Received 10 June 2016
Received in revised form 27 September
Accepted 5 October 2016
Available online xxx
The Oligocene was a period of profound climatic and biotic changes, coinciding with a shift from a mostly ice-free
warmhouse world at the Eocene-Oligocene boundary to a globally cooler, more seasonal climate. The Rauenberg locality
(Baden-Württemberg, Germany) is one of the most significant early Oligocene fossil assemblages in Europe, containing
both marine and terrestrial elements of fauna and flora. Preservation is often superb and comprises complete and articu-
lated skeletons with soft tissue preservation. The diverse assemblage provides critical insights into the paleoenvironment
of the Upper Rhine Valley. We reassess diversity at the locality, resulting in a list of 302 taxa found at the site. 207 of
302 are marine (52% of taxa represented by macrofossils), the rest are interpreted as originating from the coastal envi-
ronment. Molluscan, echinoderm, and plant macrofossil diversity are assessed here for the first time. Based on these new
results, we interpret Rauenberg as representing a fully marine assemblage deposited in a moderately shallow, low-energy
tropical-subtropical environment. Productivity was high, and seafloor anoxia was intermittently developed. There is no
evidence for long-term brackish influence or mangrove swamps, and no direct evidence for the development of seagrass
meadows. On land, warm, frost-free conditions permitted the development of prevailingly evergreen broad-leaved mes-
ophytic forests along with pine and palm-rich coastal forests on sandy soils. The marine invertebrate fauna shows more
northerly affinities, whereas the vertebrate fauna is distinctly Paratethyan.
© 2016 Published by Elsevier Ltd.
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx
Contents lists available at ScienceDirect
Palaeogeography, Palaeoclimatology, Palaeoecology
journal homepage: www.elsevier.com
The Rauenberg fossil Lagerstätte (Baden-Württemberg, Germany): A window into
early Oligocene marine and coastal ecosystems of Central Europe
Erin E. Maxwell a, ⁎, Stefanie Alexander b, Günter Bechly a, Kristina Eck c, Eberhard Frey c, Kirsten Grimm d,
Johanna Kovar-Eder a, Gerald Mayr e, Norbert Micklich f, Michael Rasser a, Anita Roth-Nebelsick a,
Rodrigo B. Salvador a, g, Rainer R. Schoch a, Günter Schweigert a, Wolfgang Stinnesbeck h,
Karin Wolf-Schwenninger a, Reinhard Ziegler a
aStaatliches Museum f
r Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany
bKarlsruhe Institute of Technology, Germany
cStaatliches Museum f
r Naturkunde Karlsruhe, Germany
dNaturhistorisches Museum Mainz/Landesammlung f
r Naturkunde Rheinland-Pfalz, Reichklarastr.10, 55116 Mainz, Germany
eForschungsinstitut Senckenberg, Senckenberganlage 25, 60325 Frankfurt am Main, Germany
fNaturgeschichtliche Abteilung, Hessisches Landesmuseum Darmstadt, Friedensplatz 1, 64283 Darmstadt, Germany
t, Eberhard Karls Universit
e 10, 72076 T
r Geowissenschaften, Universit
t Heidelberg, Germany
The Oligocene was a period of profound climatic changes. Its be-
ginning, the Eocene-Oligocene boundary, marked the transition from
a mostly ice-free warmhouse world to an interval of globally cooler
⁎⁎ Corresponding author.
Email addresses: firstname.lastname@example.org (E.E. Maxwell); Stefanie.
email@example.com (S. Alexander); firstname.lastname@example.org (G.
Bechly); email@example.com (K. Eck); firstname.lastname@example.org (E.
Frey); email@example.com (K. Grimm); firstname.lastname@example.org (J.
Kovar-Eder); Gerald.Mayr@senckenberg.de (G. Mayr); norbert.micklich@hlmd.
de (N. Micklich); email@example.com (M. Rasser); anita.rothnebelsick@
smns-bw.de (A. Roth-Nebelsick); firstname.lastname@example.org (R.B.
Salvador); email@example.com (R.R. Schoch); guenter.schweigert@
smns-bw.de (G. Schweigert); Wolfgang.Stinnesbeck@geow.uni-heidelberg.de
(W. Stinnesbeck); firstname.lastname@example.org (K.
Wolf-Schwenninger); email@example.com (R. Ziegler)
climate and Antarctic ice-sheet formation (Coxall et al., 2005; Miller
et al., 1998; Pusz et al., 2011; Zachos et al., 2008). Oligocene climate
at middle and higher latitudes particularly differed from conditions
during the Eocene by increasing seasonality, i.e. the development of
cool winter seasons (Roth-Nebelsick et al., 2014). Fossil remains re-
flect this trend. Oligocene fossil plant assemblages, for example, show
increasing invasion of arctotertiary floristic elements (Kvaček and
Walther, 2001), and a turnover occurred within the terrestrial verte-
brate community (“Grande Coupure”) (Delfino et al., 2003; Stehlin,
The clay pit “Unterfeld”at Rauenberg (Baden-Württemberg, Ger-
many) contains one of the most significant early Oligocene fossil as-
semblages in Europe, and is an important locality for reconstruct-
ing early Oligocene floral and faunal diversity. Although fossils, es-
pecially those of fishes, have been known for a long time, scien-
tific excavations at the site were intensified after the discovery of the
first Old World fossils of modern-type hummingbirds (Mayr, 2004a).
0031-0182/© 2016 Published by Elsevier Ltd.
2 Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx
Preservation is often superb and comprises complete and articulated
skeletons with soft tissues preserved. The Rauenberg site contains
both marine and terrestrial elements of the fauna and flora. The di-
verse assemblage provides critical insights into the paleoenvironment
of the Upper Rhine Valley and is of particular significance, because
it preserves marine and terrestrial organisms across a broad taxo-
nomic spectrum. As such, the Rauenberg site adds to an understand-
ing of early Oligocene ecosystems in Central Europe. Located at a
slightly lower latitude than its current location (paleolatitude 43.74°N,
12.81°E, Paleobiology Database rotation file: Wright et al. (2013);
current: 49.27°N, 8.67°E), the locality lies in a region potentially more
heavily influenced by sea level dynamics and tectonic evolution than
latitudinally-driven climatic zonation. However, this is controversial
and remains to be rigorously examined in a cohesive framework. In
the past, independent studies have attempted to reconstruct the pale-
oenvironment and sedimentation regime of this locality using various
techniques (e.g., micro- and macropaleontology, stratigraphy and geo-
chemistry), resulting in inconclusive and partially contradictory inter-
pretations. Here, we present new data and synthesize existing results
to generate a more complete picture of a marine and coastal paleoenvi-
ronment in the early Oligocene of Germany. We provide a summary of
the taxonomic diversity at the Rauenberg locality based on reexamina-
tion of previously collected but as-yet undescribed material, and also
reassess the published literature in order to compile an up-to-date list
of taxa from the locality. Some groups, such as plankton, fishes, and
birds, are already well-known from Rauenberg (see Section 5 for de-
tails); others, such as plants and macroinvertebrates, have never been
studied in detail. The synthesis of information from these different
groups provides a wealth of new information for paleoenvironmental
The geology of the Paleogene clay pits on the eastern edge of the
Upper Rhine Graben, mined since the beginning of the 19th century,
has been studied by Wagner-Klett (1919), Weiler (1931, 1966), Sittler
(1967), Doebl (1976), Schweizer (1982), Weiss (1988), Trunkóand
Munk (1998), and Grimm et al. (2002).
