 7Margery C. Coombs (Amherst, Massachusetts,
USA), 8Josef Egger (Mayor of Mainburg), 9Henriette Jechorek
(Go
¨rlitz), 10 Thorsten Kowalke (Munich), 11 Dietmar Jung (Munich),
12 Reinhard Ziegler (Stuttgart), 13 Gerhard Doppler (Munich), 14
Wolfgang Witt (Gu
¨ndlkofen), 15 Pierre Mein (Lyon), 16 John
Damuth (Santa Barbara, CA, USA), 17 Norbert Schmidt-Kittler
(Mainz), 18 Jan van der Made (Madrid, Spain), 19 Ralph Annau
(Munich), and 20 Kurt Heissig (Munich). Participants not shown in
the picture: Madelaine Bo
¨hme (Munich), Hans Hinle (Kelheim),
Thomas Kaiser (Hamburg), Markus Moser (Munich), Doris Nagel
(Vienna), Jerome Prieto (Munich), Bettina Reichenbacher (Munich),
Zbigniew Szyndlar (Krakow), and Clara Stefen (Dresden). bDuring
session in the City Hall of Mainburg. cOpening ceremony of the
information panels at the filled-in Sandelzhausen pit. dReconstruc-
tion of the landscape of Sandelzhausen 16 million years ago by artist
Wenzel Balat. Shown are the rhinos Plesiaceratherium (foreground)
and Lartetotherium (background), and the chalicothere Metaschizo-
therium (right, with hind legs that are too short; M. Coombs, oral
communication, at symposium 2005). The painting is also shown on
the left panel in Fig. 4c. eVolker Fahlbusch with Josef Egger, first
Mayor of Mainburg and generous supporter of the field campaigns of
1994–2001 and the meeting in 2005, with his Sandelzhausen digging
shirt (cartoon painting by Bettina Schenk, formerly Munich, now
Vienna)
b
The fossil lagersta
¨tte Sandelzhausen (Miocene; southern Germany) 17
123
attempts to correlate terrestrial sections on a European scale
with the continuous marine magnetostratigraphic scales are
not conflict-free among themselves or compared with other
correlation methods (see refs. above; for example, the Ibe-
rian faunas from Vargas and Fuente Sierra are attributed to
MN4 and dated as young as 16.0 Ma, thus clearly postdating
several MN5 localities in western and middle Europe).
Most recently, Abdul Aziz et al. (2008) provided
magnetostratigraphic data for Sandelzhausen. Within the
section, they identified an inverse to normal reversal event
and correlate it with the reversal between magnetochron
C5Cn.2r-2n of the Astronomically Tuned Neogene Time
Scale (ATNTS04), corresponding to an absolute age around
16.47 Ma (Lourens et al. 2004, p. 470). However, a corre-
lation to C5Cn.1r-1n (16.27 Ma) is possible too, and
appears to be not unlikely, considering that the classical
fauna of Puttenhausen (close to Sandelzhausen) (Wu 1982;
Heissig 1997; Abdul Aziz et al. 2008) is biochronologically
and lithostratigraphically slightly older than Sandelzhausen
(both belong to the late part of OSM C ?D), but cannot be
older magnetostratigraphically than C5Cn.2r, as the fauna
of Puttenhausen is biochronologically younger than that of
Oberga
¨nserndorf (basal MN5; Austria; Daxner-Ho
¨ck 1998),
which belongs to magnetochron C5Cn.3n and is bracketed
well by marine biostratigraphy (Steininger 1999; Harzha-
user et al. 2002).
