REVIEW ARTICLE
Vertebrate tracksites in the Obernkirchen Sandstone
(late Berriasian, Early Cretaceous) of northwest
Germany— their stratigraphical, palaeogeographical,
palaeoecological, and historical context
Jahn J. Hornung • Annina Bo¨hme •
Torsten van der Lubbe • Mike Reich •
Annette Richter
Received: 26 November 2010 / Accepted: 25 October 2011 / Published online: 14 March 2012
 The Author(s) 2012. This article is published with open access at Springerlink.com
Abstract The northern German Lower Cretaceous
Bu¨ckeberg Formation yields numerous dinosaur tracksites,
some of which have produced material of impressive qual-
ity. Stratigraphically, the localities are concentrated in the
Obernkirchen Sandstone, a thin subunit within this forma-
tion. The Obernkirchen Sandstone represents mainly a
sandy barrier to back-barrier and lagoonal setting within a
limnic deltaic facies complex, which was deposited during
the late Berriasian (Cypridea alta formosa ostracod sub-
zone) in the southeast of the Lower Saxony Basin, northwest
Germany. A few tracksites occur more proximally in coeval
fluvial deposits. Dinosaur footprint assemblages were left by
ornithopods, theropods, sauropods, ankylosaurs, and small,
bipedal ornithischians. Other vertebrate tracks are those of
turtles and, possibly, crocodilians. Due to the decrease in
sandstone quarrying in recent decades, many old tracksites
are inaccessible today. Additionally, historical descriptions
of the tracks were of highly variable quality and often
published in remote and today nearly unobtainable sources.
Here we provide a catalogue of 13 tracksites compiled from
the literature and some new observations. Of these 13
tracksites, only five are still accessible and currently under
study. Descriptions of each locality are provided, with a
comprehensive compilation of existing data on lithofacies,
stratigraphy, palaeogeography and palaeoecology of the
Obernkirchen Sandstone and equivalent strata. A short
review of the track-bearing lithofacies assemblage indicates
that the outcrop areas have distinctly different facies and
environments, and, therefore, track-bearing horizons can
only be correlated stratigraphically between adjacent out-
crops. For this reason, the identification of a megatracksite
in the Obernkirchen Sandstone is currently regarded as
premature and uncertain.
Keywords Vertebrate tracks  Dinosauria  Cretaceous 
Berriasian  Obernkirchen  Mu¨nchehagen  Germany
Kurzfassung Die unterkretazische Bu¨ckeberg-Formation
Nordwest-Deutschlands ist reich an Fundstellen von Di-
nosaurierfa¨hrten, von denen einige hervorragend erhaltenes
Material lieferten. Diese Fundstellen sind stratigraphisch
auf ein geringma¨chtiges Intervall, den Obernkirchen-
Sandstein, konzentriert. Der Obernkirchen-Sandstein
repra¨sentiert eine sandige Barrieren- und Lagunen-Fazies
innerhalb eines limnisch-deltaischen Fazieskomplexes, der
wa¨hrend des spa¨ten Berriasiums (Cypridea alta formosa
Ostrakoden-Subzone) im Su¨dosten des Niedersa¨chsischen
Beckens abgelagert wurde. Einige Fa¨hrtenfundstellen lie-
gen auch in etwas proximaleren, gleichaltrigen, fluviatilen
Ablagerungen. Erzeuger der Dinosaurierfa¨hrten waren
Ornithopoden, Theropoden, Sauropoden, Ankylosaurier
J. J. Hornung (&)  A. Bo¨hme
Geowissenschaftliches Zentrum, Abt. Geobiologie,
Georg-August-Universita¨t Go¨ttingen, Goldschmidtstr. 3,
37077 Go¨ttingen, Germany
e-mail: jhornun@gwdg.de
A. Bo¨hme
e-mail: aboehme@gwdg.de
T. van der Lubbe  A. Richter
Niedersa¨chsisches Landesmuseum Hannover,
Willy-Brandt-Allee 5, 30169 Hannover, Germany
e-mail: DragonsClaw@gmx.net
A. Richter
e-mail: Annette.Richter@nlm-h.niedersachsen.de
M. Reich (&)
Geowissenschaftliches Zentrum, Museum, Sammlungen
and Geopark, Georg-August-Universita¨t Go¨ttingen,
Goldschmidtstr. 1-5, 37077 Go¨ttingen, Germany
e-mail: mreich@gwdg.de
123
Pala¨ontol Z (2012) 86:231–267
DOI 10.1007/s12542-012-0131-7
sowie kleine, bipede Ornithischia. Weitere Wirbeltier-
Fa¨hrten wurden von Schildkro¨ten und mo¨glicherweise
Krokodilen hinterlassen. Aufgrund des Ru¨ckgangs im
Sandstein-Abbau im Verlauf der vergangenen Jahrzehnte
sind viele der alten Fa¨hrten-Fundstellen heute nicht mehr
aufgeschlossen. Historische Beschreibungen sind von un-
terschiedlicher Qualita¨t und ha¨ufig nur in schwer erha¨ltli-
chen Quellen publiziert. Hier pra¨sentieren wir einen
Katalog mit 13 Fundstellen, zusammengestellt aus Litera-
turquellen und neuen Untersuchungen. Nur fu¨nf Fundstellen
sind heute noch zuga¨nglich und derzeit Gegenstand wei-
terer Forschungen. Sa¨mtliche Fundstellen werden vorge-
stellt und der Kenntnisstand u¨ber Lithofazies, Stratigraphie,
Pala¨ogeographie und Pala¨oo¨kologie des Obernkirchener
Sandsteins zusammengefasst. Lithofazies-Daten zeigen,
dass eine feinstratigraphische Korrelation individueller
Horizonte innerhalb des Sandstein-Komplexes nur zwi-
schen sehr eng benachbarten Aufschlu¨ssen mo¨glich ist und
deutliche Faziesunterschiede zwischen den Fa¨hrtenvor-
kommen bestehen. Daher erscheint die derzeitige
Datengrundlage als nicht hinreichend, um den Obernkirchen-
Sandstein in seiner Gesamtheit als Megatracksite zu
charakterisieren.
Schlu¨sselwo¨rter Wirbeltier-Fa¨hrten  Dinosauria 
Kreide  Berriasium  Obernkirchen  Mu¨nchehagen 
Deutschland
Introduction
The Berriasian Bu¨ckeberg Formation is well-known for its
wealth of dinosaur tracks (Ballerstedt 1905; Dietrich 1927;
Lehmann 1978), but their documentation is mostly con-
fined to isolated specimens or localities. Important dis-
coveries of the last two decades are mostly reported in
general overviews or abstracts (Wings et al. 2005a;
Lehmann 2006; Lehmann et al. 2006; A. Richter et al.
2007; U. Richter et al. 2007; Richter et al. 2009; Richter
2007; Bo¨hme et al. 2009; van der Lubbe et al. 2009).
Nearly all tracksites from the Bu¨ckeberg Formation are
located stratigraphically within the Obernkirchen Sand-
stone, a thin subunit exposed in the northern and western
vicinities of Hannover. At least since early medieval times,
this sandstone has been a valued dressing stone and
building material. Intensive quarrying since the 11th cen-
tury AD led to the use of the nearly pure quartzose sand-
stone in hundreds of buildings and monuments across
northern Germany, Europe, and in places as far abroad as
Jakarta, Indonesia (Hamm 1938; Graupner 1977; Harnack
1989; Broschinski 2004). Since the beginning of the 19th
century, this formation became increasingly well-known as
a lagerstaette for rare but well-preserved Cretaceous
vertebrates, including turtles, crocodiles, and dinosaurs
(e.g. Roemer 1836; von Meyer 1841, 1857, 1859; von
Meyer in Dunker 1846).
Although extremely abundant and probably known to
quarry workers for centuries, reliable accounts of dinosaur
tracks were not reported before the middle of the 19th
century. According to Grabbe (1881), the mining supervi-
sor Heidtmeier reported the occurrence of a trail of con-
secutive tracks more than 100 m long which was
uncovered in a quarry near Wendthagen in the 1850s.
Unfortunately, there are no other records of this find.
In 1879, the first dinosaur tracks reported from Germany
rather sneaked into the scientific record. At the December
meeting of the Deutsche Geologische Gesellschaft (Ger-
man Geological Society), held on 3 December 1879, in
Berlin, Wilhelm Hauchecorne presented a letter and a
drawing by mining engineer Fritz F. von Du¨cker, in which
he reported the discovery of ‘‘bird tracks’’ in the Bad
Rehburg area. At the same meeting, Wilhelm Dames
commented that these tracks were very similar to those
from the English Wealden, which had been identified as
dinosaur tracks shortly before. Both contributions consisted
of a single sentence each, included in the published pro-
ceedings of the meeting (Dames in Beyrich et al. 1879;
Hauchecorne in Beyrich et al. 1879).
The geologist Carl Struckmann was the first researcher
to provide a scientific description of these tracks. During
the summer of 1879, more tracks were found at Wo¨lping-
hausen (Rehburg mountains, see locality 4 below) and
immediately vividly discussed in local press reports
(Anonymous 1879a, b). These early discussions, led mostly
by Struckmann, centred around a ‘‘bird-like’’ animal as a
potential trackmaker. This view was not left unchallenged–
in fact, the ornithologist Wilhelm Pralle suggested the tri-
dactyl imprints had been made by the mani (!) of giant
anurans (Anonymous 1879b). In 1880, Struckmann sub-
stantiated his results and published the first papers on the
‘‘Ornithoidichnites’’, referring them to the dinosaur
Iguanodon (Struckmann 1880a, b).
To corroborate his identification, Struckmann sent a
gypsum cast of one of the tracks to Louis Dollo, in Brus-
sels, who was the most important authority on Iguanodon
at the time, and was studying the famous ‘‘herd’’ of this
dinosaur found at the Bernissart coal mine in 1878. Dollo
(1883: 113) reported a perfect match of this track with the
pedal skeleton of a specimen he identified as Iguanodon
mantelli.
At least two specimens of Struckmann’s material are
still preserved in the Niedersa¨chsisches Landesmuseum,
Hannover. One right pes impression of a large ornithopod
(NLMH 105.747, Fig. 1) can be identified as the specimen
which was deemed an exception by him, as he believed it
was tetradactyl instead of tridactyl (Struckmann 1880a: 99,
232 J. J. Hornung et al.
123
pl. V, fig. 1; here Fig. 1b). This identification of the
specimen seems to be well substantiated, although the
figure by Struckmann was quite schematic in some details
and the proportions of the footprint and the supposed
‘‘hallux’’ seems not to have been depicted entirely correctly
by him. Subsequent studies (e.g. Grabbe 1883; Koken
1887; Ballerstedt 1914; Dietrich 1927) doubted the inter-
pretation of the ‘‘hallux’’. The specimen as preserved today
(Fig. 1a) supports the observation by Ballerstedt (1914)
that the ‘‘first digit’’ is in fact a poorly preserved footprint
over-stepped and deformed by the large, typically tridactyl,
ornithopod footprint (Fig. 1c). It is the historically oldest
find of a dinosaur track from Germany still preserved in a
public collection. Identification of the second specimen
(NLMH 105.746) is not as straightforward, because it
cannot be directly identified as one of those depicted by
Fig. 1 Right pes impression (hypichnial relief) of a large ornithopod
(NLMH 105.747), Bu¨ckeberg Formation, Obernkirchen Member,
Obernkirchen Sandstone. Most probably an original specimen of
Struckmann (1880a), collected in the summer of 1879 from the quarry
of A. Spo¨rl near Wo¨lpinghausen, Rehburg Mountains, Lower Saxony
(locality 4). This is, historically, the first footprint to be identified as
having been left by a dinosaur in Germany and is still preserved in a
public collection. a Specimen as preserved today; b Specimen as
drawn by Struckmann (1880a: pl. V, fig. 1). The structure below the
footprint was interpreted as a hallux impression by C. Struckmann.
This interpretation was rejected by subsequent scientists; c New
interpretation of the specimen (drawing). The grey-shaded structure is
interpreted as the remains of a much smaller tridactyl(?) footprint,
over-stepped by the ornithopod heading in the opposite direction
Vertebrate tracksites in the Obernkirchen Sandstone 233
123
Struckmann. However, he collected far more material than
he figured, and the size, proportions, and morphological
details of the specimen suggest that it belonged to a
trackway from which he figured two other, subsequent
footsteps, preserved on a single sandstone slab which is
currently lost (Struckmann 1880a: 98–99, pl. IV, fig. 1,
1880b: 127, pl. IV).
Struckmann’s discoveries were followed shortly by
those of Grabbe (1881, 1883), who identified dinosaur
tracks in many localities in the Bu¨ckeberge and Harrl areas.