2.1. Tectonic setting
The former clay pits of Unterfeld, Frauenweiler-Wiesen, and
Rohrlach (Fig. 1A, B), referred to hereafter as the Rauenberg clay
pits, or simply Rauenberg, are located in the Upper Rhine Graben
(Oberrheingraben: Fig. 1D), a region that underwent complex de-
velopment during the Tertiary. The main rifting phase began in the
Oligocene, with continuing northbound thrust of the Alps and the
Pyrenees (Dürr and Grimm, 2011). Perpendicular to the compression
narrowing the graben in a NNE-SSW direction, there was east-west-
to WNW-ESE-oriented crustal strain and spacious trench formation.
Subsidence of the Upper Rhine Graben took place between two prin-
cipal, approximately parallel, faults; subsidence was more or less con-
stant at 4.5 km, with an opening 4–7 km wide (Dürr and Grimm,
2011). The Mainz, Hanau, and Neuwied Basins began to form during
these main rifting stages.
The main rifting phase ended in the later Rupelian–Chattian. It was
a time of large-scale restructuring in the area of the western Mediter-
ranean, and the main compression shifted south and east (Dürr and
Grimm, 2011). In the late Oligocene, strong subsidence of the central
Heidelberg-Mannheim Trench began, while subsidence of the mar-
ginal plates (Mainz Basin, southern Taunus foothills, Hanau Basin)
stopped and the southern Rhenish Slate Mountains gradually rose.
The Upper Rhine Graben briefly formed a dextral shear zone in
a NE-SW-directed compressive stress field, in which the Heidel-
berg-Mannheim Graben subsided as a right-lateral transtensional
basin (Schumacher, 2002; Villemin and Bergerat, 1987). Today, the
Paleogene of Rot-Malsch, on the eastern edge of the Upper Rhine
Graben, forms a tectonic fold ridge which is subdivided by numerous
step faults (Barth, 1970; Wagner-Klett, 1919).
During the second and third Rupelian transgressions (sensu
Hardenbol et al., 1998), marine to brackish sediments were deposited
in the area of the northern Upper Rhine Graben. These sediments,
described as the Graue Schichtenfolge (Doebl, 1967, 1970) or the
Froidefontaine Subgroup (Grimm et al., 2011; Grimm, 2005), are dif-
ferentiated into clayey basinal facies with pelagic faunal elements and
sandy-gravelly coastal facies (Fig. 1E).
The Froidefontaine Subgroup extends throughout the Upper Rhine
Graben and neighboring areas. It overlies the deposits of the Pechel-
bronn Group or encroaches on pre-Tertiary subsurface, and is overlain
by the colorful, sandy marls of the Niederroedern Formation (Grimm
et al., 2011). In the central southern Upper Rhine Graben, the Froide-
fontaine Subgroup is usually around several hundred meters thick. In
the northern Upper Rhine Graben, the thickness in the central trench
decreases to < 150 m.
The Froidefontaine Subgroup is a marine-brackish sequence con-
sisting of an alternation of bituminous, dark grey to light grey green-
ish mudstones, clay marls and siltstones with locally fine sandstones
which interfinger at the borders with the coastal sands and grav-
els of the Alzey Formation. A lithologic subdivision into the Alzey
Formation (= Meeressand), Bodenheim Formation (= Rupelton with
Foraminiferenmergel, Fischschiefer and Upper Rupelton), Meletta
Beds and Cyrenenmergel is possible.
The deposits of the Bodenheim Formation document the second
Rupelian transgression (Hardenbol et al., 1998) in the Upper Rhine
area. The Bodenheim Formation consists of grey to blackish-brown
finely bedded to laminated clays, calcareous clays, and clay marls,
which can subdivided into the Wallau, Hochberg, and Rosenberg
Members (Grimm et al., 2011). In the central and southern part of
the Upper Rhine Graben, e.g., near Wiesloch, sporadically turbiditic
calcareous sandstone were occasionally deposited. In addition, fine
sand beds, layers with bioturbation, and a layer with concretions (the
so-called “lower geode horizon”of Trunkóand Munk (1998)) are dif-
The section in the clay pit was described in detail by Grimm et al.
(2002) (Fig. 1C). In the Rauenberg clay pits, the Wallau and Hochberg
Members of the Bodenheim Formation are present. The sediments in
the clay pit consist of clay- to siltstones, which vary in consolida-
tion and generally show clear bedding and lamination. The subdivi-
sion of the profile may be caused by changes in particle size, carbon-
ate content, color, or alternation between laminated and unstratified
beds. Lamination is attributable to different causes: most lamination
is caused by changes in particle size due to rhythmic sedimentologi-
cal events, but also infrequently by arrhythmic events. Other laminae,
such as calcareous mud layers or calcareous sand layers (“Kalkhaut”
of Trunkóand Munk (1998)) are common and consist of organic lay-
ers of mass occurrences of calcareous nannoplankton. Trunkóand
Munk (1998) identify two thin horizons with concretions in the pit, de-
scribed as the Lower and Upper Geode horizons. The concretions are
isolated or fused together, consisting of medium grey, weathered yel-
lowish ocher dolomitic mudstone containing fossil remains. The con-
cretions show polychaete burrows and are developed early in diagen-
esis (Grimm et al., 2002).
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx 3
Fig. 1. Locality and stratigraphy. A, Location of Rauenberg, Baden-Württemberg, Germany. B, Location of the clay pits referred to in the text relative to the city of Rauenberg. 1-
Rohrlach, 2- Frauenweiler-Wiesen, 3- Unterfeld. C, Stratigraphic section from the Hochberg Member, Unterfeld clay pit. Parts B and C redrawn from Grimm et al. (2002). D, Early
Oligocene (Rupelian) paleogeography of Europe, showing important Oligocene localities discussed in the text. Map modified from Meulenkamp et al. (2000) based on Spiegel et al.
(2007). E, Lithostratigraphy of the Upper Rhine Graben, modified after Rasser et al. (2008). F, Estimated biostratigraphic age of the Hochberg Member (Fischschiefer); produced
with TSCreator (http://tscreator.org). Abbreviations. A, Allschwil, Switzerland; Be,S, Bechlejovice and Seifhennersdorf; Bu,E-K, Budapest and Eger-Kiseged, Hungary; FS A, Fis-
chschiefer A; FS B, Fischschiefer B; Fl, Flörsheim, Mainz Basin; G, Canton Glarus, Switzerland; Ra, Rauenberg; K, Kundratice; L, Linz, Austria; P, southern Poland; Q, Phosphorites
du Quercy; R, Rupel, Belgium; W, Weisselster Basin, Germany.
4 Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx
The carbonate content in the profile is very irregular, but up to
95–100% originates from calcareous microfossil remains (calcareous
nannoplankton and foraminifera). The dolomite content varies be-
tween 1 and about 87 wt% (Grimm et al., 2002).
The color varies from light grey to dark grey based on minor varia-
tions in organic carbon content (TOC). The TOC content in the exam-
ined clay pit is 4.8–5.4 wt% (uncorrected, Grimm et al. (2002)).