Currently, no GSSP for the base of the Langhian stage
(base of the Middle Miocene) has been defined, and dif-
fering concepts have been applied; traditionally, the base of
the Langhian either is defined at the first appearance datum
(FAD) of the planktonic foraminifer Praeorbulina glome-
rosa, which is almost coincident with the beginning of
magnetochron C5Cn.1n at 16.27 Ma, or at the FAD of
Praeorbulina sicana, which in Mediterranean areas is
hiatus
A
Obergänserndorf
early MN5, Karpatian
C5Cn.3n, 16.5-16.7 Ma
l
anoiti
d
a
r
t
()MB
S
OSM E'
OSM E
OSM
C+D
OSM B
OSM A
Sandelz-
hausen
ei
reSerel
t
t
iM
eireSeretlÄ
Ries
impact
14.88 Ma
MN6
MN5
MN4
Sansan
type MN6
Pontlevoy-Thenay
type MN5, Langhian
La Romieu
type MN4, Burdigalian
main
bentonite
14.55 Ma
Martinsbrünneli
Frohberg
bentonite 14.6 Ma
bentonite 15.3 Ma
FCHD
bentonite 14.2 Ma
Rümikon
biochronological
correlation
marine
biostratigraphical
bracket
-arreS n
a
i
l
l
av
n
aihgnaL
)
e
n
e
c
oiM
e
ld
di
M
(
na
i
lag
idruB
)
e
n
ecoi
M
r
ew
o
L(
13.65 Ma
magnetostratigraphical
correlation
Beaugency-Tavers
Langhian
OSM F
Mühlbach
late MN5, Badenian
15.1 Ma
boundary of
C5Cn.1n-1r
16.27 Ma
boundary of
C5Cn.2n-2r
16.47 Ma
FAD
16.27 Ma
Praeorbulina
glomerosa
??
??
top of
C5Cn.1n
15.97 Ma
hiatus
Fig. 6 Age of the Sandelzhausen fossil lagersta
¨tte. A framework of
biochronological, biostratigraphical, and magnetostratigraphical
cross-correlations with localities in France (F), Switzerland (CH),
and Austria (A) allows dating of the localities of the Upper Freshwater
Molasse (OSM) in Bavaria. Subdivision of OSM into series, and
faunal units after Dehm (1951) and Heissig (1997), updated by
Bo
¨hme et al. (2002) and Abdul Aziz et al. (2008). Absolute ages of
bentonites in Switzerland after Bolliger (1994), new ages of main
bentonite and Ries impact glasses in Bavaria after Abdul Aziz et al.
(2008). Absolute dates for Austrian localities as reported by Daxner-
Ho
¨ck (2003) and Harzhauser et al. (2002,2003). Absolute dates for
the boundaries of the global marine stages (right column) after
Lourens et al. (2004), the stippled lines indicate the provisional status
of the proposed boundaries (see text for details). Attribution of faunas
to Mammal Neogen divisions (MN) after Heissig (1997) and Daxner-
Ho
¨ck (1998,2003). Sandelzhausen belongs to (early) middle MN5
and can be correlated to either magnetochronological reversals
C5Cn.1r-1n or C5Cn.2r-2n and to either the topmost Burdigalian
(Lower Miocene) or the lowermost Langhian (Middle Miocene)
stages of the global time scale. Accordingly, the absolute age can be
given as 16.47 or 16.27 Ma
18 M. Moser et al.
123
about 16.3 or 16.4 Ma in age (within magnetochron
C5Cn.2n), but dated as old as 16.97 Ma (amidst mag-
netochron C5Cr) in Atlantic cores (Berggren et al. 1995;
Sprovieri et al. 2002;C
´oric
´et al. 2004; Lourens et al.
2004; Kova
´c
ˇet al. 2007). However, giving these uncer-
tainties, Lourens et al. (2004) arbitrarily suggested setting
the boundary at the top of C5Cn.1n at 15.97 Ma with no
biostratigraphical background. Until this issue is settled
(the ICS has scheduled a golden spike for 2008), it is
therefore not possible to state whether Sandelzhausen is
clearly Lower or Middle Miocene in age, but it is certainly
very close to the boundary of these subseries. It may be
noted here that several reptile taxa from Sandelzhausen
such as Eoanilius (Szyndlar 2009 this volume), Palaeobl-
anus,Palaeocordylus, and Amblyolacerta (Bo
¨hme 2009),
are typical for Late Oligocene and Early Miocene localities
elsewhere.
In summary (Fig. 6), Sandelzhausen magnetostrati-
graphically correlates to either the C5Cn.1r-1n or the 2r-2n
reversal event and biochronologically with either the top-
most Burdigalian (Lower Miocene) or the basalmost
Langhian (Middle Miocene) stages, corresponding in
absolute ages to somewhat more than about 16 Ma.
Acknowledgments The investigation of the Sandelzhausen fossil
site has been enthusiastically supported for over 40 years by a large
number of individuals: students, scientists, neighbors and property
owners, volunteers, and sponsors, among whom first we would like to
name in most deserved gratitude our colleagues Winfried Werner and
Renate Liebreich; then, Kurt Heissig, Norbert Schmidt-Kittler, as well
as the volunteers Hans Hinle, Friedrich Reinfelder, Jo
¨rg Herrlen, and
Dieter Dernbach for their long-lasting contribution to the fieldwork.