Over the following decades, numerous reports were
published of tracksites in the outcrop area of the Obern-
kirchen Sandstone. Most of these appeared in hard-to-
obtain local journals, some of them even in newspapers,
and they are almost exclusively written in the German
language. Additionally, in many cases, the original mate-
rial was considered lost, hampering modern revisions
(Haubold 1971, 1974, 1984; Lockley 2000; Thulborn
2001).
The historical collection of the Gymnasium Adolfinum
Bu¨ckeburg and the Max Ballerstedt private collection are
highly important in investigations of Berriasian dinosaur
tracks from Germany. Although a few specimens from
these collections date back to the 1840s, the bulk of the
material was collected between ca 1900 and 1940 by the
teacher and amateur palaeontologist Max Ballerstedt
(1857–1945). The collection includes a large number of
footprints (mostly natural casts), some of which were ori-
ginal specimens of Ballerstedt’s publications (see also
Thulborn 2001). However, after Ballerstedt’s death in
1945, the collection suffered from lack of care and atten-
dance shortly after World War II (Probst and Windolf
1993; Hornung and Reich 2007). Most of what survived is
stored in the collections of the Geoscience Centre of the
University of Go¨ttingen. A detailed inventory or catalogue
of the Ballerstedt collection is currently incomplete. In an
ongoing project, this gap is to be filled and many of
Ballerstedt’s specimens, thought to be lost, have recently
been located and are, in fact, not lost (Hornung and Reich
2006, 2007; Hornung et al. 2007).
In addition to classical works and material, new evi-
dence has been gathered from the Obernkirchen Sandstone,
beginning with the discovery of sauropod tracks in 1979
(Hendricks 1981; Fischer 1998) and culminating in
continuing excavations at several localities, immensely
increasing the amount of data (A. Richter et al. 2007; U.
Richter et al. 2007; Richter et al. 2009; Bo¨hme et al. 2009;
van der Lubbe et al. 2009).
The objective of this paper is to comprehensively survey
of the known localities of vertebrate tracks in the Obern-
kirchen Sandstone and to provide an up-to-date account of
the stratigraphical and palaeoenvironmental data of this
remarkable formation. This should serve as easy access to
older locality descriptions and as a basis for further
investigation of ichnological issues in this formation. The
identification of these occurrences as a megatracksite in the
Obernkirchen Sandstone (as proposed by Diedrich 2004) is
also discussed.
Abbreviations
Institutional abbreviations BGR Bundesanstalt fu¨r Geo-
wissenschaften und Rohstoffe, Hannover, Germany;
GPMH Geologisch-Pala¨ontologisches Museum, Universi-
ta¨t Hamburg, Germany; GZG Geowissenschaftliches Zen-
trum, Georg-August-Universita¨t Go¨ttingen, Germany;
NLMH Niedersa¨chsisches Landesmuseum, Hannover,
Germany.
Palaeogeographical and stratigraphical abbreviations
DP: ‘Dachplatte’, hf: Hauptflo¨z (main coal seam), LSB:
Lower Saxony Basin (Niedersa¨chsisches Becken), OHS:
Oberer Hauptsandstein (Obernkirchen Sandstone sensu
stricto), UHS: Unterer Hauptsandstein.
General abbreviations H: Hochwert (northing, Gauss-
Kru¨ger coordinate system), R: Rechtswert (easting, Gauss-
Kru¨ger coordinate system).
Geology and environment of the Bu¨ckeberg formation
Exposure and stratigraphy
The Bu¨ckeberg Formation is buried beneath younger
deposits across wide areas of Lower Saxony but is exposed
in local uplifts, for example the mountainous areas of the
Bu¨ckeberge, the Deister, the Rehburg Mountains, the
southeastern Su¨ntel, the Osterwald, and the Hils mountains
(Fig. 2). South and west of the Weser Valley, it has been
completely eroded. The Obernkirchen Sandstone forms a
thin interval within the Obernkirchen Member, the lower
member of the Bu¨ckeberg Formation, stretching in a SW to
NE striking belt between the River Weser in the west and
Hannover in the east. In most areas where outcrops of
Obernkirchen Sandstone are revealed it was quarried,
especially for high-quality building stones. Additionally,
thin coal seams are intercalated with the sandstones and
have been mined in shallow subsurface mines since the
13th century AD until the 1960s (Falke 1944; Graupner
1980). However, following the decline of both industries,
mining has been completely abandoned and quarrying
substantially reduced, leading to a decrease in accessible
outcrops.
The Bu¨ckeberg Formation (Casey et al. 1975, tradi-
tionally referred to as the ‘German Wealden’, Hoffmann
1830, Fig. 3) comprises the middle Berriasian through
early Valanginian sedimentary infill of the Lower Saxony
234 J. J. Hornung et al.
123
Basin (LSB, Fig. 4). It is underlain by evaporitic marl-
stones of the Tithonian through lower Berriasian Mu¨nder
Formation and overlain by Valanginian marine deposits
(Kemper 1973; Strauss et al. 1993; Mutterlose 1997a,
2000). The predominantly limnic nature of the Bu¨ckeberg
Formation hampers exact correlation with the marine
Berriasian outside the basin (Mutterlose 1997a). Internal
correlation is provided by ostracod biostratigraphy (Wol-
burg 1949, 1959; Elstner and Mutterlose 1996). Older
lithostratigraphical schemes use a subdivision into six units
(W[ealden]1 to W6, Wolburg 1949). Current schemes
(Elstner and Mutterlose 1996; Mutterlose 1997a, 2000) use
subdivision into a basal Obernkirchen Member (corre-
sponding to the W1 to W4) and the overlying Osterwald
Member (W5 to W6).
The Obernkirchen Member can be subdivided into a fine-
grained argillaceous basin facies and a marginal, sandy facies.
The fine-grained basinal facies reaches a thickness of up to
700 m in the basin centre. It consists of siltstones, mudstones,
and black shales. In the entire succession, thin carbonate
layers are occasionally interbedded, mostly composed of
mollusc coquinas (Neomiodon spp. and gastropods).
The marginal facies attains a thickness of up to 200 m
along the southeastern basin margin, and accumulates
Fig. 2 Distribution of vertebrate tracksites in the Bu¨ckeberg Formation. Numbers refer to the locality numbers in the text
Vertebrate tracksites in the Obernkirchen Sandstone 235
123
especially in two sedimentary wedges (Fig. 4), an eastern
one in the Hannover area and a western one in the Osnabru¨ck
area (North Rhine-Westphalia, Kauenhowen 1927). The
western sediment wedge (Riegel and Lill 1988; Lill and
Riegel 1991; Wilde and Schultka 1996) has so far yielded only
one tridactyl cast (hypichnium), found in a field near Hilter-
Borgloh (Anonymous 2005). This single, yet undescribed
record suggests that the southwestern part of the basin margin
may also be significant source of dinosaur tracks. However, all
other finds are from the eastern wedge, and the discussion
herein will be restricted to this region.
The eastern sediment wedge fills a southeastern exten-
sion of the LSB, the Hils Embayment (Fig. 4). Its distal-
most outreaches consist of two sandstone tongues, a lower
one (Unterer Hauptsandstein, UHS, Figs. 3, 5) and an
upper one (Oberer Hauptsandstein, OHS, Figs. 3, 5, 6b),
separated by 10–25 m of fine-grained sediments. The UHS
attains a thickness of 2–5 m, the OHS of 10–20 m. Prox-
imally, along a SE–NW trending depositional axis, an
increasing number of thin sandstone layers become inter-
calated below, between, and above the UHS and OHS. The
OHS is equivalent to the Obernkirchen Sandstone sensu
stricto and the latter term is used herein exclusively as a
synonym of the OHS.
Thin intercalated coal seams are common throughout the
Obernkirchen Sandstone and its equivalents, including the
Osterwald and Hils mountains (Falke 1944; Graupner
1980; Pelzer et al. 1992). They were used in early attempts
at detailed intraformational lithostratigraphy of the sand-
stone succession across the Rehburg Mountains, Bu¨cke-
berge, and Harrl hill (Grabbe 1883; Grupe 1933; Falke
1944; Graupner 1980). Up to five superimposed, cm to
dm-thick coal seams have been reported intercalated with
the sandstones (Figs. 5, 6). However, lateral correlation of
these coal seams spanning wider distances is questionable.
Only the most important Hauptflo¨z (main seam, hf in
Figs. 5, 6) can be traced below the Obernkirchen Sandstone
across the Bu¨ckeberge, where a prominent coal layer, up to
80 cm thick, is embedded at the base of the sandstone
complex. A similar situation is present at the Rehburg
Mountains, where the thickest seam is located at the base
of the sandstone succession, suggesting a potential corre-
lation. The coal seams in the outcrops at Harrl hill are
intercalated within a different lithofacies succession and
previous correlation attempts, for example that by Grupe
(1933), are still very weakly supported.
In many outcrops and subsurface logs, the Hauptflo¨z is
separated from the overlying Obernkirchen Sandstone by a
Fig. 3 Stratigraphy, lithofacies,
and environment of the basal
Early Cretaceous in NW
Germany. Ostracods:
C. = Cypridea,
M. = Macrodentina,
P. = Pachycytheridea. After
Kemper 1973; Strauss et al.
1993; Elstner and Mutterlose
1996; Gramann et al. 1997;
Mutterlose 1997a, 2000; and
others, modified
236 J. J. Hornung et al.
123
0.5–1.5 m fine-grained interval, the ‘Dachplatte’ (DP,
Figs. 5, 6b). This unit is locally very fossiliferous and has
yielded especially rich plant and vertebrate remains
(Grabbe 1883; Dames 1884a).
The distal sandy complex correlates biostratigraphically
with the Cypridea alta formosa ostracod subzone (upper
Berriasian, Elstner and Mutterlose 1996) and is nested well
within the lower part of W3. Diedrich (2004) assigned the
upper part of the Obernkirchen Sandstone at the Obe-
rnkirchener Sandsteine GmbH quarry (locality 8) to the W4
(C. jonesi/C. parallela ostracod biozone). However, this is
not in agreement with other stratigraphical schemes (Jor-
dan 1979), and because Diedrich (2004) did not provide
any data or rationales for his correlation, it is not followed
herein.
South of Hannover, the sandy facies increasingly dom-
inates and conglomerate layers are rarely intercalated
(Struckmann 1880a). Because of the lack of adequate
lithostratigraphical and biostratigraphical marker-beds,
exact correlation of the UHS and OHS with the more
proximal sandy beds toward the SE is not possible.
The overlying Osterwald Member shows a retrograda-
tional facies trend with a margin-ward onlap of the fine-
grained basin facies. A marginal, coarse-grained facies is
restricted to the south-easternmost region of the Hils
Embayment (Pelzer 1998).
Palaeogeography
The LSB (Fig. 4) is an adjacent subbasin of the North
German Basin, which stretches across most of central
northern Germany and westwards into the eastern part of
The Netherlands (Schott et al. 1967, 1969). Its subsidence
began during the Late Jurassic and continued into the early
Late Cretaceous, when it was terminated by basin inversion
(Betz et al. 1987; Bachmann and Grosse 1989).
Communication of the LSB with the main North Ger-
man Basin was maintained only through narrow gateways
in the SW, W, and E (Schott et al. 1967, 1969). The basin
fill facies was governed primarily by eustatic sea level
fluctuations and/or local tectonic opening of gateways
which connected or unconnected the LSB from the Tethys
and Boreal Ocean (Kemper 1973, 1992; Elstner and Mut-
terlose 1996; Mutterlose and Bornemann 2000). Following
disconnection from marine influx, after deposition of the
marine-brackish to hyperhaline upper Mu¨nder Formation
(early Berriasian, Arp and Mennerich 2008), short-time
ingressions occurred from the W and SW (Holland) during
the latest W2 and throughout W3 (Elstner and Mutterlose
1996; Mutterlose and Bornemann 2000; Berner et al. 2010;
Berner 2011) resulting in a W–E-decreasing salinity gra-
dient across the LSB. The eastern part remained oligoha-
line until the latest Berriasian or earliest Valanginian (W4)
when marine microfaunas occur temporarily in the eastern
part of the basin also (Martin 1961a; Mutterlose and
Bornemann 2000). On a secondary scale, the LSB facies
and palaeogeography were controlled by autocyclical pro-
cesses, including local tectonics (including halotectonics
and subrosion of Permian evaporites), differential subsi-
dence, and intra-basinal, climatically controlled base-level
fluctuations during phases of disconnection from the global
sea-level (Mutterlose and Bornemann 2000). These factors
dominated the sedimentation within the LSB from the
Jurassic/Cretaceous boundary well through the latest
Berriasian.