The dark bituminous shales contain abundant pyrite, marcasite,
and other metal sulfides. The majority of the metal sulfides consist
of pyrite in the form of small cubes and blooms. Marcasite occurs in
the form of spheres, aggregates and fillings of burrows (Trunkóand
Munk, 1998). Phosphatic skeletal grains (fish remains, teeth, and co-
prolites) are common and of black-brown color. The clay mineral il-
lite dominates, followed by kaolonite and chlorite. Secondary gypsum
crystals are also present (Weber, 1951).
Among the calcareous nannoplankton, the occurrence of Transver-
sopontis pygmaea indicates an Oligocene age between NP 23 and NP
24 (after Köthe, 1986). The section comprising the Hochberg Mem-
ber is correlated to NP 23 (after Martini, 1971; modified from Müller,
1978): from the last occurrence of Reticulofenestra umbilica to the
first occurrence of Cyclicargolithus abisectus and/or Helicosphaera
recta. Consistent with the calcareous nannoplankton, dinoflagellate
cysts in the sediments can be placed in the Subzone D 14 a (lower
Oligocene, Rupelian). This subzone is marked from the first occur-
rence of Chiropteridium galea to the last occurrence of Enneadocysta
pectiniformis. Corresponding well with the microfossils, the mam-
malian creodont genus Apterodon is associated with MP 22–MP 23
(Frey et al., 2010) (Fig. 1F).
The foraminiferal association is also typical of the Rupelian. Char-
acteristic and frequent forms are Cyclammina placenta,Bathysiphon
tauriensis,Stilostommella ewaldi and Bolivina beyrichi. It is possible
to subdivide the Hochberg Member at Rauenberg into the foram-rich
Fischschiefer A and the foram-poor Fischschiefer B (Grimm, 1991,
1994). Fischschiefer B cannot be further subdivided, however Fis-
chschiefer (= FS) A shows mass occurrences of Bolivina beyrichi
and Stilostomella ewaldi (Beds 19–13: Fig. 1C), which correspond
to the Bolivina beyrichi-Stilostomella ewaldi Abundance Zone (after
Grimm, 2002) and FS 2 and FS 3 (after Grimm, 1991, 1994). The di-
vision between Fischschiefer parts A and B does not correspond to a
shift in dinoflagellate cysts or other microflora.
Some of the calcareous nannoplankton have been reworked from
older Eocene, Cretaceous, or Jurassic strata. Reworking cannot be
identified based on the state of preservation, but is indicated by the
presence of calcareous nannoplankton from NP 11–NP 12. Reworked
dinocysts were recovered from Eocene, but not Cretaceous, species.
4. Materials and methods
Macrofossils occur mainly as scattered remains in the sediment
at the Rauenberg locality. Consequently, many years of intensive
excavations, mainly by private collectors but more recently by the
Staatliches Museum für Naturkunde Karlsruhe (SMNK), were nec-
essary to assemble the collections at the Hessisches Landesmuseum
Darmstadt (HLMD), Institut für Geowissenschaften Universität Hei-
delberg (GEOW), SMNK, and the Staatliches Museum für
Naturkunde Stuttgart (SMNS) on which this study is based. Fossil
plant material has also been studied from the collection of the Natur-
museum Augsburg. The material was collected by many different in-
dividuals over many years, and so stratigraphic information is for the
most part lacking, making it impossible to assess potential changes at
the site over time.
The collection of fossil plant material at the SMNS comprises
> 600 specimens; that of the SMNK has approximately 300, and that
of the Naturmuseum Augsburg > 200 specimens. Unfortunately, the
private collectors focused on well-preserved plant material, largely
neglecting strongly fragmented material and small remains. Conse-
quently, such material has only been sampled since 2007 when J. Ko-
var-Eder contacted the collectors.
Leaves are preserved as carbonized compressions suitable for cu-
ticular analysis (Figs. 2–3). The material collected prior to 2007 was
usually coated with a varnish, which hampers cuticle studies. Cuticle
preparations have been performed from > 280 specimens, however, of
which almost 200 were successful. Cuticles of some taxa are well pre-
served, however most are heavily infected by fungi (Fig. 3).
The fossil insects from Rauenberg are generally poorly preserved
as a very thin and brittle coaly film that is, in many specimens, frag-
mented into tiny rectangular blocks like a mosaic (Suppl. Fig. 1B),
most likely due to unsuitable preparation and storage. Many speci-
mens have been coated with cellulose varnish, which further deterio-
rated the visibility of the fossils.
5. Systematic overview
For a complete list of taxa recovered, refer to Appendix A. The fol-
lowing sections are intended as a brief overview.
5.1. Plankton and benthic foraminifera
Calcareous nannoplankton and dinoflagellate cysts from the
Rauenberg clay pit were investigated by Grimm et al. (2002);
foraminifera have also been studied (Grimm et al., 2002; Weiss,
1988). The taxonomic diversity of the calcareous nannoplankton flora
is low. Dinoflagellate cysts are common in the investigated section.
Planktonic foraminifera are rare, only one short planktonic bloom oc-
curs over the studied interval. Foram diversity is typical for a Rupelian
benthic fauna, with low numbers of species but some species occur-
ring at high abundance. Other calcareous microfossils such as calci-
spheres, interpreted as crustacean eggs (described as Orbulina bitumi-
nosa: Grimm and Grimm (1996)) and tintinnids (Pseudarcella rhum-
beleri) are found in such high abundance that they exceed forams in
some layers (Grimm et al., 2002).
5.2. Macroflora and pollen
Plant macrofossils are abundant at the Rauenberg locality and com-
prise not only leaves and, more rarely, fructifications of the terres-
trial flora, but also marine algae (Fig. 2). A brief overview of plant
macrofossils with several ambiguous determinations was provided by
Winterscheid and Gregor (2008) (Suppl. Table 1). Sittler (1965) pro-
vided a synthetic list of pollen and spores from the Hochberg Mem-
ber in the Alsace region of the Upper Rhine Graben based on drill
cores, and Pross et al. (1998) later studied the palynomorphs from the
Mainz Basin (Middle Rupelton, NP 23, drill core Bodenheim), but
the palynomorphs from the Rauenberg section have not been studied.
The macro remains (leaves and fruits) have recently been evaluated
(Kovar-Eder, submitted for publication).