We are indebted to the City of Mainburg (Mayor Josef Egger, the City
Council, and co-workers), which provided its property for excavation,
water, power, and technical help on numerous occasions without
which the exploitation of the fossil site would not have been possible.
The City also generously hosted and organized the Sandelzhausen
Symposium 2005. Financial support for the first digging period
(1969–1975) and the Sandelzhausen symposium 2005 in Mainburg
came from the Deutsche Forschungs-Gemeinschaft (DFG, GZ 4850/
88/05). The second digging period (1994–2001) was made possible
almost exclusively by generous sponsors: the Freunde der Bayeris-
chen Staatssammlung fu
¨r Pala
¨ontologie und historische Geologie
(Mu
¨nchen), the Bayerische Akademie der Wissenschaften, the com-
panies Wolf Klimatechnik (Mainburg), Deutsche Gleis- und Tiefbau
(Berlin), Pinsker Druck und Medien (Mainburg), Bosch-Siemens-
Haushaltsgera
¨te (Mu
¨nchen), the financial institutes Raiffeisenbank
Hallertau/Mainburg, Sparkasse Kelheim and Stadtsparkasse Mu
¨n-
chen, and many others. Additionally, technical and material support
came from the companies Thyssen-RO
¨RO (Mu
¨nchen), Ziegelwerke
Leipfinger und Bader (Puttenhausen), Volkswagen (Abteilung Nutz-
fahrzeuge, Mu
¨nchen), Elektrofirma Burger (Mainburg), and the Isar-
Amper-Werke (now Eon, Mu
¨nchen). Last, but not least, we would
like to thank the neighbors of the digging site for their generous
hospitality towards our crews during both digging periods. We cor-
dially thank Renate Liebreich and Winfried Werner (both Munich)
who kindly reviewed an earlier draft of this work, Gudrun Ho
¨ck and
Mathias Harzhauser (both Vienna) for their valuable improvements,
and John Damuth (Santa Barbara, CA, USA) for the improvement of
the English.
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The fossil lagersta
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... The Sandelzhausen fossil site is one of the most important Cenozoic continental sites in Europe (MoSeR et al. 2009a) and its rich record includes hundreds of gastropods. Still, only two works dealt specifically with the mollusks: gall (1972), who identified 49 gastropods and two bivalves in the material recovered, but based his work heavily on younger faunas; and MoSeR et al. (2009b), who dealt with paleoecological questions. ...
... Still, only two works dealt specifically with the mollusks: gall (1972), who identified 49 gastropods and two bivalves in the material recovered, but based his work heavily on younger faunas; and MoSeR et al. (2009b), who dealt with paleoecological questions. Here is presented the third and last part of the taxonomic treatment of the pulmonate snails from Sandelzhausen (for which harbors the stratigraphical group known as Upper Freshwater Molasse (Ober Süßwassermolasse, OSM; MoSeR et al. 2009a). The fossils from Sandelzhausen belong to a member of the OSM called Nördlicher Vollschotter, composed primarily of marl and gravel (MoSeR et al. 2009a). ...
... Here is presented the third and last part of the taxonomic treatment of the pulmonate snails from Sandelzhausen (for which harbors the stratigraphical group known as Upper Freshwater Molasse (Ober Süßwassermolasse, OSM; MoSeR et al. 2009a). The fossils from Sandelzhausen belong to a member of the OSM called Nördlicher Vollschotter, composed primarily of marl and gravel (MoSeR et al. 2009a). The age of these deposits was established by stratigraphic, biostratigraphic and magnetostratigraphic correlations: the Middle Miocene Burdigalian/Langhian boundary (~16.47-16.27 ...
... In the present study, we took advantage of the Miocene fossil lagerstätte Sandelzhausen in Germany, which has revealed an exceptional high percentage of remains of juvenile individuals for many species (Fahlbusch 1976(Fahlbusch , 2003Moser et al. 2009). In particular, the rhinoceros Prosantorhinus germanicus is remarkable for an extraordinary high number of juvenile dentaries at different developmental stages including highly fragile tooth germs (Fahlbusch et al. 1972;Peter 2002). ...