After local highstand and marine flooding of most of the
incipient LSB during the Late Jurassic (Gramann et al.
1997; Ka¨stner et al. 2008), a major fall in sea level and
Fig. 4 Berriasian
palaeogeography (after Schott
et al. 1967, 1969; Kemper 1973,
Pelzer 1998, modified) and
vertebrate tracksite occurrences
in the Lower Saxony Basin
Vertebrate tracksites in the Obernkirchen Sandstone 237
123
Fig. 5 Lithostratigraphical correlation of outcrops in the Bu¨ckeberge
and Harrl areas. Note the weakly supported correlation of the track-
bearing beds of Harrl hill with those of the central Bu¨ckeberge.
Encircled numbers refer to locality numbers in text. Lithological data
after Dietrich 1927, Grupe 1930, Pelzer 1998, and our own
observations
238 J. J. Hornung et al.
123
early basin subsidence led to isolated, shrinking perimarine
water bodies in the developing LSB during deposition of
the Tithonian through earliest Berriasian Mu¨nder Forma-
tion. These beds were regarded as the equivalent of the
English Purbeck beds in the 19th and early 20th century
literature (Struckmann 1880a). The lower part of the
Mu¨nder Formation is dominated by marlstones, locally
evaporitic (Katzberg Member) and contains low-diversity
faunal assemblages with dominant local biostromes of the
sedentary polychaete Serpula coacervata (Blumenbach,
1803) and increasing influx of freshwater in the upper part
of the Mu¨nder Formation (Serpulite Member, Borberg
Member; Arp and Mennerich 2008).
During early to mid-Berriasian times, precipitation
increased and a lacustrine water body began to spread in
the basin centre, fed by river systems draining the basin
margins and uplands (Pelzer 1987; Pelzer and Wilde 1987;
Pelzer et al. 1992; Wilde et al. 1995). The river systems
reached substantial dimensions with meandering beds and
extensive belts of well-vegetated floodplain deposits,
backswamps, and associated lakes, as documented by
related deposits in the southeast of the LSB (Pelzer 1984;
Pelzer et al. 1992). The relatively undisturbed succession
of fine-grained deposits in the basin centre (Kemper 1973)
suggests that a central lake was continuously present
throughout the Berriasian. However, transgressive–regres-
sive patterns of this lake are not well known. Transgressive
sequences can be traced in parts of the succession and an
overall expansion of the lake is assumed for most of its
existence. The fluvial facies is exposed in the Hils and
Osterwald mountains (Pelzer 1984, 1998; Pelzer et al.
1992; Riegel et al. 1986).
It is of importance to note that the vertebrate track-
bearing deposits of the Obernkirchen Sandstone are all
located within this limnic depositional system, contrary to
a few publications (Diedrich 2004; Lockley et al. 2004)
which regard this as marginal-marine in origin.
In the Osterwald mountains, fluvial large-scale cross-
stratified, up to 12 m thick (channel-fill?) sandstones
(Obernkirchen Member) are abruptly overlain by a thin
conglomerate, interpreted as a transgressional beach
deposit, passing upwards into perideltaic brackish-marine
sediments (Osterwald Member; Waldeck 1969; Pelzer
1998). Palynological data suggest that the marginal river
systems were still active in Valanginian times (Pelzer
1984) when the basin centre was reoccupied by marine
waters because of eustatic sea level rise.
At least one fluvial system apparently occupied the axis
of the Hils Embayment. The system terminated down-
stream in the Bu¨ckeburg–Bad Rehburg–Hannover area,
where an extensive, shallow-water deltaic system
(Kauenhowen 1927; Graupner 1980; Pelzer et al. 1992;
Pelzer 1998) became established during the mid- to late
Berriasian. This system is represented by the distal reaches
of the Hannover sedimentary wedge. The sandstones and
intercalated fine-grained sediments preserve a diverse
assemblage of sedimentary facies, representing several
subenvironments. Pelzer (1998) proposed a barrier/back-
barrier setting for the Obernkirchen Sandstone with severe
storm events as an important landward transport mecha-
nism for reworked and matured terrigenous sand in this
microtidal environment. Current work suggests that the
Obernkirchen Sandstone encompasses several facies
assemblages, including back-barrier storm deposits in the
Rehburg Mountains and a landward deltaic succession in
the southern and western Bu¨ckeberge (see the discussion
below).
On the landward side of the lagoons, swamps existed,
passing into peat-bogs and vegetated areas on emergent
and stabilised interchannel and interlagoonal islands
(Wilde et al. 1995; Pelzer et al. 1992; Pelzer 1998).
The dinosaurs apparently passed across the shore-face,
back-barrier, deltaic and alluvial flats, and through shallow
lagoonal waters. The preservation of multiple, vertically
stacked barrier and lagoonal deposit successions within the
Obernkirchen Sandstone and abundant interbedded palae-
osols and authochthonous coal seams indicate a long-term
net aggrading sedimentary regime under transgressive
conditions, favouring also the preservation of vertebrate
tracks. However, erosional discontinuities on a local scale
have been reported in some outcrops (Fischer 1998; Pelzer
1998).
Palaeoecology
The only well-studied microfaunal group are the ostracodes,
which provide the biostratigraphic framework of the non-
marine Berriasian (Martin 1940, 1958, 1961b; Wolburg 1949,
1959, 1962, 1971; Schudack and Schudack 2009).
The invertebrate macrofauna in the lagoonal deposits
is dominated by mollusc assemblages (predominantly
Unionidae, Neomiodontidae and various gastropods; Dun-
ker 1846, 1849; Struckmann 1880a; Huckriede 1967).
Modern detailed taxonomic and diversity studies of these
groups are still lacking. Abundant endobenthic invertebrate
trace fossils (Schwennicke 1998) indicate normally oxy-
genated conditions for the lagoonal water bodies in the
Mu¨nchehagen area.
Aquatic vertebrates are represented by elasmobran-
chians (hybodontoid, petalodontid, and neoselachian
sharks) and several species of actinopterygian fishes,
including members of Semionotiformes, Pycnodontifor-
mes, Amiiformes, and Pholidophoriformes (Dunker 1846;
Struckmann 1880a; Branco 1887; Martin and Weiler 1954;
Schultze 1970; Lo´pez-Arbarello et al. 2007, Nyhuis and
Herbig 2009). A maximum of fish diversity is known from
Vertebrate tracksites in the Obernkirchen Sandstone 239
123
240 J. J. Hornung et al.
123
concentrated, tempestitic bone-beds, deposited in the
Osterwald Member in the western, deeper basin (Nyhuis
and Herbig 2009). In the southeastern, sandy facies of the
Obernkirchen Member, the known fish fauna is so far
confined to hybodontoid sharks (fin spines, teeth, and egg
capsules) and semionotids (including some splendid spec-
imens of ‘‘Lepidotes’’ mantelli Agassiz, 1833; see Branco
1887; Lo´pez-Arbarello et al. 2007).
Plesiosaurs also inhabited the LSB, including Branca-
saurus brancai Wegner, 1914 known from a rather com-
plete skeleton from the Osterwald Member of Gronau/
Westphalia (Wegner 1914). Isolated plesiosaur skeletal
elements are furthermore known from various localities
and facies (Koken 1887, 1896). Semiaquatic vertebrates
are mostly represented by the very abundant and diverse
turtles (at least seven species of basal cryptodirans; Grabbe
1884; Peitz 1998; Karl et al. 2007a, b) and by the common
mesoeucrocodylian crocodile genera Goniopholis Owen,
1842 and Pholidosaurus von Meyer, 1841 (von Meyer in
Dunker 1846; Koken 1883, 1886, 1887, 1896; Jaffe´ 1912;
Edinger 1938; Salisbury et al. 1999).
The fauna of fluvial and alluvial environments is less
well-known, although mollusc shell-beds and bone-bed
layers are present (Pelzer 1984, 1998). Fine-grained over-
bank deposits preserved a few terrestrial arthropods, for
example insects (Coleoptera) and possibly arachnids
(Struckmann 1880a; Pelzer 1984; Pelzer et al. 1992).
The terrestrial vertebrate fauna was mainly composed of
dinosaurs, but only the small ornithischian Stenopelix vald-
ensis von Meyer, 1857 is represented by a largely complete,
articulated postcranium (Schmidt 1969; Sues and Galton
1982; Butler and Sullivan 2009). Besides the track record,
Ornithopoda, Ankylosauria, Theropoda, and Sauropoda have
also been identified from isolated bone fragments (Dames
1884a, b; Koken 1887; Struckmann 1894; Sachs 1997;
Windolf 1998; Hornung unpublished data).
A single specimen of pterosaur, the holotype of
Ctenochasma roemeri von Meyer, 1851, was found in the
Deister mountains. Although considered to be Late Jurassic
in age, for example by Wellnhofer (1991), the information
provided by von Meyer (1851) supports an origin from the
Serpulite Member of the Mu¨nder Formation and therefore a
basal Berriasian age, as suggested by Bennett (2007). So
far no pterosaurs are known from the limnic mid- to late
Berriasian deposits.
The palaeoflora is known from leaf impressions in fine-
grained sediments, stem-impressions in sandy sediments,
macro- and microremains from lignites, and root-traces,
including strongly rooted palaeosol horizons, and, rarely,
amber. Palynomorphs and macroremains indicate various
floral communities, including pioneer, swamp and flood-
plain vegetation, and forested areas (Pelzer 1984, 1998;
Pelzer et al. 1992). Aquatic plants are represented by
dinoflagellates and charophytes (Schudack 1996; Pelzer
1998). Major terrestrial plant taxa include Coniferae,
Equisetales, Ginkgophyta, Cycadophyta, Pteridophyta,
Bryophyta, and Fungi (Lipps 1923; Ma¨gdefrau and Rudolf
1969; Hiltermann 1949; Pelzer 1984, 1998; Riegel and
Wilde 1984; Wilde and Heunisch 1990; Wilde 1991; Wilde
et al. 1995). Various morphotypes of root-traces, rhizomes
of Equisetales and log impressions are very common plant
remains associated with the track-bearing sandy facies
(Pelzer 1998).
Facies and lithology of vertebrate track-bearing
horizons
The vast majority of vertebrate (mostly dinosaur) tracks
from the Obernkirchen Member are from the limnic del-
taic-estuarine deposits, more precisely from the OHS del-
taic-estuarine sandstones. However, some tracks have also
been described from the fluvial and alluvial facies in the
Osterwald mountains (Naumann 1927).
Dinosaur track horizons in the deltaic-estuarine envi-
ronment commonly occur in mm- to dm-thick sandstone
beds interlayered with mm- to cm-thick sandy or silty
claystones and mudstones. The sandstones are often mas-
sive or internally ripple to large-scale cross-stratified,
horizontally laminated or flaser-bedded with oscillation or
unidirectional ripple-marks at the top. Thick-bedded
sandstone layers are often composed of amalgamated
sublayers.
Lithofacies data indicate that vertebrate tracks were left
in various subenvironments of the deltaic-estuarine system.
In the Mu¨nchehagen area (localities 1–3, Fig. 7a), the
tracks were left on top of dm-thick sandstone beds, which
have a slightly erosive base and a predominantly massive
or diffusely horizontally stratified texture. The uppermost
centimetres are wave-reworked and have well-preserved
oscillation-ripple cross-lamination and intensive inverte-
brate bioturbation, overlain by 1–10 cm thick drapes of
siltstones to mudstones. The thick bedding and massive
texture indicates rapid deposition from voluminous highly
concentrated density flows with limited turbulence at the
flow base. The best preserved dinosaur tracks and ripple-
marks occur at the contact between the top of sandstone
beds and the base of relatively thick (2–5 cm) siltstones
to claystones. These boundaries contain sub-millimetric,
smooth, carbonaceous films covering the surface of
the oscillation ripple-marked sandstones. The films are
Fig. 6 a Geographical overview of the tracksites near Mu¨nchehagen
(localities 1–3, Rehburg Mts.); b Simplified lithological logs of
localities 1–3, showing lateral correlation and track horizons.
Lithological data after Wilde et al. 1995, Fischer 1998, Pelzer
1998, and our own observations. Track symbols are eplained in Fig. 2
b
Vertebrate tracksites in the Obernkirchen Sandstone 241
123
Fig. 7 Typical dinosaur track-bearing lithofacies of the Obernkirchen
Sandstone. a Thick-bedded, massive sandstone beds with oscillation
ripple-marks at the top, intercalated by mm to cm-thick sandy siltstone to
claystone layers. In the foreground a large ornithopod track. The
sandstone beds in the background have a thickness of 70–100 cm.