5.2.1. Aquatic plants
Thalli of at least three different marine algae can be distinguished,
probably belonging to the Phaeophyceae and/or Rhodophyceae (e.g.,
Fig. 2A). One of them bears aerocysts as in, for instance, the modern
genus Cystoseira. Fragments of monocotyledons that might be inter
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx 5
Fig. 2. Plant macrofossils from Rauenberg; all specimens figured are housed at the SMNS. A, Algae gen et sp. indet., slender linear thallus, several times forked; P 1952/1; B, Pinus
(Pinus) cf. thomasiana (Goeppert) Reichenbach, cone, P 1952/176; C, Tetraclinis salicornioides (Unger 1841) Kvaček 1989, branched twig with cupressoid foliage, P 1952/118; D,
Ceratozamia floersheimensis (Engelhardt) Kvaček, leaf fragment with parallel running venation, P 1952/456; E, Platanus neptuni (Ettingshausen) Bůžek, Holýand Kvaček, P 1952/
32; F, Myrica longifolia Unger, P 1952/50; G, Comptonia diformis (Sternberg) Berry, P 1952/166; H, Hydrangea microcalyx Sieber, calyx of a flower, P 1952/145; I, Engelhardia
macroptera (Brongniart) Unger, involucrum of a fruit, P 1952/154; J, Sloanea olmediaefolia (Unger) Kvaček and Hably, P 1952/340; K, Laurophyllum pseudoprinceps Weyland and
Kilpper, P 1952/48; L, Daphnogene cinnamomifolia (Brongniart) Unger, P 1953/7; M, Symplocos volkeri Kvaček, P 1952/65; N, Palmacites lamanonis Brongniart, P 1952/113; O,
Phoenicites sp., rhachis of a pinnate leaf showing the attachment of the pinnae, P 1952/12. Scale = 20 mm (A), 10 mm (B, D–M, O), 5 mm (C), and 50 mm (N).
preted as reeds, sedges or seagrasses (all monocotyledons) are ex-
tremely rare. Remains of reeds and reed-like plants such as sedges or
seagrasses were not identified; those remains preliminarily identified
as “reeds”by Micklich and Hildebrandt (2010) may represent algae or
fragmentary palm (Phoenicites), cycad, or conifer remains.
5.2.2. Land plants (macrofossil record)
The land plant record is diverse. 66 species have been identified,
including 14 new species, some of them of cryptic systematic affinity
(Suppl. Table 2; Kovar-Eder, submitted for publication). At the fam
6 Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx
Fig. 3. Leaf cuticle preparations from the Rauenberg locality. A, Ceratozamia floersheimensis (Engelhardt) Kvaček, stomata-bearing cuticle and short, strongly cutinized cells
arranged in short rows; this structure excludes the reed or sedge-like nature of these remains (compare A–D), P 1952/456/2; B, Platanus neptuni (Ettingshausen) Bůžek, Holýand
Kvaček, stomata-bearing abaxial cuticle with peltate trichomes, P 1952/32/1; C, Daphnogene cinnamomifolia (Brongniart) Unger, stomata-bearing abaxial cuticle, P 1952/76/2; D,
Laurophyllum pseudoprinceps Weyland and Kilpper, stomata-bearing abaxial cuticle, P 1953/48/2; E, Symplocos volkeri Kvaček, stomata-bearing abaxial cuticle, fungal infection via
stoma (arrow) and hyphae, P 1952/65/1; F, Myrica longifolia Unger, tetracellular trichome bases, P 1952/50/1; G, Sloanea olmediaefolia (Unger) Kvaček and Hably, abaxial cuticle,
stomata characteristically faint, P1952/340/1; H, Phoenicites sp. brachyparacytic stomata with parallel orientation, P 1952/12/1. Scale bar = 50 μm.
ily level, Lauraceae are most diverse, represented by at least nine
species, and at the generic level Pinus is most diverse (6 species
represented by fascicles, 2 species by cones: e.g., Fig. 2B). At the
species level, Platanus neptuni is by far the most common element
(Figs. 2E, 3B) followed by Daphnogene cinnamomifolia (Figs. 2L,
3C). Some species are documented by several remains, such as Lau-
rophyllum pseudoprinceps (Figs. 2K, 3D), Tetraclinis salicornioides
(Fig. 2C), and the palms Palmacites lamanonis (Fig. 2N) and Sa
balites major. Other species are represented by few specimens. These
include Betula,Carya,Comptonia difformis (Fig. 2G), Craigia,Dis-
tylium sp., Engelhardia (Fig. 2I), Eotrigonobalanus furcinervis hasel-
bachensis,Hydrangea,Laurus abchasica,Myrica div. sp. (Figs. 2F,
3F), Phoenicites sp. (Figs. 2O, 3H), Arecaceae (? Calamoideae), Pop-
ulus germanica,Sloanea olmediaefolia (Figs. 2J, 3G), S. artocarpites,
Symplocos volkeri (Figs. 2M, 3E), Ceratozamia floersheimensis (Figs.
2D, 3A), Doliostrobus taxiformis,Sequoia abietina,
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx 7
Taxodium sp., and fern frond fragments such as Lygodium kaulfussii,
Approximately 35% of all terrestrial taxa were presumably tall
trees or trees and around the same percentage encompasses taxa that
were likely small trees or shrubs (Suppl. Table 2). Roughly 13% of
plant species represent lianas or climbers. Taxa with anemophilous
and zoophilous pollen dispersal are almost equally represented
(~ 40%). In the fruit vector, zoochorous dispersal (~ 41%) appears
slightly more abundant than anemochorous dispersal (37%); addition-
ally 13% of species probably had a dyschorous or autochorous fruit
More than 60% of the woody angiosperm species were proba-
bly evergreen, bearing either entire-margined (Lauraceae) or minutely
toothed leaves (Distylium,Engelhardia,Myrica,Sloanea, Symplocos).
Less than 20% of woody angiosperms were likely deciduous and for
10–15% it remains ambiguous whether they shed their leaves reg-
ularly. Taphonomic bias is suggested by the low representation of
Fagaceae (a single specimen of Eotrigonobalanus furcinervis hasel-
bachensis), since 8 types of Fagaceae pollen (17% of recovered
pollen) have been reported from more westerly localities in the Rhine
Graben (Sittler, 1965). Deciduous woody angiosperms are not diverse
and, with the exception of Platanus neptuni, are mainly represented by
fructifications (Betula dryadum,Carya quadrangula,Craigia bronnii)
or calyx remains (Hydrangea microcalyx) (Fig. 2H).
Especially noteworthy is the occurrence of Ceratozamia floer-
sheimensis (Zamiaceae) (Fig. 2D, 3A). This cycad is known from
Flörsheim (Fig. 1D), an important early Oligocene locality in the
Mainz Basin (2 fragments), Rauenberg (1), Budapest (1), and Tri-
bovlje (1) (formerly Trifail, Slovenia). The first report of Ceratozamia
was from Flörsheim (Kvaček, 2002), where it was originally described
as Iris floersheimensis by Engelhardt (1911). In the Cenozoic, cycads
were already relicts in the European flora.
Another important finding is Laurus abchasica, a Lauraceae
species commonly occurring as an accessory element in late
Oligocene and especially early to middle Miocene floras of Central
Europe. Along with a record from Flörsheim (one fragment: Kvaček
(2004)), the three fragments from Rauenberg are the oldest reported
records of L. abchasica.
Sloanea artocarpites (Elaeocarpaceae) has been recognized unam-
biguously from Flörsheim (Kvaček et al., 2001). Sloanea olmediaefo-
lia is very abundant in the Tard Clay (Hungary) while S. artocarpites
is also known from Northern Bohemia (ČeskéStředohoří Mountains).
All these records derive from the Oligocene. The most similar living
relatives are restricted to subtropical regions in eastern China.
In general, the flora from Rauenberg has closer affinity to the
Oligocene and even Miocene record of Europe than that of the Eocene.