... Although Sandelzhausen mainly yielded disarticulated vertebrate skeletons, the preservation of fragile fossils and the absence of transport marks on most skeletal remains indicate that transport over long distances is unlikely (Fahlbusch et al. 1972). The sediments of the Upper Freshwater Molasse in which the fossils occur represent a limnofluvial paleoenvironment with seasonal inundations (Fahlbusch et al. 1972;Schmid 2002;Moser et al. 2009;Böhme 2010). The mammal remains were washed in from the moist subtropical woodland habitat and deposited within a relatively short time (Fahlbusch et al. 1972 and references therein;Fahlbusch 2003;Moser et al. 2009). ...
... The sediments of the Upper Freshwater Molasse in which the fossils occur represent a limnofluvial paleoenvironment with seasonal inundations (Fahlbusch et al. 1972;Schmid 2002;Moser et al. 2009;Böhme 2010). The mammal remains were washed in from the moist subtropical woodland habitat and deposited within a relatively short time (Fahlbusch et al. 1972 and references therein;Fahlbusch 2003;Moser et al. 2009). There is evidence for accumulation of the fossils through deposition of episodic influx within a fluvial plain (Fahlbusch et al. 1972;Fig. ...
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Fossil evidence of complete sequences of dental ontogeny in extinct mammals is rare but contains valuable information on the animal’s physiology, life history, and individual age. Here, we analyzed an exceptionally high number of juvenile dentaries at different developmental stages including highly fragile tooth germs of the extinct rhinoceros Prosantorhinus germanicus from the Miocene fossil lagerstätte Sandelzhausen in Germany. We used dental wear stages, eruption stages, and tooth germ development in order to reconstruct the tooth replacement pattern for P. germanicus. The results allow for the distinction of 11 dental eruption stages and document a tooth eruption sequence of (d2, d3), (d1, d4), m1, m2, p2, p3, p4, m3; a pattern identical to that reported for the extant African rhinoceros, Diceros bicornis. Moreover, our findings indicate that P. germanicus falls into the life history category of slow-growing, long-living mammals. The dental eruption stages of the fossil rhinoceros were correlated with data of living rhinoceroses in order to gain insight into the age-at-death distribution of P. germanicus at Sandelzhausen. The juvenile mortality profile of P. germanicus shows a trend of selective mortality at an inferred age range of about 3 months to 3 years. As this age range represents a life phase of increased natural risk of mortality, our findings indicate a gradual accumulation of corpses (attritional fossil assemblage). This result supports the interpretation of a taphocenosis found at the Sandelzhausen fossil site.
... The Sandelzhausen fossil site is one of the most important Cenozoic continental sites in Europe (MoSeR et al. 2009a) and its rich record includes hundreds of gastropods. Still, only two works dealt specifically with the mollusks: gall (1972), who identified 49 gastropods and two bivalves in the material recovered, but based his work heavily on younger faunas; and MoSeR et al. (2009b), who dealt with paleoecological questions. ...
... Still, only two works dealt specifically with the mollusks: gall (1972), who identified 49 gastropods and two bivalves in the material recovered, but based his work heavily on younger faunas; and MoSeR et al. (2009b), who dealt with paleoecological questions. Here is presented the third and last part of the taxonomic treatment of the pulmonate snails from Sandelzhausen (for which harbors the stratigraphical group known as Upper Freshwater Molasse (Ober Süßwassermolasse, OSM; MoSeR et al. 2009a). The fossils from Sandelzhausen belong to a member of the OSM called Nördlicher Vollschotter, composed primarily of marl and gravel (MoSeR et al. 2009a). ...
... Here is presented the third and last part of the taxonomic treatment of the pulmonate snails from Sandelzhausen (for which harbors the stratigraphical group known as Upper Freshwater Molasse (Ober Süßwassermolasse, OSM; MoSeR et al. 2009a). The fossils from Sandelzhausen belong to a member of the OSM called Nördlicher Vollschotter, composed primarily of marl and gravel (MoSeR et al. 2009a). The age of these deposits was established by stratigraphic, biostratigraphic and magnetostratigraphic correlations: the Middle Miocene Burdigalian/Langhian boundary (~16.47-16.27 ...