Mu¨nchehagen/F. Wesling GmbH quarry (ch) (locality 1), 2005;
b Medium-bedded, partly amalgamated and channellised (ch) fine-
grained sandstone beds, intercalated by mm to cm-thick siltstone to
claystone layers. Note the overall thinning-upward trend. The surface at
the base of the image is the so-called ‘‘chicken yard’’ track horizon
(Fig. 15). ‘‘Upper’’ and ‘‘lower unit’’ refer to a lithological subdivision,
which can be traced between adjacent quarries at a local scale (see locality
description for further details). Scale: 2 m. Obernkirchen/Obernkirchener
Sandsteinbru¨che quarry (locality 8), 2009; c Channellised (ch), medium
to thick-bedded sandstone, ‘‘lower unit’’, Obernkirchen/Obernkirchener
Sandsteinbru¨che quarry (locality 8), 2009; d Thin to medium-bedded,
scoured, massive sandstone beds, overlain by a soft succession of sandy
siltstone to mudstone and capped by a thin coal-seam (cs). Pelzer (1998)
tentatively identified a track horizon (TH) in this succession. Bu¨ckeburg/
abandoned quarry at the W’ side of Harrl hill (locality 5), 2009
242 J. J. Hornung et al.
123
probably the remnants of microbial mats (Eriksson et al.
2007) which grew on the freshly deposited sand grounds
before being covered by the subaqueous deposition of pe-
litic sediments. They probably helped to prediagenetically
stabilize ichnia and sedimentary structures by providing
increased cohesion. Stacked and partly amalgamated
sandstone layers are in places separated by wavy-irregular,
often laterally discontinuous boundary surfaces, often
covered by very thin clay vails. Seen from above, these
surfaces have a reticulated pattern of mm-wide grooves
separated by sharp ridges. Similar structures have been
described to be formed by desiccated microbial mats in
ephemeral pools on supratidal sandbars in extant environ-
ments (Bose and Chafetz 2009). Together with extremely
rare desiccation cracks these structures are indicative of
temporary emersion of the sandy deposits.
These deposits are interpreted to be associated with
storm-reworked barrier sands, splayed into back-barrier
lagoons (Pelzer 1998). Such an event-based sedimentation
is concordant with the presence of cluster-like colonies of
fully grown, gregarious, infaunal unionoid bivalves (Fig. 8,
see also Schwennicke 1998). Growth to several centimetres
in shell-length required years to decades before the popu-
lation was extinguished or forced to relocate by rapid
burial. The 101 to 102 yr frequency of high-energy depo-
sition supports an association with severe storm events.
The lacustrine setting, in concordance with the absence
of clear indicators of regular rhythmic tidal currents (e.g.
herring-bone cross-stratification) and rarity of signs of
sustained unidirectional currents, precludes astronomical
tides as sediment-transport mechanism. However, storm-
induced tides, piling water in shallow littoral regions as the
Hils Embayment may explain temporal inundation of
intertidal to supratidal areas after the deposition of storm-
mobilized sediments. Subsequent desiccation of such iso-
lated supratidal water bodies can explain the occurrence of
features such as the reticulated microbial mat structures
(compare Bose and Chafetz 2009).
The relative abundance of interspersed channel- and
scour-fill sandstones, often containing root-traces and
Fig. 8 Lockeia isp. (NLMH 105.745A), resting traces of unionid
bivalve colony at the underside of a dm-thick, massive sandstone layer
indicating sudden burial under rapidly deposited sediment after
prolonged period of undisturbed growth. Bu¨ckeberg Formation,
Obernkirchen Member, Mu¨nchehagen/F. Wesling GmbH quarry
(locality 1). Scale bar 5 cm
Vertebrate tracksites in the Obernkirchen Sandstone 243
123
separated by knobby-irregular, clay-draped boundary surfaces
at Obernkirchen (Bu¨ckeberge, locality 8, Fig. 7b, c) indicate a
facies setting rather different from that at the Rehburg Moun-
tains. The knobby surfaces are always associated to rooted
horizons. These surfaces, which were first described as stacked
palaeosols by Grupe (1931), are very similar to structures
formed by desiccated microbial mats on vegetated, supratidal
sandbars (Bose and Chafetz 2009). They indicate subaerial
exposure which lasted long enough to enable the growth of a
rather dense vegetation cover. Channel-fill deposits occasion-
ally have large-scale trough cross-stratification and coarser-
grained basal lags. Together with the diffusely horizontally
stratified or structureless, laterally extensive sandstone beds,
these observations suggest a shallow-water deltaic mouthbar
complex, in which the tops of mouthbars became emersed and
vegetated following the drop of the local water-table after
periods of very strong, probably seasonal discharge. Dinosaur
tracks occur on the top of channel complexes and on rooted-
knobby surfaced and plain sandstone layers. The quality of
track preservation varies substantially, but is generally inferior
to that in the Mu¨nchehagen area. Track-covered surfaces were
probably at least partially subaerially exposed and degraded
because of the lack of a protective mud layer, vegetation
growth, and surface modification by desiccating microbial
mats (compare Marty et al. 2009).
At Harrl hill (localities 5 and 6, Fig. 7d), at least one
track horizon is situated near the top of a thinning-upward,
scoured sandstone succession, passing upwards into several
dm of ripple cross-laminated siltstones to mudstones and
capped by a coal seam (Pelzer 1998). The facies assem-
blage suggests an aggrading mouthbar under transgressive
conditions, overlain by proximal shallow-lagoon sedi-
ments, when the base-level rise forced retrogradation of the
system or the mouthbar became inactive by avulsion.
Finally, a peat-bog swamp became established under
highstand conditions. The dinosaur tracks were left on top
of the shallowly submerged mouthbar.
The dinosaur tracks from fluvial deposits at the Osterwald
mountains (localities 12 and 13) are preserved at the base of
medium-bedded, partly current-ripple cross-laminated sand-
stones, overlying partly coal-bearing clay- and mudstones rich
in plant macrofossils (Naumann 1927). Crevasse-splay
deposited sand probably rapidly covered the fine-grained
overbank mud in which the tracks were made.
Tracks are commonly preserved as hyporeliefs on the
lower surface of sandstone beds. The fine-grained intervals
are poorly inundated and the trackway impressions (posi-
tive epireliefs) are often destroyed during exposure if not
preserved by artificial casting (Grabbe 1881; Hamm 1938;
Lehmann 1978). However, this type of preservation may be
over-represented in historical collections as it was logisti-
cally easier to collect blocks with isolated hypichnia than
to remove the track-bearing layer. Thin mudstone interca-
lations support the preservation of epireliefs on an under-
lying sandstone surface either by penetration of the mud
layer by the foot or by undertracks. Thinly bedded sand-
stones with intercalating mudstone laminae also favour the
preservation of undertracks.
Vertebrate track inventory of the Bu¨ckeberg Formation
(Table 1)
As this work is not intended as a thorough ichnotaxo-
nomical revision of the vertebrate ichnofauna, which will
be given elsewhere, only a short overview and
Table 1 Distribution, taxonomy, and abundance of dinosaur tracks across the localities discussed herein
Region Rehburg Mountains Harrl hill Bu¨ckeberge Osterwald Mts.
Locality 1 2 3 4 5 6 7 8 9 10 11 12 13
SAUROPODA ?
THEROPODA
‘Bueckeburgichnus’ maximus ? ? ?
‘Megalosauripus’ isp. ? ? ?
Didactyle morphotype ?
ORNITHOPODA ? ?? ?? ? ?? ?? ?? ? ? ?
ANKYLOSAURIA
Metatetrapous valdensis ?
UNID. TRIDACTYLE
DINOSAUR TRACKS
? ? ? ?
CHELONIA ?
CROCODYLIA ?
Abundance: (?) rare to very rare, (??) abundant; (?) identification uncertain
244 J. J. Hornung et al.
123
terminological framework for the locality section will be
given here. Several dinosaur ichnotaxa have been formally
established on the basis of material from the Bu¨ckeberg
Formation; however, virtually none of these are correctly
diagnosed or undisputed.
Sauropoda (Fig. 9a)
Sauropod trackways (Fig. 9a) are known only from the
‘‘Dinosaurier-Park Mu¨nchehagen’’ open-air museum
(locality 2; Hendricks 1981; Lockley et al. 2004). A new
ichnotaxon, Rotundichnus muenchehagensis Hendricks,
1981, has been established for these tracks, but this has not
found wide acceptance, because the tracks do not preserve
many morphological details (Lockley et al. 1994, 2004;
Wright 2005). Their general morphology is similar to that
of the wide-gauge ichnogenus Brontopodus Farlow, Pitt-
man and Hawthorne, 1989. Despite confirming its status as
nomen dubium, Wright (2005) proposed characters to
distinguish Rotundichnus from Brontopodus and denied the
referral of the Mu¨nchehagen tracks to the latter. The main
differences pointed out by Wright (2005) were subcircular
Fig. 9 Dinosaur track
morphotypes from the
Obernkirchen Sandstone.
a Brontopodus
isp. = (Rotundichnus
muenchehagensis Hendricks,
1981), sauropod trackway,
locality 2 (from Lockley et al.
2004, modified);
b ‘Bueckeburgichnus’ maximus
Kuhn, 1958, large theropod
track, locality 6 (from Lockley
2000, modified);
c ‘Megalosauripus’ isp.,
moderately sized theropod
track, locality 8 (from Diedrich
2004, modified); d Large
ornithopod pes track, the
presence of a vestigial hallux in
this track morphotype is
questionable, locality 7 (after
Dietrich 1927; Kuhn 1958,
modified); e Large ornithopod
trackway with manus imprint,
locality 2 (from Lockley et al.
2004, modified); f Large
ornithopod trackways, adult and
subadult/juvenile tracks, locality
8 (from Lehmann 1978,
modified); g Metatetrapous
valdensis Nopcsa, 1923,
ankylosaur track, locality 6
(from Ballerstedt 1922,
modified). Scale bar 1 m, note
different scale in a and b–g
Vertebrate tracksites in the Obernkirchen Sandstone 245
123
manus impressions (narrowly arc-shaped in Brontopodus) and
a manus width circa two times the manus length (nearly equal
in Brontopodus). However, the manus impressions are more
crescent-shaped in Rotundichnus than illustrated in Wright
(2005: fig. 9.5J and 9.5G respectively, compare with Lockley
et al. 2004: figs. 4–5 and to Fig. 9a herein).
An occurrence of Brontopodus isp. sauropod tracks at
Harrl hill as indicated by Diedrich (2004: fig. 1) is not
supported by previous reports or the original material.
Theropoda (Figs. 9b, c, 16b, c, 18a, b)
Theropod tracks (Fig. 9b, c) are common in the LSB but
have been misidentified as ornithopod tracks on various
occasions. The first description of theropod tracks from the
Obernkirchen Sandstone were given by Grabbe (1883: 20),
who noted moderately sized tridactyl footprints (footlength
ca 25–30 cm) with an asymmetric metatarsal region and
distinct claw marks. Unfortunately, he did not provide
figures or a detailed locality (?Harrl hill). The first indis-
putable figures of theropod tracks were published by
Ballerstedt (1905); these were from Harrl hill.
Theropod footprints occur in various size classes and
morphotypes. One distinct morphotype is represented by
large, functionally tridactyl footprints, which often preserve
the small digit I on deep casts and show a broad, robust digit II
(Fig. 9b). This morphotype has been named Bueckeburgich-
nus maximus Kuhn, 1958 (see Lockley 2000; Lockley et al.
2004). However, Thulborn (2001) reasonably argued that the
ichnogenus Megalosauripus Lessertisseur, 1955 is available
as a valid senior synonym of Bueckeburgichnus. We tenta-
tively retain here the name ‘Bueckeburgichnus’ for the sake of
distinction between those theropod tracks which have been
included in the ichnotaxonomic concept of the ichnogenus
Megalosauripus Lockley, Meyer and dos Santos, 1998 non
Lessertisseur, 1955. Because a second morphotype of thero-
pod tracks (Figs. 9c, 16b, 18a, b) is also consistent with the
current definition of the latter (Lockley et al. 1998), we use
here the name ‘Megalosauripus’ to distinguish these from
‘Bueckeburgichnus’. However, both designations are unsat-
isfactory in the long term, and this plexus of theropod ichno-
taxa is in desperate need of revision.
Yet unpublished material includes other types of the-
ropod tracks which will be listed as ‘‘theropod tracks’’
generally. These include recently discovered didactyl
trackways at Obernkirchen, apparently left by theropods
with a specialised sickle-shaped claw on digit II (probably
troodontids, van der Lubbe et al. 2009, Fig. 16c).