Our study provides the first survey of gastropods and bivalves from
the Rauenberg locality. The studied material comes from various his-
torical and modern excavations, and since the stratigraphic origin of
the material from within the Rauenberg succession is usually unknown
it is difficult to interpret which of the described benthic invertebrate
taxa occurred together and therefore might have formed benthic com-
munities. The molluscan fauna of the Bodenheim and Alzey Forma-
tions has been little-studied in the Rhine Valley, but is well-known
from neighboring localities in Germany (e.g., Mainz Basin), Bel-
gium, and the Netherlands (e.g., Boekschoten, 1963; Glibert, 1957a,
1957b; Marquet, 2010; Neuffer, 1973, 1984;
Sanderberger, 1858-1863; van den Bosch et al., 1975). However, few
works compare all these localities, which might have resulted in an in-
flation of synonymous names.
The gastropod assemblage (entirely epi- or infaunal) is very un-
usual: of the seven identified genus- or species-level taxa, five are
from predatory families. Of the remaining two, the aporrhaid
Drepanocheilus cf. D. speciosus is considered a semi-infaunal detri-
tivore (Roy, 1994; Saul and Squires, 2015) and the batillariid Gran-
ulolabium plicatum? either an epifaunal grazer or a deposit feeder
(Ozawa et al., 2009).
Muricids and naticids are predators, and the holes they bore in
the shells of their molluscan prey are well-documented in the fossil
record (e.g., Bromley, 1981; Fretter and Graham, 1962; Zonneveld
and Gingras, 2014). Both groups are present in the Rauenberg de-
posits, represented by Muricopsis sp. (Muricidae) and Euspira cf. E.
micromphalus (Naticidae). The Fasciolariidae (represented by Strep-
tochetus sp.) are also malacophagous (also feeding on polychaetes),
but feed by wedging open the valves of bivalves or the operculum of
snails (Modica and Holford, 2010; Wells, 1958) and therefore leave
no trace fossils.
Cassidae (represented by Galeodea depressa) are known predators
of echinoderms (especially sea urchins), leaving boreholes in the tests
(Hughes and Hughes, 1981).
Epitoniids (possibly represented by Opalia? sp.) feed on corals and
sea anemones (Kokshoorn et al., 2007; Robertson, 1963); however
there are no fossils of cnidarians known from Rauenberg.
Of the ten bivalve species, only two (Chama weinheimensis and
Isognomon sp.) are from epifaunal families; the rest are infaunal and
include some possible deep infaunal representatives (Panopea? sp.
and Pholadomya weissi?). Seven species are from families of suspen-
sion-feeding habits; nuculids (represented by Nucula duchasteli; Fig.
4B) might be either suspension-feeders or deposit-feeders, and yoldi-
ids (Portlandia deshayesiana; Fig. 4C) are deposit-feeders. The family
Thyasiridae (represented by Thyasira benedeni; Fig. 4A) has species
with and without chemosymbiotic sulfide-oxidizing bacteria (Dufour,
2005; Rozemarijn et al., 2011; Southward, 1986). Symbiont-bearing
thyasirid species tend to be larger and to live in shallower waters than
species lacking symbionts (Dufour, 2005). The fossil species T. mich-
elottii, from the Austrian Miocene (Badenian), was considered to be
chemoautotrophic by Zuschin et al. (2001). In Rauenberg, T. benedeni
occurs exclusively in life position in the pyrite horizon (i.e., in sul-
fide-rich sediments, Bed 9, Fig. 1C); this suggests chemosymbiosis, as
this species is known from pyrite-bearing horizons in other localities
(e.g., Boekschoten, 1963).
Overall, the most abundant mollusks are the gastropods Strep-
tochetus sp. and Drepanocheilus cf. D. speciosus and the bivalves
Thyasira benedeni (Fig. 4A), Nucula duchasteli (Fig. 4B) and Port-
landia deshayesiana (Fig. 4C).
Only one species of irregular sea urchin was found (Fig. 4F), rep-
resented by 26 specimens in the SMNK collections. They are mostly
fragmented and deformed by sediment pressure and therefore identi-
fication is difficult. The general morphology and the presence of two
goniopores suggests, however, referral to Ova (Spatangoida, Schizas-
Decapod crustaceans from Rauenberg are very rare, represented
only by a few poorly preserved, undescribed specimens of the crab
Coeloma taunicum (Brachyura: Goneplacoidea; Fig. 4G). This species
was widespread in the Rupelian seaway of the Upper Rhine
8 Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx
Fig. 4. Marine macroinvertebrates from Rauenberg. A, Thyasira benedeni; SMNK-PAL 7585, shell length 10.4 mm. B, Nucula duchasteli, with borehole (ichnospecies Sedilich-
nus paraboloides); SMNK-PAL 7343, shell length 13.8 mm. C, Portlandia deshayesiana, with borehole (ichnospecies Sedilichnus paraboloides); SMNK-PAL 7585, shell length
19.9 mm. D, Operculum of a neogastropod (likely Muricoidea), embedded in the rock matrix; SMNK-PAL 8058, greater length 11.9 mm. E, A balanid colony consisting of two
individuals with broken shell showing the internal shell structure (SMNK PAL 7352); scale bar = 10 mm. F, The irregular sea-urchin Ova sp. (SMNK PAL 7796); scale bar = 10 mm.
G, Coeloma taunicum (v. Meyer, 1862), SMNS 106866. Total width of specimen including pereiopods = 100 mm.
Graben and is also known from clayey deposits of Central Germany
(Freess, 1992; Polkowsky, 2005; von Fritsch, 1870). The extreme flat-
tening of the specimens of C. taunicum from Rauenberg and the ab-
sence of the pleon in ventral view indicates that these fossils do not
represent carcasses but moults. It is likely that preservation biases or
low benthic oxygen levels are responsible for the absence of other de-
capods such as shrimps and squat lobsters which would be expected
in this type of environment. From elsewhere in the region (Dämmel-
wald I and II and Dammstücker clay pits northeast of Wiesloch),
Wagner-Klett (1919) described additional crustaceans. Reappraisal of
these remains suggests that, in addition to C. taunicum, ghost shrimps
(Callianassidae gen. et sp. indet.), and the lobster Hoploparia klebsi
(Noetling) were present. Hoploparia klebsi is widely distributed in
Oligocene sediments from Northern Germany.
Barnacles are also known from Rauenberg. They are represented
by shells of two unattached individuals in the SMNK collection (Fig.
4E). Two further individuals attached to the entoplastron of a che-
loniid turtle were figured by Alexander and Frey (2010). The shape
of the shells is most similar to the genus Protochelonibia, which
is thus far the oldest representative of the extant sea-turtle fouling
Chelonibiidae (Cirripedia: Balanomorpha; Harzhauser et al. (2011)).
The SMNS collection additionally houses an unattached colony of an
unidentified smaller type of barnacle as well as four samples with
poorly preserved shells of goose barnacles (Pedunculata).
A relatively rich assemblage of fossil insects is known from
Rauenberg (Fig. 5; Suppl. Fig. 1), however the material is largely un-
described, aside from the work of Monninger and Frey (2010) and
a brief mention in recent newspaper reports (Anonymous, 2014; Ott,
2014). 340 specimens were examined from the SMNK and SMNS col-
lections as part of the current study (Appendix B). Although many of
these were collected by private individuals, most notably H. and A.
Oechsler, it is important to note that the collection does not seem to
be affected by collection bias, as every item remotely resembling a
fossil was collected and preserved. Only very few of the fossil insect
specimens are associated with stratigraphic data. However, anecdotal
information provided by the fossil collectors suggests that insects are
mostly found in the upper beds.