Article
Full-text available
Sandelzhausen is an Early/Middle Miocene (Mammal Neogene zone MN5) fossil site near Mainburg, southern Germany, home to a very rich fossil record. Hundreds of fossil continental mollusks, almost exclusively pulmonates snails, were recovered during the excavations, but received scarce attention by researchers. Here is presented the third and last part of a taxonomical treatment of the fossil pulmonates from Sandelzhausen, dealing with the superfamilies Lymnaeoidea, Planorboidea, Punctoidea, Gastrodontoidea and Limacoidea. The following species were found in the material: Galba dupuyiana, Lymnaea dilatata and Radix socialis (Lymnaeidae); Ferrissia deperdita, Gyraulus albertanus, Gyraulus dealbatus, Hippeutis sp., Planorbarius cornu and Segmentina lartetii (Planorbidae); Discus pleuradrus (Discidae); Lucilla subteres (Helicodiscidae); Janulus supracostatus (Gastrodontidae); Archaeozonites sp. (Gastrodontoidea incertae sedis: Archaeozonitinae); Limax sp. (Limacidae); Vitrina sp. (Vitrinidae).
... The Miocene Sandelzhausen Fossil-Lagerstätte with an absolute age of somewhat more than 16 Ma (MN5) is located near Mainburg, 60 km north of Munich (Bavaria, Germany; Moser et al. 2009). The locality was discovered in 1959 (Fahlbusch and Gall 1970) and several digging campaigns yielded remains of more than 120 vertebrate taxa (Fahlbusch 2003;Moser et al. 2009). ...
... The Miocene Sandelzhausen Fossil-Lagerstätte with an absolute age of somewhat more than 16 Ma (MN5) is located near Mainburg, 60 km north of Munich (Bavaria, Germany; Moser et al. 2009). The locality was discovered in 1959 (Fahlbusch and Gall 1970) and several digging campaigns yielded remains of more than 120 vertebrate taxa (Fahlbusch 2003;Moser et al. 2009). The remains of three rhinoceros species are the most abundant large mammal findings in the Sandelzhausen locality (Heissig 1972;Fahlbusch et al. 1974). ...
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Prosantorhinus germanicus is a small, short-legged, teleoceratine rhino from the Miocene of Sandelzhausen (Bavaria, Germany). P. germanicus shows a high variation in some of its carpal bones. A unique modification of the articulation of Intermedium and Carpale 4 is described here. Special emphasis is given to additional articulation facets at the palmar processes of both bones. These additional contacts, working as stop facets, are unique among rhinos and restrict the flexion of the mid-carpal joint. Some individuals show these additional facets which prohibit the flexion within the wrist and therefore stiffen the carpus. Carpale 4 specimens without the additional facets show knob-like structures instead. These knobs are most likely precursory structures of those facets and the facets are fully developed in heavier males. A skeletal sexual dimorphism is not visible in the sample as all bones are in the same size range. The wrist stiffening in the mid-carpal joint supports a greater bodyweight and therefore could coincide with P. germanicus as a proposed dwarfed rhinoceros species. The stiffening can also be interpreted in favor of a semiaquatic mode of life. The stiffened carpus is more resistant against injuries while walking on muddy grounds in a wet environment.
... Christine Böhmer*, Kurt Heissig** und Gertrud E. Rössner** Die miozäne Fossilfundstelle Sandelzhausen ist berühmt für ihre Säugetiere und hat seit Bestehen des Vereins einen besonderen Stellenwert im Hinblick auf Fördermaßnahmen. Bergung, Erhaltung und Auswertung der Fossilien haben maßgeblich davon profitiert (siehe Fahlbusch 1995;Fahlbusch et al. 1997Fahlbusch & Liebreich 1999siehe außerdem Fahlbusch 1976siehe außerdem Fahlbusch , 2003Moser et al. 2009) und wurden zu einer der wichtigsten Informationsressourcen für den Übergangsbereich von Früh-zu Mittel-Miozän in Eurasien. Alle Sandelzhausen-Fossilien werden an der Bayerischen Staatssammlung für Paläontologie und Geologie unter der Sammlungsnummer 1959 II aufbewahrt. ...
... Wie schon lange bekannt, repräsentieren die Sandelzhausener Sedimente Teile der sogenannten Oberen Süßwassermolasse, deren Ablagerungsraum eine vielfältige Flusslandschaft war (Fahlbusch et al. 1972;Schmid 2002;Fahlbusch 2003;Rössner & Nose 2008;Moser et al. 2009;Böhme 2010). Es gibt Hinweise darauf, dass sich die außerordentliche Fossiliendichte in Sandelzhausen durch saisonale Überschwemmungen angehäuft hat, aber es besteht Uneinigkeit über die Anreicherungsdauer im Rahmen Abb. ...