Ornithopoda (Figs. 1, 9d, e, f, 11, 12, 14, 16e, 17a, 18a, b, 19)
Ornithopods are by far the most abundant dinosaurs in the
Bu¨ckeberg Formation ichnofauna (Fig. 9d, e, f). Even their
earliest discoveries have been associated with the genus
Iguanodon Mantell, 1825 from the Barremian through
Aptian of southern England (Struckmann 1880a, b).
Although Struckmann used the descriptive term ‘‘Ornith-
oidichnites’’ coined by Beckles (1852, 1854) for similar
tracks from England, most subsequent authors addressed
them simply as ‘‘Iguanodon tracks’’ or even as ‘‘Iguan-
odon’’ (Stechow 1909; Ballerstedt 1914; Lehmann 1978;
Richter 2007). It had already been noted by Kuhn (1958)
and others that such a designation is taxonomically incor-
rect, because:
1. it cannot be demonstrated positively that the (geolog-
ically younger) orthotaxon Iguanodon had produced
the tracks; and
2. naming tracks by the same formal taxonomic group as
their producer is only admissible for extant tracks.
Alternatively, the term ‘‘iguanodontid tracks’’ was also
used (first by Grabbe 1881). However, this spelling is
problematical too, because a monophyletic family Igua-
nodontidae was not recognised by recent analyses of
ornithopod relationships (Norman 2004; Paul 2008; Butler
et al. 2008). Nonetheless, attribution of the tracks to
(a) relatively large-sized, basal member(s) of Iguanodontia
(in the sense of crown-group Ornithopoda, e.g. Norman
2004) is, of course, indisputable.
More recent papers (Lehmann 2003, 2006; Diedrich
2004; U. Richter et al. 2007; Hornung and Reich 2007)
recognised the ichnogenus Iguanodontipus Sarjeant, Delair
and Lockley, 1998, however without a proper rationale for
this assignment. As this ichnogenus was introduced by
Sarjeant et al. (1998) for a distinct morphotype of tridactyl
ornithopod tracks and not intended as a catch-all ichno-
taxon for Lower Cretaceous iguanodontian tracks, its
general use for the LSB large ornithopod tracks is rejected
herein. Indeed the morphological diversity suggests the
presence of at least 2–3 morphotypes among the large
ornithopod tracks from the LSB (Hornung unpublished
data). These include a new, recently discovered morpho-
type with two additional metapodial pad impressions at the
foot base from Obernkirchen (Bo¨hme et al. 2009, unpub-
lished data).
Because of these complications, ornithopod tracks will
not be further differentiated at ichnogenus level in the
locality descriptions. Several observations on well-pre-
served trackways have shown that some of the large
ornithopod trackmakers were quadrupedal and various
ontogenetic stages are present (Lockley et al. 2004; Died-
rich 2004; A. Richter et al. 2007; Bo¨hme et al. 2009;
Figs. 9e, f, 16e).
Ballerstedt (1905, 1921a) reported the existence of a
didactyl ornithopod track which he named Struthopus
schaumburgensis Ballerstedt, 1921. The location of the
246 J. J. Hornung et al.
123
type material of this ichnotaxon is currently not known, but
from the published figures it can be inferred that it is not
identical to the newly discovered didactyl theropod tracks
from Obernkirchen. Given the variation observed in
ornithopod tracks from the Obernkirchen Sandstone, it is
highly probable that S. schaumburgensis is an extramor-
phologically modified tridactyl track or one left by a
pathological individual (as proposed, for example, by Abel
1935). In the latter case, Ballerstedt’s ichnotaxon would be
justified and valid; however, this can only be decided with
the original material at hand.
Small bipedal ornithischians (Fig. 16d)
Recently, small trackways, possibly left by bipedal cursorial
ornithischians have been found at Obernkirchen (Fig. 16d).
Their generalised morphology potentially agrees with various
groups of ornithischians, including derived ornithopods, basal
thyreophorans, and marginocephalians. Detailed studies of
these tracks are in preparation.
Ankylosauria (Fig. 9g)
The first trackway of a quadruped dinosaur from the
Bu¨ckeberg Formation was described by Ballerstedt (1921b,
1922) from Harrl hill near Bu¨ckeburg (Fig. 9g). It was
named Metatetrapous valdensis and identified as the
trackway of a thyreophoran ornithischian by Nopcsa
(1923). The validity and identity of this ichnotaxon were
long doubted (Schmidt 1969; Haubold 1974), as the only
known documentation of the trackway was a sketch by
Ballerstedt (1922). However, recently some parts of the
ichnoholotype have been identified from the Ballerstedt
collection confirming the observations by Ballerstedt and
Nopcsa and supporting its description as an ankylosaurian
trackmaker (Hornung et al. 2007, unpublished data).
Non-dinosaurian vertebrate tracks
Non-dinosaurian vertebrate tracks are quite rare in the
Bu¨ckeberg Formation. Recently, a well-preserved cast of a
manus imprint from a large turtle (possibly Hylaeochelys
menkei) from the Harrl hill locality was identified in the
Ballerstedt collection (Hornung et al. 2008). Diedrich
(2004) noted scratch-marks (?swimming-track) made by a
crocodilian from the Dinosaurier-Park Mu¨nchehagen open-
air museum quarry but gave no further details or figures.
A strong discrepancy in diversity and composition exists
between the vertebrate fauna represented by tracks and by
body fossils from the same lithological unit (Table 2).
Crocodiles and turtles are very abundant (with a very
scarce track record), whereas large ornithopods and
theropods (for which an extremely abundant track record
exists) are only represented by a few teeth and isolated
bones (Dames 1884a, b; Koken 1887; Struckmann 1894;
and unpublished material). This suggests high taphonomi-
cal control of both body and trace fossil preservation. The
dinosaur tracks represent a local fauna at least temporarily
autochthonous to the delta and shoal system (large orni-
thopods, theropods, rare sauropods, and ankylosaurs).
Some body fossils (e.g. Stenopelix valdensis) are allo-
chthonous components from other habitats, which must
have existed in the catchment area of local fluvial systems.
Vertebrate tracksites in the Bu¨ckeberg formation
In the sections below, all data about the published dinosaur
tracksites from the Obernkirchen Sandstone are compiled.
Some emphasis is given to those localities which are currently
not accessible and for which data are compiled from the dis-
persed literature. Localities with current excavation and
research coverage are only rather briefly introduced, because
their detailed description will be the objective of future work.
The locality numbers refer to the map in Fig. 2.
It must be stressed that the rather deliberate use of
locality names in the past may have led to confusion. For
example, Diedrich (2004) distinguished between two
tracksites in the Rehburg Mountains, ‘‘Bad Rehburg’’
(based on Struckmann 1880a, b and Stechow 1909) and
‘‘Mu¨nchehagen’’ (based upon Fischer 1998). In fact,
Struckmann (1880a, b) referred to specimens found at
Wo¨lpinghausen (here locality 4) and Stechow (1909) and
Fischer (1998) to material from Mu¨nchehagen. Bad Reh-
burg, situated 1.5 km E of Mu¨nchehagen, has not yielded
any tracksite yet. Its use in the older references was
Table 2 Fossil record and taphonomy of tetrapods from the Obern-
kirchen Sandstone
Fossil record Body fossils Trace fossils
Vertebrate Taxa Preservation Abundance Abundance
DINOSAURIA
Sauropoda (f) ? ?
Theropoda (f) ? ??
Ornithischia
basal Iguanodontia (f) ? ???
Stenopelix valdensis (a) ? ?
Ankylosauria (f) ? ?
CROCODYLIA (a) ?? ?
CHELONIA (a) ??? ?
Preservation coded as (a) articulated material, (f) fragmentary and/or
isolated material
Abundance: (?) single specimen, extremely rare, (??) few speci-
mens, rare, (???) numerous specimens
Vertebrate tracksites in the Obernkirchen Sandstone 247
123
because of its status as an administration centre (commu-
nity of Rehburg-Loccum).
Localities 1 to 4: Rehburg Mountains
Three of the four known tracksites from the Rehburg Moun-
tains (localities 1–3) are clustered in the immediate vicinity E
of the town of Mu¨nchehagen. They expose nearly the same
stratigraphical interval. At all three outcrops, the succession
begins ca 1 m below the outcrop base with the Hauptflo¨z coal
seam, which is overlain by a sandstone layer with a rugged,
eroded surface on top. This discontinuity is found at the same
level in all three outcrops (Wilde et al. 1995), and the
boundary surface bears dinosaur tracks in two of them
(localities 2 and 3). The overlying sandstones are laterally
discontinuous and cannot be correlated with certainty.
1. Mu¨nchehagen/F. Wesling GmbH quarry (Figs. 6, 7a, 10, 11)
Coordinates N 5226037.3900, E 912005.9000; R: 3513680,
H: 5812380; 1:25000 German Topographical Map refer-
ence grid sheet 3521 Rehburg.
Stratigraphic position Lower part of Obernkirchen Sand-
stone, ca 3–5 m above Hauptflo¨z coal seam.
Track inventory Large and medium-sized ornithopods
and theropods.
Remarks The quarry of the F. Wesling GmbH company
exposes ca 6–7 m of 0.1–1 m thick sandstone, intercalated
by 0.05–0.2 m thick mudstone and siltstone layers. Two
horizons with well-preserved trackways are present, sepa-
rated by ca 2 m of sandstone. Abundant ([15) ornithopod
trackways left by bipedal and quadrupedal individuals in
various ontogenetic stages were present on both track
levels. Several of the animals walked subparallel to each
other. One ornithopod trackway consisted of at least 57
steps (Fig. 11). Medium-sized theropods are represented by
several trackways on both track horizons. The quarry is still
in production; contemporaneous to the quarrying activities,
systematic rescue excavations, surveys and casting have
been carried out by the Dinosaurier-Park Mu¨nchehagen,
the NLMH, and collaborating institutions since 2004. As
not all track-bearing layers could be preserved, contem-
poraneously to the documentation of the track-site, a ca
Fig. 10 Mu¨nchehagen, F. Wesling GmbH quarry (locality 1) and
Dinosaurier-Park Mu¨nchehagen open-air museum (locality 2, com-
pare Fig. 6a). Aerial view of the excavations and the exhibitions (in
the upper right corner) in 2005. The tracks discovered in 1979 are
partly protected by the building in the background. Photo by P. Nisi
(NLMH)
248 J. J. Hornung et al.
123
20 m2-sized slab of the lower track horizon was removed in
2005 for a future exhibit at the GZG. Casts from the
2004–2005 excavation campaigns are stored at the NLMH.
Other original material from this locality, already found
before systematic excavation at the site began, is on
exhibition at the nearby Dinosaurier-Park Mu¨nchehagen
open-air museum (locality 2) and was partly figured by
Lockley et al. (2004). Since 2010, newly excavated at
locality 1, excellently preserved theropod and ornithopod
trackways from the lower track horizon are reassembled
Fig. 11 Mu¨nchehagen/F. Wesling GmbH quarry (locality 1), upper
track-horizon (large ornithopod trackways) during excavation in
2005. Note the long trackway on the left side and well-preserved
oscillation ripple-marks. After documentation, this area was lost to
quarry operations; view towards N
Vertebrate tracksites in the Obernkirchen Sandstone 249
123
and permanently exhibited in the Dinosaurier-Park Mu¨n-
chehagen protection hall, including currently (spring 2011)
a [15 m long consecutive trackway of a medium-sized
theropod.
References Athen et al. (2005); Wings et al. (2005a, b);
Lehmann (2006); Lehmann et al. (2006); A. Richter et al.
(2007); U. Richter et al. (2007); Richter (2007).
2. Mu¨nchehagen/Dinosaurier-Park Mu¨nchehagen open-air
museum (Figs. 6, 10)
Coordinates N 5226033.2000, E 912002.5100; R: 3513720,
H: 5812190; 1:25000 German Topographical Map refer-
ence grid sheet 3521 Rehburg.
Stratigraphic position Lower part of Obernkirchen
Sandstone, ca 1 m above the Hauptflo¨z coal seam.
Track inventory Sauropods, unidentified tridactyl, bipe-
dal dinosaur, ?crocodile swimming-track.
Remarks The outcrop exposes 7–8 m of thin- to thick-
bedded sandstone, interbedded with cm-thick mudstone to
siltstone. The Hauptflo¨z coal seam was found in a core well
ca 1 m below the sandstone layer forming the floor of the
quarry (Fischer 1998).
256 individual footprints belong to eight sauropod
trackways, seven of which are oriented roughly subparallel.
19 additional footprints form a single trackway of a large
tridactyl biped on a single bedding surface. All tracks were
pre-diagenetically eroded and further worn by postexca-
vational damage on the quarry floor before protection.
Because of the poor preservation, the producer of the tri-
dactyl track cannot be determined (Lockley et al. 2004).