With few exceptions (i.e., cicada, SMNS 112 and jewel beetle,
SMNS 140; Fig. 5A,D) there are no details preserved in the fossil
insects from Rauenberg, apart from a shadowy outline of the insect
body. Some of the better-preserved specimens include a few larger
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx 9
Fig. 5. Fossil insects from the Oligocene of Rauenberg. A, Auchenorrhyncha, SMNS 112, the only insect from this locality with faintly preserved wing venation. B, Auchenorrhyn-
cha, wing, SMNK-PAL-unnumbered. C, Coleoptera, SMNS 39b (the fossil is black, the reflection is due to coating with cellulose varnish). D, Coleoptera, Buprestidae, SMNS 140.
E, Coleoptera, Carabidae, SMNS 4. F, Odonata, Zygoptera, SMNK-PAL-8379. G, Odonata, Zygoptera, Calopterygidae, SMNK-PAL-8089 (note the complete interpleural suture).
H, Coleoptera, Curculionidae, SMNK-PAL-6565. I, Heteroptera, Pentatomidae, SMNS 21. J, Lepidoptera, Geometridae (note the small eyespot on distal part of wing), specimen
is prepared by transfer to resin, SMNK-PAL-6572. K, Hymenoptera, Formicidae, SMNS 161. L, Arachnida, Scorpiones?, SMNK-PAL-7798. Scale parts A–G, I, L = 5 mm; part
J = 10 mm, parts H,K = 2 mm.
beetles (Fig. 5C,H) and beetle elytra (Fig. 5E), a butterfly (Fig. 5J),
and a complete fossil damselfly (Fig. 5F).
Among the 340 fossil insects in the studied collections, 122 spec-
imens (36%) are indeterminate insects. Among the remaining 218
specimens, representatives of 8 insect orders (Odonata, Orthoptera,
Auchenorrhyncha, Heteroptera, Hymenoptera, Coleoptera, Diptera
and Lepidoptera) could be recognized. The record of the superorder
Amphiesmenoptera by Monninger and Frey (2010) was confirmed as
Lepidoptera-Geometridae. Family-level determination was only pos-
sible for 33 specimens, representing 11 families or superfamilies
(Calopterygidae, Carabidae, Staphylinidae, Elateridae/-oidea,
Buprestidae, Cerambycidae, Curculionidae/-oidea, Pentatomidae/-
oidea (Fig. 5I), Formicidae (Fig. 5K), Bibionidae, and Geometridae).
Of the 218 specimens that could be determined to order-level, the vast
majority (154 specimens: 71%) are Coleoptera. Nine specimens repre-
senting larvae or pupae have also been recovered.
10 Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx
In the SMNK collection, there are four specimens of the order
Odonata that can be attributed to the suborder Zygoptera (Fig. 5F,G).
Even though few details of the wing venation are preserved, all spec-
imens except SMNK-PAL-8379 can be determined to family level as
Calopterygidae based on the complete interpleural suture (Asahina,
1957) of the very robust pterothorax (Fig. 5G), which is otherwise
only known from the exclusively Neotropical family Polythoridae.
Similar size indicates that all four specimens could represent the same
taxon (Suppl. Table 3). At least one specimen (SMNK-PAL-8088,
and probably SMNK-PAL-8379) is a female, as the ovipositor pouch
is visible on the abdomen. Because calopterygids oviposit into sub-
merged aquatic vegetation, it can be presumed that these specimens
are all females that drowned during misguided oviposition attempts on
Otherwise, most of the insects are terrestrial pterygotes that were
probably transported by wind into the marine environment. The large
percentage of complete insects (ca. 75% complete vs. 25% fragments
or isolated elytra), including delicate insects such as Diptera and Hy-
menoptera, suggests that the animals drowned and rapidly sank to
the bottom. Beetles probably had a longer drifting time because of
air trapped beneath the elytra. During decomposition of the animals,
the elytra became detached (Fig. 5E). A strange phenomenon is the
preservation of both elytra together without any other remains of the
body in 10 of the fossil beetles. The lack of damage to the elytra ex-
cludes the possibility that this fragmentary preservation is due to pre-
dation. The most likely explanation is incomplete decay, where the
elytra have been detached from the thorax but remained in connection
to each other.
There are only three terrestrial arthropods from Rauenberg that are
not pterygote insects. All three are arachnids; myriapods are absent.
We also discovered a large arachnid (SMNK-PAL-7798) originally
misidentified as a beetle. This specimen is about 23 mm long with
seven of eight legs preserved, segmented opisthosoma, half-ovoid pro-
soma/carapax, long pedipalps (1 preserved) with two elongate basal
segments, and apparently orthognathous chelicerae (Fig. 5L). At first
glance, it resembles a spider of the suborder Mesothelae; however,
the shape of the prosoma/carapax, the pattern of segmentation of the
opisthosoma, and the broad connection between these two tagmata re-
fute an attribution to Araneae. This specimen most likely represents a
poorly preserved and partly decayed scorpion lacking the tail and the
pincers (chelae) (P. Selden pers. comm., 2015).
5.4.1. Chondrichthyes and Actinopterygii
Fish remains (Figs. 6–7) are the dominant faunal elements of the
Rauenberg clay pits, both in the field and in museum collections
(Micklich and Hildebrandt, 2010; K. Eck, pers. observ.). Fishes from
Rauenberg were first mentioned in the literature by Wagner-Klett
(1919) and later revised by Weiler (1931), who identified a total of
19 taxa. In a subsequent publication, Weiler (1966) emphasized the
general significance of this fish fauna for paleobiological reconstruc-
tion and paleogeography. He recognized a total of 31 fish taxa, with
slight taxonomic deviations from his earlier publication. However, ac-
cording to the maps from these publications, these authors referred
to records from the ancient Dämmelwald I and II and Dammstücker
clay pits NE of Wiesloch (Wagner-Klett, 1919), and the Rohrlach and
Frauenweiler-Wiesen clay pits east of the small village of Frauen-
weiler, now a suburb of Rauenberg (Micklich and Hildebrandt, 2005;
Weiler, 1966) (Fig. 1B). The Unterfeld clay pit was first mentioned as
a fossil fish locality by Eikamp (1983), but without any faunistic de-
tails. A detailed synopsis of the fishes from this locality was pub-
lished by Micklich and Parin (1996), who distinguished 52 distinct
taxa, and was updated by Micklich (1998), who corrected some er-
rors and briefly characterized some new faunal elements. The num-
ber of teleosts recognized from the locality was further increased
by Micklich and Hildebrandt (2010), who included 72 taxa in their
paleobiological analyses. The chondrichthyan fauna was revised by
Hovestadt et al. (2010). The updated faunal list reflecting ongoing
study of the material now comprises 74 discrete taxa (Appendix A;
Suppl. Table 4). It must to be stressed that this synopsis is a snapshot,
as several of the listed taxa are only preliminary identifications and are
in need of detailed revision.