... pantherae, Vertigo angulifera and Vitrea ammoni) do not provide much information with respect to the paleoecological analysis. In contrast, species occurring only in Sandelzhausen, indicate a much more eschweizerbart_xxx diverse freshwater environment (see also moser et al. 2009b;sAlvAdor & rAsser 2014). ...
Article
Full-text available
This study presents a taxonomic study and paleoecological reconstructions of the continental gastropod assemblages from two new Miocene localities in the German part of the North Alpine Foreland Basin, Bavaria. Riedensheim (next to the city of Rennertshofen) and Fasanerie (next to the city of Adelschlag). The fossil rich deposits belong to the Upper Freshwater Molasse (OSM), correlated to the regional biostratigraphic unit C+D (Burdigalian/Langhian; MN 5 in the European mammal Neogene zone). Seventeen gastropod species (almost exclusively pulmonates) were found in Adelschlag-Fasanerie and 35 species in Riedensheim. Thirteen species are shared by both assemblages. Of these, the following species have their distribution expanded in Germany: Azeca peneckei, Carychium (Carychium) galli, Vitrea ammoni and Urticicola perchtae; and possibly also Stagnicola cf. praebouiletti and Truncatellina cf. pantherae. Paleoecological reconstructions indicate a rich freshwater gastropod community in Riedensheim, with a well-vegetated area immediately surrounding the water body (which diminished in size with time), and a sourrounding environment dominated by humid forest. The fauna from Adelschlag-Fasanerie, which was much impoverished in comparison, might represent flood plain and fluvial channel deposits. Finally, a comparison to the mollsucan faunas of coeval and nearby localities (Adelschlag, Attenfeld, Sandelzhausen and Undorf) is provided.
... The Desmaninae ended their long absence with the return of Mygalea jaegeri (Seemann, 1938). On the other hand, the locality Sandelzhausen is no fissure filling, but consists of Molasse sediments, including freshwater deposits (Moser et al. 2009). Therefore, the desmans' absence in the Southern German localities might just be the result of the absence of freshwater deposits during MN 3 and 4. The Scalopini continued to be present. ...
Article
Full-text available
In the Miocene, the talpid diversity was much higher than today. Especially the semifossorial and ambulatory moles were farther distributed and more common. Here, the talpids from the Bavarian fissure filling Petersbuch 28 (Early Miocene, MN 3) are described. The uropsiline Desmanella engesseri Ziegler, 1985 is the most frequent species, followed by Myxomygale hutchisoni (Ziegler, 1985), an urotrichine less specialized towards burrowing. The fossorial talpids Talpa sp. and Proscapanus intercedens Ziegler, 1985 are represented by humeri, P. intercedens by a few teeth. Some teeth could not be determined to species level. Among the rich postcranial material, the humerus of Desmanella engesseri was found for the first time. Compared to other Miocene localities, Petersbuch 28 is unique in the dominance of only superficially burrowing and semifossorial talpids. In the time from MN 2 to MN 7 + 8 (Early to Middle Miocene), the fossorial talpids were less dominant during MN 2 and most of MN 3, became more frequent during MN 4 and were predominant from MN 5 onward.
... This species was initially represented only by the teeth (a right m3, holotype) (Roger, 1896). Addi-tional specimens of Lagomeryx pumilio teeth were found especially from the fossil site Sandelzhausen, southern Germany, in a layer correlated with the biozone MN5 (Moser et al., 2009). The holotype, a right m3, displays an enlarged entoconulid forming a third lingual cusp (Roger, 1896). ...