Diedrich (2004) reported ‘‘crocodile scratch-marks’’ from
this locality but gave no further details or figure.
On discovery of these tracks in 1979, this tracksite
enjoyed great public interest, because it was the first time
since the 1920s that long and numerous in situ trackways of
dinosaurs had been revealed. It was also the first time
sauropod tracks, apparently left by a ‘herd’, had been
identified from the Obernkirchen Sandstone (for overviews
see Hendricks 1981; Probst and Windolf 1993; Fischer and
Thies 1993, 2000; Fischer 1998). The tracks are protected
by conservation law and form the centrepiece of the
Dinosaurier-Park Mu¨nchehagen open-air museum. Some
areas of the tracksite are sheltered and publicly accessible
(see also To¨nebo¨hn and Kulle-Battermann 1989a, b, c).
The tracks have been studied extensively by Hendricks
(1981), Kleinschmidt (1986), Fischer (1998) and Lockley
et al. (2004). Production in the quarry ceased in 1972.
References Hulke (1980); Hendricks (1981); Staesche
(1981); Look (1986); Look et al. (1988); Kleinschmidt
(1986); Fischer (1987, 1998); Meyer (1987); Fischer et al.
(1988); To¨nebo¨hn and Kulle-Battermann (1989a, b, c),
Fischer in Boenigk (1990); Probst and Windolf (1993);
Fischer and Thies (1993, 2000); Haubold in Lockley
(1993); Wilde et al. (1995); Mutterlose (1997b); Lockley
and Meyer (2000); Lockley et al. (2004); Wright (2005);
U. Richter (2007).
3. Mu¨nchehagen/Stadtla¨nder quarry (Fig. 6)
Coordinates N 5226032.5700, E 911051.3700; R: 3513540,
H: 5812160; 1:25000 German Topographical Map refer-
ence grid sheet 3521 Rehburg.
Stratigraphic position Lower part of Obernkirchen
Sandstone, ca 4 m above Hauptflo¨z coal seam.
Track inventory Large ornithopods(?).
Remarks The quarry exposed ca 9 m of medium- to
thick-bedded sandstone, interbedded infrequently by cm-
thick mudstone layers. Layers with accumulated shark-
teeth, preserved as impressions, have been reported from
this site.
Large tridactyl hypichnia, ‘‘probably made by Iguan-
odon’’, were observed by Wilde et al. (1995) and Pelzer
(1998) at the basal horizon (corresponding to the track
horizon in locality 2) and in a second horizon ca 3 m above
the base of the outcrop. The quarry is abandoned, over-
grown by vegetation, and partly water-filled.
References Wilde et al. (1995); Pelzer (1998).
4. Wo¨lpinghausen/Wo¨lpingha¨user Berg (Figs. 1, 12)
Coordinates Wo¨lpingha¨user Berg (Wilhelmsturm),
approx. N 5225044.2200, E 913012.2800; R: 3515120, H:
5810660; 1:25000 German Topographical Map reference
grid sheet 3521 Rehburg.s
Stratigraphic position Obernkirchen Sandstone, exact
level unknown.
Track inventory Large and medium-sized ornithopods,
theropods.
Remarks The first scientifically described record of any
dinosaur track from Germany was reported from this
locality (Struckmann 1880a, b; Fig. 1). Struckmann
reported the presence of ca 40 tridactyl, ‘‘bird-like’’ tracks
Fig. 12 Wo¨lpinghausen (locality 4). Sandstone block with numerous
ornithopod and theropod tracks (hypichnial casts, GZG.IF.00100).
Size: ca 178 9 150 cm. On exhibition in the GZG Geopark,
Go¨ttingen (donation of the Principality of Schaumburg-Lippe, second
half, nineteenth century)
c
250 J. J. Hornung et al.
123
Vertebrate tracksites in the Obernkirchen Sandstone 251
123
of a biped at the quarry of A. Spo¨rl, close to the Wil-
helmsturm, ca 1.5 km NW to the town of Wo¨lpinghausen.
For some of them the footlength was ca 60 cm. Struck-
mann (1880a, b) provided depictions of isolated hypichnial
casts corresponding to large ornithopod tracks, which he
associated with the genus Iguanodon (Fig. 1b).
Unfortunately, geological information about the tracksite
provided by Struckmann is very sparse. He noted the foot-
prints from two 40–60 cm thick sandstone beds near the base
of the quarry, exposing ca 6 m of Obernkirchen Sandstone.
Today the quarry is abandoned and inaccessible.
The GZG houses a large sandstone slab from ‘‘Wo¨l-
pinghausen’’ (donation of the Principality of Schaumburg-
Lippe; mentioned in Schmidt 1959), lacking more precise
locality information. The lower surface is densely covered
with hypichnia of mid-sized to large ornithopods and
theropods, heading in various directions (Fig. 12).
References Struckmann (1880a, b); Schmidt (1959).
Localities 5 and 6: Bu¨ckeburg/Harrl hill
5. Bu¨ckeburg/Abandoned quarry on the W’ side of Harrl
hill (Fig. 7d)
Coordinates N 5215003.7000, E 903050.7100; R:
35044670 H: 5790870; 1:25000 German Topographical
Map reference grid sheet 3720 Bu¨ckeburg, ca 1.4 km SE of
Bu¨ckeburg.
Stratigraphic position Upper part of Obernkirchen
Sandstone.
Track inventory Tridactyl, bipedal dinosaurs.
Remarks The quarry exposed a ca 7.5 m thick succession
showing a tripartite subdivision (Pelzer 1998; Fig. 7d). The
lower, [4.7 m thick succession consists of medium-bed-
ded, scoured, and partly cross-stratified sandstone with
root-traces in the upper part, overlain concordantly with ca
0.8 m of sandy siltstone and mudstone with ripple-mark
cross-lamination and root horizons, topped by a thin coal-
seam. The upper, ca 2 m thick unit consists of thin to
medium-bedded, cross-stratified, scoured, fine-grained
sandstone, rich in molluscs (Neomiodontidae and uniden-
tified gastropods), beginning with a basal coarse-grained
horizon with intraformational rip-up clasts, wood impres-
sions, and abundant large bivalve shallow-burrowing traces
(Lockeia isp., personal observation).
Pelzer (1998) reported cross-sections of hypichnia in the
quarry wall, near the base of the second unit, ca 1.3 m
below the coal seam (Fig. 7d). The track-bearing horizon is
probably equivalent to the track layer in nearby locality 6,
excavated by Ballerstedt (e.g. 1905, 1921b). The quarry is
abandoned and over-grown by vegetation, but still acces-
sible (personal observation 08/2009).
References Pelzer (1998).
6. Bu¨ckeburg/Hexenteich quarry
Coordinates N 5215001.1600, E 904007.6200; R: 3504770
H: 5790734; 1:25000 German Topographical Map refer-
ence grid sheet 3720 Bu¨ckeburg, ca 1.6 km SE of
Bu¨ckeburg.
Stratigraphic position Upper part of Obernkirchen
Sandstone.
Track inventory Large ornithopods (including Struthopus
schaumburgensis Ballerstedt, 1921), theropods (‘Buecke-
burgichnus’ maximus Kuhn, 1958), ankylosaurs (Metatet-
rapous valdensis Nopcsa, 1923), turtle footprint.
Remarks During the 19th century, this locality has yiel-
ded important body fossils, for example the holotypes of
the crocodylian Pholidosaurus schaumburgensis and the
dinosaur Stenopelix valdensis. First discoveries of tracks in
the Harrl hill area date back to Grabbe (1881, 1883), but
these lack detailed locality information. Around 1900, an
extensive track horizon was discovered by Ballerstedt
(1905). The material includes large ornithopods and large
theropods, among them the type material of ‘Bueckeburg-
ichnus’ maximus Kuhn, 1958 and Struthopus schaum-
burgensis Ballerstedt, 1921. Isolated hypichinia from this
track horizon survived in the Ballerstedt collection at the
GZG and in other places.
In 1921, Ballerstedt reported a trackway of a quadruped
from the same locality and horizon (the first definite evi-
dence of a quadruped known from the Obernkirchen
Sandstone at the time). This trackway is, historically, the
earliest named trackway attributed to an ankylosaur
(Metatetrapous valdensis Nopcsa 1923; Hornung et al.
2007).
The strata exposed in this outcrop follow the strike
direction of the strata from locality 5, which is located only
350 m to the W. The beds can be partly correlated between
the two outcrops. The lower part of the exposed section
consists of[8 m of scoured, medium-bedded, partly cross-
stratified, fine- to medium-grained sandstone, overlain by
ca 0.6 m of thin-bedded sandstone to mudstone, capped by
a cm-thick coal seam. The coal is truncated erosively and
overlain by ca 1.7 m of thin- to medium-bedded, fining-
upward, cross-stratified sandstone that forms very gently
dipping foresets. Above the sandstone, ca 1 m of siltstone
to mudstone is followed by sandstone 0.5 m thick.
According to Grupe (1933), the succession is underlain
by another thin coal seam and the base of the sandstone
252 J. J. Hornung et al.
123
above the coal-seam is rich in stem-fragments of the pte-
ridophyte Tempskya schimperi Corda, 1845 and contains
rip-up coal clasts. The dinosaur tracks were confined to the
section below the upper coal seam. The central section,
including the coal seam and the underlying fine-grained
deposits, correlates with the exposure at locality 5. The
lithology of the track casts preserved at the GZG suggests that
they were found in this subunit below the coal seam, from
were Pelzer (1998) also reported track casts at locality 5.
The quarry near the Bu¨ckeburg Castle was owned by the
Count of Schaumburg-Lippe in the 19th century. Therefore
it was cited in older literature as ‘‘Fu¨rstlicher Steinbruch’’
(‘‘comital quarry’’) or simply as ‘‘the large quarry on the
Harrl (hill)’’ (von Meyer 1841, 1857; Ballerstedt 1905,
1914, 1921a, 1921b, 1922). Regular operations in the
quarry ceased around 1870, and it was exploited only
occasionally (on demand) into the first half of the 20th
century (Ballerstedt 1921). Today the quarry is partly over-
grown by vegetation and occupied by the ‘Hexenteich’
(‘witch pond’) but is still accessible (personal observation
08/2009).
References von Meyer (1841, 1857, 1859); Ballerstedt
(1905, 1914, 1921a, b, 1922); Wegner (1913, 1926); Grupe
(1933).
Localities 7 to 11: Bu¨ckeberge
7. Obernkirchen/Schauenstein glass factory quarry
(Figs. 13, 14)
Coordinates N 5215037.9300, E 908036.8000; R: 3509887
H: 5791918, 1:25000; German Topographical Map
reference grid sheet 3720 Bu¨ckeburg, ca 1.5 km SE of
Obernkirchen.
Stratigraphic position Upper part of Obernkirchen
Sandstone.
Track inventory Large ornithopods (tridactyl and tetra-
dactyl(?) tracks).
Remarks Dietrich (1927) gave a short description of the
locality as of June 1926 (Fig. 13): The succession shows a
subdivision into two superimposed lithofacies. The upper,
track-bearing lithofacies consists of thin- to medium-bed-
ded, laterally irregular-continuous or lenticular, partly
cross-stratified sandstone, intercalated with thin-bedded
sandy mudstone. This succession was underlain by thick-
bedded sandstone and overlain by a thin, discontinuous
coal-seam. Dietrich (1927) reported a thickness of 12 m,
implying that this thickness referred to the upper, cross-
stratified horizon. However, comparison with other data
from surrounding outcrops and the schematic log by Grupe
(1933: fig. 1), suggest that this corresponded rather to the
total exposed thickness in the quarry. The cross-stratified
lithofacies (below the coal-seam) therefore accounts for
[3–4 m and the underlying thick-bedded facies for ca
8–9 m, of which only the upper ca 5 m were exposed in the
quarry. The underlying strata down to the UHS were
identified from a well in the quarry area (Grupe 1933).
The quarry exposed at least three track horizons, one of
which (from the upper lithological unit) was mapped in
detail by Dietrich (1927). He identified approximately
40–45 individual tridactyl tracks but was unable to identify
longer trackways of consecutive footsteps (Fig. 12). Hamm
Fig. 13 Obernkirchen/
Schauenstein glass factory
quarry (locality 7). Uppermost
track horizon as mapped by
Dietrich 1927: fig. 1. A,
B Covered areas of the track
horizon; C track surface,
sandstone with local oscillation
ripple-marks; vertically striated
areas were covered by some cm
of mudstone, thinning in the
stippled areas; a–e single
trackway of a large ornithopod,
f, g various other tridactyl tracks
Vertebrate tracksites in the Obernkirchen Sandstone 253
123
(1957) published a contemporary photograph, apparently
showing a part of Dietrich’s track horizon, and some
measurements (Fig. 14).