Recently, several taxonomic revisions of the Rauenberg fishes
have been undertaken. The shark Triakis kelleri was described by
Hovestadt and Hovestadt-Euler (2002) from the Unterfeld clay pit,
and Carcharhias gustrowensis is also present at this locality, although
most records are from younger (Chattian) localities (Hovestadt and
Hovestadt-Euler, 2010). The record of an almost-complete juvenile
basking shark (Hovestadt and Hovestadt-Euler, 2012) is also of strik-
ing importance, as little is known concerning the spawning behav-
ior and presumed nursery grounds of these sharks (McCandless et
al., 2007). Weissobatis, an eagle ray, was described as coming from
the lower parts of the Meletta Beds (Hovestadt and Hovestadt-Euler,
1999). If so, this would be one of the youngest vertebrate fossils
from the locality. It is significant for myliobatid systematics and phy-
logenetic relationships. Among the Syngnathoidei (revised by Parin
and Micklich, 1996a; Parin and Micklich, 1996b), Frauenweilersto-
mus synarcualis, a genus and species which is not rare but is ex-
clusively reported from Rauenberg, is particularly noteworthy. Ae-
oliscus distinctus is also only known from Rauenberg. In addition,
the presence of the syngnathids Microphis and Nerophis extend the
stratigraphic range of these taxa from essentially Recent (Microphis:
Serravallian-Langhian (Bachmayer, 1980); Nerophis: early Miocene
(Sergienko, 1971)) into the Paleogene. Musculopedunculus (Parin and
Astakhov, 2007) is of special interest as it represents a new family of
the Trichiuroidae, and Oligoremora rhenana, a new genus and species
of Echeneidae, has recently been described which, based on meris-
tic data, most closely resembles representatives of the extant genus
Remora (Micklich et al. in press). The presence of tholichthys larvae
(Fig. 7F) represents the oldest record of butterfly fishes worldwide
(Micklich et al., 2009). Interestingly, and in contrast to their great rar-
ity in extant tropical plankton and midwater trawl samples –according
to Leis (1989), they have a mean frequency of 0.09% of all larvae cap-
tured –two additional records were found in 2012 (N. Micklich, pers.
As previously suggested (Micklich, 1998, 2005), some of the taxa
listed in Appendix A may be “waste-basket”taxa comprising more
than one nominal genus and/or species. Others remain undescribed.
The total diversity of the fish fauna is therefore undoubtedly greater
than the current state of the art. For instance, a new lophiid genus and
species is recognized, closely related to the extant Sladenia and the
fossil Caruso (G. Carnevale, pers. comm.). Ongoing study of Prop-
ercarina (Figs. 6L, 7J) indicates that the Rauenberg material differs
from the other known species and may represent a new species (T.
Přikryl, pers. comm., N. Micklich, pers. observ.), and a similar situ-
ation exists concerning the presumed Leiognathoides (A. Bannikov,
Y. Yabumoto, pers. comm., N. Micklich, pers. observ.). However,
the largest ongoing work is a revision of the scombroids (K. Mon-
sch, pers. comm., N. Micklich, pers. observ.), including documenta-
tion of material in existing collections and detailed systematic revision
of selected taxa. Currently, about 30 genera and species can be distin-
guished (K. Monsch, pers. comm.). Some of them, e.g., those resem
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx 11
Fig. 6. Typical fishes and fish remains from the Rupelian of Rauenberg. A, Basking shark, Keasius parvus (HLMD-SMFF 32), bundle of gill rakers from filtering apparatus; B,
Requiem shark, Physogaleus latus, lower lateral tooth in lingual view (HLMD-WT 705); C, Eagle ray, Myliobatidae indet. aff. “Myliobatus”var. oligocaena, symphysal tooth
(HLMD-WT 707); D, Herring, “Sardinella”sardinites (HLMD-SMFF 667); E, Shrimpfish, Aeoliscus heinrichi (HLMD-WT 861); F, Pipefish, Doryrhamphus sp. (HLMD-SMFF 7);
G, Pair of sea bass, Oliganodon budensis (HLMD-SMFF 15a); H, Bigeye, Priacanthus spinosus (HLMD-WT 687); I, Jack, Archaeus glarisianus (HLMD-WT 37); J, Bonito, Sarda
brachycephala (HLMD-WT 41); K, Bill fish, Palaeorhynchus cf. P. glarisianus (HLMD-SMFF 27); L, Stromateid-like fish, Propercarina sp., arrows indicate the vertebral column
of prey fish in the gut (HLMD-SMFF 23); M, Stromateid-like fish, Pinichthys pulcher (HLMD-WT 29). Scale bars A–C, F–G = 5 mm, scale bars D–E, H–I, L–M = 10 mm, scale
bars J–K = 50 mm.
bling Diplospinus and Neoepinnula, were previously known exclu-
sively from extant representatives. Others (e.g., Abadzekhia) were
only known from localities in the eastern Paratethys, and more are
completely new and as yet undescribed.
The fish fauna is typically Cenozoic, specifically Rupelian. Aside
from the presumed sphyraenids, no actinopterygian species range ex-
tends earlier than the Paleocene (Suppl. Fig. 2). About 40 are more
or less restricted to the Rupelian or slightly younger layers, and 11
species are known exclusively from Rauenberg. This number will in
12 Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx
Fig. 7. Early juvenile stages of fishes from the Oligocene of Rauenberg. A, Herring, cf. “Sardinella”sardinites (HLMD-WT 764); B, Shrimpfish, Aeoliscus heinrichi (HLMD-WT
937); C, Scorpionfish, cf. Sebastes sp. (HLMD-SMFF 93b); D, Seabass, cf. Oliganodon budensis (HLMD-WT 200); E, Bigeye, Priacanthus spinosus (HLMD-WT 361); F, Butterfly-
fish tholichthys larval stage (HLMD-WT 410a); G, Jack, cf. Archaeus glarisianus (HLMD-WT 815); H, Spanish mackerel, Scomberomorus cf. S. lingulatus, arrow indicates prey fish
in mouth (HLMD-WT 803b); I, Bill fish, Palaeorhynchus glarisianus, arrow indicates prey fish in mouth (HLMD-WT 807); J, Stromateid-like fish, Propercarina sp. (HLMD-WT
224); K, Stromateid-like fish, Pinichthys pulcher (HLMD-WT 224); L, Flatfish, Oligoscophthalmus weissi (HLMD-WT 254). Scale bars = 5 mm.
crease following reevaluation of some of the groups discussed above
(e.g., Lophiidae, Scombroidei), as well as those specimens currently
identified only to genus or family. 14 taxa are also known from ex-
tant forms. For at least six of them, the records from Rauenberg ex-
tend their stratigraphical range from (almost) Recent into the lower
In the mudstones of the clay pits around Rauenberg, turtles are
rare. Due to their predominantly fragmentary nature, little work has
been undertaken on this clade. Only 19 specimens are known to date.
There are two near-complete skeletons and four fragments at the
SMNK, two partial skeletons and 10 fragments at the HLMD, one
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx 13
isolated humerus at the GEOW, and seven specimens at the SMNS. Of
these, only the more complete material housed at the SMNK has been
described (Alexander and Frey, 2010).