Article
The primitive deer (subfamily Lagomerycinae) Lagomeryx and Stephanocemas are characterized primarily by their palmate antlers. Two lagomerycines, Lagomeryx manai, sp. nov., and Stephanocemas rucha, are described for the first time from Q and K coal layers of the late middle Miocene (13.4–13.2 Ma) Mae Moh Basin in northern Thailand. A species-level phylogeny of the Ligeromeryx-Lagomeryx clade, based on cranial appendages, reconstructs Lagomeryx manai, n. sp., as a derived species of Lagomeryx, sister group of Lagomeryx complicidens. This study suggests that the large species of Lagomeryx are restricted geographically to Asia and dispersed to Southeast Asia at the latest during late middle Miocene, where they are represented by Lagomeryx manai, n. sp. The paleoenvironmental studies of five Mae Moh mammalian taxa, a cervid (Lagomeryx manai, n. sp.), an indeterminate bovid, a suid (Conohyus thailandicus), a rhinoceros (Gaindatherium sp.), and a proboscidean (Stegolophodon sp.), investigated with stable carbon and oxygen isotope analyses of tooth enamel, indicate that the Mae Moh mammals inhabited a wide range of habitats from woodlands to grasslands in a C3-plant-dominated environment. The new species of Lagomeryx seems to have been living in an open environment, contrary to its European relatives. The serial isotopic samples also support that Mae Moh herbivores probably lived in a low-seasonal climate during the late middle Miocene of northern Thailand.
Article
The fossil molluscan fauna of Sandelzhausen (Early/Middle Miocene, SE Germany) comprises a total of 44 species, mostly terrestrial pulmonate snails. Herein we present a paleoecological analysis of this fauna based on an actualistic approach and on data on stable isotopes of carbon and oxygen (assessed from specimens of the freshwater lymnaeid snail Galba dupuyiana and the terrestrial clausiliid snail Pseudidyla moersingensis). The paleoecological reconstruction achieved here is in line with previous works, with some novelties and minor modifications. The basal sediment layers point to a swampy area with ponds and/or oxbow lakes (closed system, as indicated by the covariation between oxygen and carbon isotopic signals of G. dupuyiana), prone to seasonal flooding events. This environment would then gradually transition into a perennial lake, as indicated by: the proportion of planorbids, the appearance of aquatic species intolerant to desiccation, and the decoupling of the covariation between oxygen and carbon isotopic signals of G. dupuyiana. The terrestrial habitat would have developed from a more open environment (semi-arid/sub-humid scrubland) to a sub-humid/humid denser forest afterwards. Still, species from drier and more open environments are present throughout all the layers, suggesting that these habitats persisted in the lake’s hinterland. The mean annual temperature, calculated from the oxygen isotopic composition of P. moersingensis, ranges from 18.5 to 20.5 °C, but with no significant trend of change throughout the layers.
Article
Blanquatere 1 is the filling of a fissure opened in a limestone that had already provided a rich fauna (more than 6300 isolated teeth, 33 species) of small mammals including mainly rodents among which several new species of Cricetidae and Gliridae, referred to the Middle Miocene. The present paper brings an addition to this fauna with the description of the Cricetidae Eumyarion, Democricetodon, and Pseudofahlbuschia, of the Eomyidae Ligerimys et Keramidomys, and of the Sciuridae Spermophilinus and Heteroxerus. This highly diverse fauna must be definitely considered as a reference for the first part of the Middle Miocene (MN 4/5) in South Western Europe. It testifies of the occurrence of both new taxa and allochtonous ones, as the gerbillid Dakkamys sp. the occurrence of which being likely related to the closure of the Tethys in the Middle East during the Langhian. The co-occurrence of Ligerimys and Keramidomys is also documented as the oldest one. This fauna also illustrates the interest of karstic fissure fillings when they are exceptionally rich: they can deliver remains of species either very rare or of short duration. © Publications Scientifiques du Muséum national d'Histoire naturelle, Paris.
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The first record of an isolated distal phalanx of a fossil pangolin (Pholidota) is reported from the rich vertebrate locality of Sandelzhausen (Upper Freshwater Molasse, Bavaria, MN 5). The somewhat fragmentary claw is similar in size and morphology to specimens from Wintershof-West (MN 3) and Vieux Collonges (MN 5). It is to be determined as Necromanis sp. Whether it belongs to hand or foot is unknown as well as the number of the finger/toe. It is the youngest record of Pholidota from the Miocene of Southern Germany.
Article
When in 1984 an intensive exploration of the exposures of the Upper Freshwater Molasse started, there existed two biostratigraphical scales more or less independent from one another. The first was the subdivision by Dehm (1951, 1955) in three units on the basis of the occurrence of different proboscidean species, the “Ältere Serie” with only the small Gomphotherium angustidens (Cuvier), the “Mittlere Serie” with the presence of both G. angustidens and Deinotherium bavaricum v. Meyer, and the “Jüngere Serie” or“Hangendserie” with big species of both genera. Several local lithostratigraphical units have been assigned to these only paleontologically defined “series,” namely in the east of the Molasse basin.