The tracks were identified as having been produced by a
single taxon of large ornithopods but their shape and
quality strongly depends on the local thickness of the sandy
mud in which the tracks were left. In some areas they were
impressed through the mud-layer into relatively clean sand,
where only subtle marks were left. From isolated hypich-
nia, Dietrich (1927) also figured an allegedly tetradactyl
ornithopod pes print, which was named Wealdenichnites
iguanodontoides by Kuhn (1958; Fig. 9d). The existence of
this morphotype was questioned by Haubold (1974, 1984).
Based upon the vertical succession of lithofacies, the
track-bearing section can be correlated with the upper part
of the exposure in the nearby active quarry of the Obe-
rnkirchener Sandstein (locality 8, Fig. 5). The Schauen-
stein quarry was operated by the F.C.H. Heye glass
manufacturing company from 1827. It is now abandoned
and re-filled as an earth dump.
References Dietrich (1927); Grupe (1933); Hamm
(1957).
8. Obernkirchen/Obernkirchener Sandsteinbru¨che GmbH
quarry (Figs. 7b, c, 15, 16)
Coordinates Main site: N 5215045.0200, E 912024.1600;
R: 3514120 H: 5792220; nearby small, inactive quarry: N
52 15037.2200, E 912000.3400; R: 3513720, H: 5791940;
1:25000 German Topographical Map reference grid sheet
3721 Auetal, ca 5.3 km ESE of Obernkirchen.
Stratigraphic position Middle to upper part of Obern-
kirchen Sandstone.
Track inventory Small, bipedal ornithischians, several
morphotypes of large ornithopods and theropods
(‘Bueckeburgichnus’ maximus, ‘Megalosauripus’ isp., and
didactyl tracks).
Remarks The active quarry of the Obernkirchener
Sandsteinbru¨che GmbH is currently the most important
production site for the Obernkirchen Sandstone. The
exposed section comprises more than 8 m and can be
subdived into two vertically stacked lithofacies units
(Fig. 7b). The lower, at least 6 m thick, unit consists of
massive, locally channellised (Fig. 7c), thick bedded
sandstone with irregular bedding surfaces, abundant rooted
palaeosol horizons, and few cm-thick mudstone intervals.
The upper unit comprises more than 2 m of scoured or
channellised, lenticular, or planar-bedded, laterally dis-
continuous sandstone, interbedded with mm- to cm-thick
mudstone layers.
At least six to seven track horizons are present: five to
six in the lower lithofacies unit and one in the upper. The
lowermost track level, ca 1.5–2 m above the base of the
exposure, comprises hundreds of tri- and didactyl theropod
and tridactyl ornithopod tracks of different sizes and
morphologies (Figs. 7b, 16a), some still showing long,
superimposed trackways. This horizon, which was
Fig. 14 Obernkirchen/
Schauenstein glass factory
quarry (locality 7). Large
ornithopod trackway, foot
length ca 35 cm, stride length ca
140 cm. Photographed in 1925,
measurements from Hamm
1957
254 J. J. Hornung et al.
123
nicknamed ‘chicken yard’ during excavation (Richter et al.
2009; van der Lubbe et al. 2009), can be traced throughout
wide areas in the northern, deepest part of the quarry. Circa
1 m above the ‘‘chicken yard’’ level, another track horizon,
with large ornithopod tracks, has been recognised in a few
places.
The uppermost track horizon within the lower unit at the
active quarry contains large ornithopod tracks of a new
morphotype yet to be described in detail (Bo¨hme et al.
2009). The stratigraphically youngest track level at
Obernkirchen is located within the upper unit. Finally, at
least one horizon with ornithopod tracks has recently been
found in the southern part of the active quarry. The exact
stratigraphic position of this horizon has not yet been
established, because this area is affected by some faulting,
but it seems to be situated in the lower unit.
The track levels reported by Diedrich (2004) are not
exposed in the main quarry but in a small, currently inac-
tive quarry ca 500 m to the SW. We suppose that the
bipartite lithofacies subdivision observed by Diedrich
(2004: figs. 2–3) at this site corresponds to the lower and
upper lithofacies units in the main quarry. Only his ‘‘track
bed I’’ was found at the site, as indicated in Diedrich
(2004), ca 2.5 m below the top of the lower lithological
unit. A second track layer occurs ca 1 m lower in the
succession than indicated in Diedrich (2004: fig. 3) and
nearly 2 m below the position indicated in Diedrich (2004:
fig. 2). Diedrich’s ‘‘track bed II’’ could not be found in any
of the positions given. In the same locality, isolated large
ornithopod tracks were found at the top bed of the lower
unit, indicating a third track level.
The gross lithostratigraphic succession in the Obe-
rnkirchener Sandsteinbru¨che GmbH quarry is very similar
to that in the Schauenstein glass factory quarry (locality 7,
Dietrich 1927; Grupe 1933), and the boundary between the
lower and upper subunits may correlate. Unfortunately, no
coal seam is exposed in the former in direct succession
with the track horizons. However, a 10–30 cm thick seam
(probably the Hauptflo¨z seam) is tectonically uplifted in the
southeastern area of the quarry in relation to the main part,
suggesting that it underlies the lower facies unit. The
presence of the Hauptflo¨z coal seam ca 11 m below the
ground level (ca 2–4 m below the quarry base level) is also
confirmed by various wells drilled in the vicinity of the
quarry (LBEG 2009).
In recent years, especially since 2006, the dinosaur track
inventory (Fig. 16b–e) of this location has been shown to
be of enormous richness in quantity and diversity. U. Leh-
mann (1978) provided the first extensive description of
ornithopod tracks from the Obernkirchen Sandstone since
Ballerstedt’s era based upon a large track slab from this
locality (now at the GPMH). Diedrich (2004) described a
large track slab with ornithopod and theropod (‘Megalo-
sauripus’ isp.) tracks, discovered in 1987. Haderer and
Neef (2004) reported new material of ‘Bueckeburgichnus’
maximus from this locality, for the first time since the
Fig. 15 Locality map of the
Obernkirchener
Sandsteinbru¨che GmbH quarry
(locality 8) figured by Diedrich
(2004: figs. 2-3) is situated in
the inactive quarry to the SW.
Most other discoveries,
including the excavations since
2006, were made in the large,
active quarry. A map legend is
given in Fig. 6a
Vertebrate tracksites in the Obernkirchen Sandstone 255
123
Fig. 16 Obernkirchen/
Obernkirchener
Sandsteinbru¨che GmbH quarry
(locality 8). The ‘chicken yard’
track horizon as excavated in
2008–2009. a Overview of the
bedding surface with hundreds
of dinosaur footprints, preserved
as epichnial impressions;
b–e Examples of dinosaur
footprint morphotypes
preserved on the ‘chicken yard’
track surface; b cf.
‘Megalosauripus’ isp.,
moderately sized, tridactyl
theropod pes track. Light shed
from the right; c Small, didactyl
theropod pes track, probably
from a troodontid. Light shed
from the right; d Small, biped
ornithischian pes track. Light
shed from the lower right;
e Large ornithopod track,
manus/pes-couple. Light shed
from the left. Coin diameter in
all figures b–e is 24 mm
256 J. J. Hornung et al.
123
discovery of Ballerstedt’s (1905) original material found
near Bu¨ckeburg. However, the specimen, kept at the
NLMH, is very poorly preserved, and its theropod assign-
ment is disputable (personal observation, TvdL, AR). Iso-
lated large slabs at the quarry dump area also preserved
large theropod imprints referable to ‘Bueckeburgichnus’
(personal observation, JJH, MR).
Systematic excavations of the NLMH since 2006 have
continuously yielded new finds, among others spectacular
mass occurrences of theropod tracks, including didactyl
footprints, presumably of troodontids (van der Lubbe et al.
2009). At least four to six ichnotaxa occur at Obernkirchen,
partially in the same track layers, therefore providing the
highest known diversity of any tracksite in the Obernkir-
chen Sandstone. Large ornithopods are represented by
several trackways (Bo¨hme et al. 2009) including those that
indicate quadrupedal individuals (adult and subadult).
A proposal for the partial conservation and public
exhibition of this site is under development.
References Broschinski (2004); Diedrich (2004); Haderer
and Neef (2004); Lehmann (2003, 2006); Richter and
Stratmann (2007); Bo¨hme et al. (2009); van der Lubbe
et al. (2009); Richter et al. (2009).
9. Niensta¨dt/‘‘Dreier’s quarry’’
Coordinates Exact locality unknown. Probably SE to the
town of Niensta¨dt, 1:25000 German Topographical Map
reference grid sheet 3721 Auetal.
Stratigraphic position Unknown position within Obern-
kirchen Sandstone.
Track inventory Tridactyl, bipedal dinosaurs.
Remarks Grabbe (1881) reported moderately sized
(footlength 32.9–37.9 cm) apparently tridactyl tracks of a
biped of ‘‘distinct habitus’’. They were characterised by a
‘‘very broad ‘toe base’’’ and ‘‘triangular toes’’ of which the
middle one was the shortest. Although he provided some
measurements, his description–lacking an illustration–is
not sufficient to gain a clear picture of this type of tracks.
The allegedly short digit III is incongruent to mesaxonic,
tridactyl dinosaur pes anatomy and suggests either misin-
terpretation or an epigenetic modification of the track.
The small, now probably vanished quarry could no
longer be located with certainty but was situated SE of the
vicinity of Niensta¨dt.
References Grabbe (1881).
10. Niensta¨dt: Liekwegen/Brandshof quarry (Fig. 17)
Coordinates Approx. N 5217006.5000, E 911025.7000;
R: 3512950 H: 5794670; 1:25000 German Topographical
Map reference grid sheet 3721 Auetal, ca 1.8 km SE of
Niensta¨dt.
Stratigraphic position Unknown position within Obern-
kirchen Sandstone.
Track inventory Large ornithopods.
Remarks Grabbe (1883) reported large ornithopod tracks
(footlength 47–48 cm) from this locality, but did not pro-
vide any details on the geology or stratigraphy of this, now
inaccessible, site. He excavated more that 10 consecutive
footprints forming a trail.
Fig. 17 Liekwegen/Brandshof quarry. a Tridactyl pes track, exact
size unknown. Historical photography, ca 1940-1945, GZG museum
archive; b Present exposure of the quarry. The base of the quarry
consists of sideritic shales with bivalve coquinas, overlain by
medium-bedded sandstone. Height of exposure ca 6–7 m, 2009
Vertebrate tracksites in the Obernkirchen Sandstone 257
123
An unpublished historical photograph from the 1940s
(Fig. 17), labelled ‘‘Liekwegen-Wendthagen quarry’’,
shows a tridactyl track of a biped consisting of two pes
imprints, preserved as an epichnial relief on an irregular
sandstone surface. The morphology and proportions of the
track suggest an ornithopod trackmaker. The photograph
from the Ballerstedt archive was probably taken at the
Brandshof quarry, because of this was the largest quarry in
the area and still in production in the mid-20th century.
Only a small part of the former quarry is still accessible,
exposing ca 4–5 m of thinning-upward, medium to thinly
bedded sandstone. In some places the floor of the quarry
consists of sideritic claystone with neomiodontid coquinas
underlying the sandstone.
References Grabbe (1883).
11. Wendthagen/‘‘Maier’s quarry’’
Coordinates Exact locality unknown. In the vicinity of
the town of Wendthagen, 1:25000 German Topographical
Map reference grid sheet 3721 Auetal.
Stratigraphic position Lower or middle part of Obern-
kirchen Sandstone.
Track inventory Large bipedal dinosaurs.
Remarks Grabbe (1881) recorded an oral report by min-
ing supervisor Heidtmeier that a ca 100 m long trail of
large footprints was found ‘‘well below the top’’ of the
Obernkirchen Sandstone ca 1850–1860. The tracks were
preserved as an epirelief. This is the oldest rather detailed
and reliable account of dinosaur tracks from Germany. The
tracks were no longer preserved at the time of Grabbe’s
investigations.
References Grabbe (1881).
Localities 12 and 13: Osterwald
Tracksites in the Osterwald mountains are located in the
more proximal, fluvial and alluvial deposits of the
Obernkirchen Member (Pelzer et al. 1992; Pelzer 1998).
These deposits cannot be correlated in detail with those
of the Obernkirchen Sandstone sensu strictu or dated
more precisely than middle or late Berriasian. They are
overlain by Valanginian brackish deposits (Osterwald
Member).
12. Osterwald/Quarry 450 m south of Ahrensberg
mountain (Fig. 18)
Coordinates N52 7030.2700, E 9 38058.7000; R: 3544640
H: 5777050; 1:25,000 German Topographical Map
reference grid sheet 3823 Coppenbru¨gge (formerly Eldag-
sen as in Naumann 1927), ca 2 km NE to the town of
Osterwald.
Stratigraphic position Upper(?) Obernkirchen Member.
Track inventory Large ornithopods, theropods.
Remarks Naumann (1927) indicated an unspecified
number of large, tridactyl hypichnia at the base of a 2.1 m
thick sandstone layer interbedded within a ca 23 m thick series
of alternating sandstone, mudstone, and claystone rich in plant
macrofossils and minor coal seams. The succession is located
well below the boundary to the Osterwald Member and
probably slightly older than that in nearby locality 13. The
track horizons in both localities cannot be correlated litho-
stratigraphically. The lithofacies and flora suggest a peri-flu-
vial environment with fine-grained sediments and coal settled
in oxbow-lakes and back-swamps, while coarser-grained
sedimentary bodies represent channel-fills, natural levees, and
crevasse-splay fans (compare Pelzer 1984).
Naumann (1927: 29) explicitly associated the tracks with
Iguanodon and reported the recovery of two individual foot-
prints, one with a foot-length of 52 cm and a foot-width of
42 cm. The quarry is abandoned and over-grown by vegetation.
A ca 2.10 m 9 1.62 m large slab (Fig. 18) of thinly
bedded, fine-grained, bioturbated, current ripple-marked
sandstone was recovered from the Osterwald/Ahrensberg
area (exact provenance unknown) during the 1970s and is
now kept at the Bundesanstalt fu¨r Geowissenschaften und
Rohstoffe, Hannover (BGR). The slab shows three mid-
sized ornithopod footprints (footlength ca 30 cm), forming
the section of a trackway, and a single well-preserved left
footprint of a mid-sized theropod (footlength ca 31 cm).
The tracks are preserved as epichnia. The small fragment
of a second (right) theropod footprint is probably preserved
at the upper margin of the slab.
References Naumann (1927).
13. Osterwald/Kaiserblick (Fig. 19)
Coordinates N52 7038.0800, E 9 39016.0000; R: 3544950
H: 5777260; 1:25000 German Topographical Map refer-
ence grid sheet 3823 Coppenbru¨gge (formerly Eldagsen as
Fig. 18 Osterwald/Ahrensberg area. Bu¨ckeberg Formation, fluvial
equivalent of Obernkirchen Member. a Track slab with two,
overstepping, large ornithopod trackways (lower part) and theropod
footprint (upper right). Parts of a second theropod footprint are
located at the upper margin of the slab. Epichnial relief, low-angle
light from upper right slab-size approximately 210 cm 9 162 cm,
BGR unnumbered; b Right and left ornithopod footprints, the
trackmakers walked in opposite directions, detail of a; c Left
theropod footprint, detail of a. Scale bar 30 cm (in a), 10 cm (in
b and c)
c
258 J. J. Hornung et al.
123
in Naumann 1927), ca 2.4 km NE to the town of
Osterwald.
Stratigraphic position Upper(?) Obernkirchen Member.
Track inventory Large tridactyl tracks (?Ornithopoda).
Remarks From an abandoned quarry, ca 500 m ESE of
the Ahrensberg, Naumann (1927) noted ‘‘Iguanodon-
tracks’’ near the base of a sandstone package, embedded in
a ca 20 m thick succession of sandstone, mudstone, and
claystone, and minor coal seams. It is probably slightly
younger than that in nearby locality 12. The lithofacies data
provided by Naumann are scarcer than in locality 12 but
suggest a similar environmental setting. The quarry is
abandoned and over-grown by vegetation.
Barner (1931) published a photograph from the track-
site, taken around 1925 (Fig. 19).
References Naumann (1927); Barner (1931).
The Obernkirchen Sandstone dinosaur track localities-
a megatracksite?
Diedrich (2004) listed several of the above mentioned
dinosaur tracksites (localities 2, 4, 6, 8) as forming a
‘‘central European Berriasian megatracksite’’. This argu-
ment was mainly based on the (rough) stratigraphical and
lithofacial correlation of these localities and the assumed
lateral connection of the track-bearing lithological units.
Generally, a megatracksite is strictly defined as a
regionally extensive, single track-bearing horizon, or a
very thin package of horizons over a large area in the order
of hundreds or thousands of square kilometres (Lockley
and Pittman 1989; Lockley 1991a). Almost all of the
established megatracksites are formed by track-bearing
strata not exceeding 1 m in thickness (Lockley 1997). An
exception is provided by the Dakota Sandstone mega-
tracksite (the ‘‘dinosaur freeway’’, Lockley et al. 1992),
which consists of several track horizons, encompassing a
total thickness of ca 10 m. Regional correlation of the track
horizon(s) requires very sound stratigraphic control, as
provided, for example, by vertical relationships to major
sequence boundaries in clastic sequences or laterally
extensive bed-by-bed correlation in carbonate platform
successions (Lockley 1991b; Meyer 1993; Kvale et al.
2001).
Such tight stratigraphic control is not available for the
Obernkirchen Sandstone tracksites. Although in many
locations several trackway horizons are present, these
cannot be correlated laterally over more than a few square
kilometers at best (Figs. 5, 6b). The sandstone beds on
Vertebrate tracksites in the Obernkirchen Sandstone 259
123
which the tracks were preserved are often laterally dis-
continuous, sometimes even at outcrop scale, and lithofa-
cies assemblages differ among the outcrop regions.
Truncated channel-fill deposits indicate the presence of
intraformational discontinuities above the track-level in the
Mu¨nchehagen quarry (Fischer 1998). A discontinuity at
regional scale was also reported from the western Obern-
kirchen area by Dietrich (1927) and Grupe (1933) and
observed in the Obernkirchener Sandsteinbru¨che GmbH
quarries (personal observation).
As already discussed, major differences exist between
the lithofacies assemblages at the Rehburg Mountains and
the Bu¨ckeberge. Although both systems were probably
genetically coupled on a wider time scale, a strict iso-
chrony of the barrier deposits in the northwest and the
deltaic succession in the southwest—and especially of the
dinosaur track horizons within both—cannot be demon-
strated at present. Although similar in facies, again such a
correlation cannot be made between the occurrences at
Harrl hill and the Bu¨ckeberge. Therefore, identification of
the Obernkirchen Sandstone as a megatracksite currently
seems premature and rather uncertain.
Conclusions
13 dinosaur tracksites have been identified in the Obern-
kirchen Sandstone (Obernkirchen Member, Bu¨ckeberg
Formation, Late Berriasian) in the southeastern part of the
Lower Saxony Basin, eight of which are no longer acces-
sible. A survey of available geological and ichnological
data on these localities shows a database of quite inho-
mogeneous quality.
Correlations to current facies models (Pelzer 1998) for
the Obernkirchen Sandstone and equivalents show that
eleven tracksites are located in a distal coastal barrier/
deltaic facies assemblage.
Fig. 19 Osterwald/Kaiserblick (locality 13). Large ornithopod track (hypichnial cast). Historical photograph, 1930 by F. Hamm (from Barner
1931)
260 J. J. Hornung et al.
123
The presence of dinosaur tracks spans a number of
facies types, indicating a range of local environments.
Track occurrences in the Mu¨nchehagen area are linked to
heterolithic successions, reflecting re-occurring high-
energy depositional events interrupting long-term calm
suspension settling. These events have a local reoccurrence
frequency of at least 101 to 102 yrs, and are related to storm
events carrying matured littoral barrier sands into back-
barrier lagoons. Dinosaurs must have frequently passed
through shallow areas of these lagoons and along barriers
and shoals.
At Obernkirchen (Bu¨ckeberge), many of the track levels
occur at the top of sandbars, which show indications of
subaerial exposure and plant coverage. The laterally con-
tinuous beds are vertically and laterally intercalated with
channel-fill sandstones in places, suggesting a net aggrad-
ing, temporarily emergent mouthbar complex. At Harrl hill,
well-preserved dinosaur tracks occur on what is interpreted
as a submerging mouthbar surface.
Because of association of various facies assemblages,
lateral inconsistency, and a lack of stratigraphic correlation
of the track-bearing horizons in the Obernkirchen Sand-
stone, its characterisation as a megatracksite (Diedrich
2004) is currently regarded as speculative.
Two tracksites occur at the Osterwald mountains in
more proximal equivalents of the Obernkirchen Sand-
stone, represented by perifluvial overbank deposits further
inland.
Vertebrate clades represented by tracks include Orni-
thopoda, small, bipedal ornithischians, Ankylosauria,
Theropoda, Sauropoda, turtles, and possibly crocodiles.
Large ornithopods are most abundant and probably present
in all localities, followed in abundance by theropods (five
localities), sauropods, ankylosaurs, turtles and crocodiles
(one locality each). Several ichnotaxa have been estab-
lished for these tracks, but, with the exception of Meta-
tetrapous valdensis Nopcsa, 1923 (Ankylosauria; Hornung
et al. 2007), none is properly defined at present. There are
two or three morphotypes (partly related to different
ontogenetic stages) of large ornithopod tracks and at least
three morphotypes of theropods (corresponding to different
clades). Ornithopod and theropod tracks occur in large
numbers in several localities.
A strong discrepancy in diversity and composition exists
between the vertebrate fauna represented by tracks and by
body fossils from the same lithological unit. This suggests
high local vicariance and taphonomical control of both
body and trace fossil preservation. The dinosaur tracks
represent a local fauna at least temporarily autochthonous
to the estuary and shoal system. The nearly total absence of
articulated or associated material from these groups sug-
gests that their presence in this environment was probably
non-permanent and migratorial.
Acknowledgments The following people provided valuable dis-
cussions and logistic help: H. Schwanke (GZG), M. Sosnitza (retired,
formerly GZG), T. Stegemann (GZG), H.-V. Karl (Weimar/GZG), J.
Ko¨nig (GZG), M. Raddatz (Halle/S.), and U. Staesche (Hannover).
T. Wiese (BGR) is thanked for providing access to material in his
care. V. J. Roden (formerly GZG) is acknowledged for checking the
English spelling and grammar. The current excavation projects in the
Obernkirchen Sandstone would not be possible without ongoing and
generous support by the owners and crew of the quarry operations.
We cordially thank F. Wesling sen. (Ferdinand Wesling GmbH & Co.
KG and Dinosaurier-Park Mu¨nchehagen GmbH & Co. KG, Rehburg-
Loccum), K. Ko¨ster (Obernkirchener Sandsteinbru¨che GmbH,
Obernkirchen), and B. Wolter (retired, formerly Dinosaurier-Park
Mu¨nchehagen) for their cooperation in our research work, and for
their willingness to allow public access to and preservation of the
dinosaur tracks found on their property. F. Klebe and L. Seidel
(Landkreis Schaumburg-Lippe) also provided interest in and support
of conservation of the tracks. The research at Mu¨nchehagen is
strongly assisted by the team of the ‘‘Dinosaurier-Park Mu¨ncheha-
gen’’ open-air museum headed by N. Kno¨tschke. The support by
many others was vital to the excavation projects, including M. Sch-
mitz (formerly NLMH, currently Naturhistorisches Museum, Mainz),
A. Basse (Bremen), and U. Richter (Hannover), and the volunteer
team, joined on a regular basis by U. Stratmann, O. Gerke, U. Picht,
G. Picht, J. Picht, C. Weimann, and L. Weimann and hundreds of
short-term volunteers who have spent thousands of working-hours
uncovering the tracksites at Mu¨nchehagen and Obernkirchen since
2004. Colleagues from other research institutions also cooperated in
various stages of the project, including M. Wiggenhagen (Institut fu¨r
Photogrammetrie und Geoinformation, Leibniz-Universita¨t Han-
nover, Hannover), M. Fischmann, and K. Grobe (Fachbereich Design
und Medien, Fachhochschule Hannover, Hannover), O. Wings
(Museum fu¨r Naturkunde, Humboldt-Universita¨t zu Berlin), J. Lepper
(formerly BGR), C. Heunisch, and F. Luppold (Landesamt fu¨r
Bergbau, Energie und Geologie, Hannover), J. Lehmann (Geowis-
senschaftliche Sammlung, Universita¨t Bremen), and O. Kullmer
(Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am
Main). All of these people are warmly acknowledged for their efforts.
We also thank Richard J. Butler (Mu¨nchen) and Oliver Wings
(Mu¨nchehagen/Berlin) for reviewing the manuscript and providing
helpful comments to improve the paper.
Open Access This article is distributed under the terms of the
Creative Commons Attribution License which permits any use, dis-
tribution, and reproduction in any medium, provided the original
author(s) and the source are credited.
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