Marine turtles or Chelonioidea comprise three families: Cheloni-
idae, Protostegidae, and Dermochelyidae. The bulk of the marine tur-
tles from Rauenberg, including the material previously described by
Alexander and Frey (2010), are referred to Cheloniidae despite hav-
ing a wide range of body sizes (Fig. 8A–D). The Karlsruhe speci-
mens (SMNK-PAL 6608, SMNK-PAL 6609: Fig. 8A–B) are poorly
preserved, permitting identification only to family level. Within the
Stuttgart sample, the better articulated specimens closely resemble
“Chelonia”gwinneri, particularly a large adult (SMNS 80529/1);
SMNK-PAL 6608 is also similar to “Chelonia”gwinneri (Alexander
and Frey, 2010). The Rauenberg material highlights problems with the
assignment of juvenile cheloniid specimens to species. For instance,
SMNK-PAL 6609 strongly resembles Glarichelys knorri, however,
the secondary transversal and longitudinal sections of the neuralia
are more similar to the late Miocene taxa “Euclastes”melii and Pro-
colpochelys grandaeva (Alexander and Frey, 2010). At present,
Glarichelys knorri is based exclusively on juvenile specimens.
The nine cheloniid specimens studied here fall into a wide size
range and most probably form an ontogenetic series. Further studies
are required to (1) test this hypothesis and (2) address the question of
whether the single Rauenberg taxon is distinct from “Chelonia”gwin-
neri and Glarichelys knorri.
In addition, a single specimen (SMNS 96919: Fig. 8E) of
soft-shelled turtle (Trionychidae) is reported here for the first time,
and bears a close resemblance to Trionyx (Platypeltis)posterus. Ex-
tant trionychid turtles preferentially inhabit freshwater but also occur
in brackish water, sometimes with surprisingly high salt concentra-
tions (Ernst et al., 2000). However, they have been reported to tem-
porarily invade marine habitats (Das, 2008; Shanas et al., 2012). The
presence of a trionychid at Rauenberg may indicate the presence of a
nearby freshwater source, such as a river mouth. Trionychid and ch-
eloniid turtles are uninformative climatic indicators because of their
wide temperature tolerance. A number of extant trionychids occur at
high latitudes (45°) and hibernate (e.g., Harding, 1997; Reese et al.,
2003). Cheloniid sea turtles also occasionally invade subpolar waters
(e.g., Bustard, 1972; Carr, 1952).
Fig. 8. Cheloniid Turtles from Rauenberg. A, an almost-complete cheloniid SMNK-PAL 6608; note the plates were reassembled incorrectly (plate borders are marked in white); B,
a disarticulated cheloniid SMNK-PAL 6609; C, a juvenile cheloniid including skull and forelimbs SMNS 87449/26; D, a juvenile cheloniid carapace SMNS 80738/2; E, carapace
referable to Trionychidae (SMNS 96919). Photos parts A, B courtesy of Griener; used with permission. Scale = 1 cm (C) and 5 cm (D, E).
14 Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx
Rauenberg is one of the most significant localities for early
Oligocene fossil birds in Europe, being among the few sites that
yield well-preserved partial skeletons rather than isolated bones. The
most abundant birds from Rauenberg are procellariiforms of the taxon
Rupelornis (Diomedeoididae), of which several partial and complete
skeletons are known (Mayr, 2009a; Mayr et al., 2002; Mayr and
Smith, 2012) (Fig. 9A). Two species appear to have coexisted, whose
taxonomic status is in need of revision. One is likely to be conspecific
with R. definitus from the Rupelian of Belgium, whereas the other
probably belongs to a Rupelornis species that was first described from
the early Oligocene of France. Diomedeoidid procellariiforms had a
wide distribution in the early Oligocene epicontinental seas of Europe
and the Near East, having been reported from localities in Belgium,
France, Switzerland, Poland, and Iran (Mayr, 2009b).
Another aquatic bird from Rauenberg is the loon Colymboides?
metzleri (Gaviiformes; (Mayr, 2004b)). This species is known from a
single skeleton and seems to be closely related to a loon species from
the Rupelian of Belgium (Mayr, 2009c; Mayr and Smith, 2013).
Altogether, six species of landbirds are known from Rauenberg,
all of which provide significant insights in the evolution of their re-
spective groups. Today, the nectarivorous hummingbirds (Trochili
Fig. 9. Articulated and associated avian remains from Rauenberg. A, Rupelornis cf.
brodkorbi (Procellariformes: Diomedeoididae; SMNK-PAL 3812); B, Eurotrochilus in-
expectatus (Trochiliformes: SMNK-PAL 5591). Scale = 5 cm (A) and 1 cm (B).
dae) only occur in the New World. At the time of their description,
specimens of Eurotrochilus inexpectatus from Rauenberg represented
the earliest fossil records of hummingbirds (Mayr, 2004a, 2007; Mayr
and Micklich, 2010) (Fig. 9B). Closely related species are now known
from the early Oligocene of the Lubéron area in France and from the
Rupelian of Poland (Mayr, 2009b).
Two further landbirds from Rauenberg that closely resemble re-
lated species from the early Oligocene of the Lubéron area include
aPrimotrogon-like trogon (Trogoniformes), which is known from
a partial wing (Mayr, 2005a), and Turnipax oechslerorum, a stem
group representative of buttonquails (Turnicidae) known from a sin-
gle specimen (Mayr, 2009b; Mayr and Knopf, 2007a). Trogons have
a pantropical distribution and include insectivorous and frugivorous
species. They excavate nest cavities in tree trunks and therefore oc-
cur in forested areas and open woodlands. Buttonquails are an Old
World group of small charadriiform birds, which today occur in arid
semi-deserts, but Turnipax exhibits a much more primitive morphol-
ogy than its extant relatives and its ecological preferences may have
A stem group representative of mousebirds (Coliiformes), Oligo-
colius brevitarsus, is represented by a dissociated partial skeleton
(Mayr, 2000). The Rauenberg fossil was the first record of the distinc-
tive taxon Oligocolius, which appears to have been adapted to a more
aerial way of life than its extant African relatives. A second more re-
cently described species of the taxon from the late Oligocene of Ger-
many (Mayr, 2013) is associated with preserved stomach contents, in-
dicating that Oligocolius was a specialized frugivore.
Two partial skeletons of a tody, Palaeotodus itardiensis (Todidae;
(Mayr and Knopf, 2007b; Mayr and Micklich, 2010), have also been
recovered. Todies are very small alcediniform birds, which have a
relict extant distribution on the West Indian islands. Palaeotodus itar-
diensis was originally described from late Eocene and early Oligocene
localities of the Quercy fissure fillings in France (Mayr, 2009b).
One of the oldest representatives of the Pici, the clade includ-
ing woodpeckers and allies, is known from Rauenberg: Rupelram-
phastoides knopfi. This species is one of the smallest known repre-
sentatives of the clade (Mayr, 2005b, 2006). Rupelramphastoides does
not exhibit any of the specialized dietary or locomotory adaptations
found in its extant relatives.
Rauenberg also yielded fossils of a passerine bird, Wieslochia
weissi (Mayr and Manegold, 2004, 2006). At the time of its descrip-
tion, it was the oldest European fossil occurrence of Passeriformes,
the most species-rich clade of extant birds, which includes more than
half of all extant avian species, although in the past years equally
old fossils have been described from the early Oligocene of Poland
(Bocheński et al., 2011, 2013). Wieslochia weissi is among the earliest
well-represented fossil passerines.
Mammalian remains from Rauenberg are extremely rare, with very
few specimens known. These include a creodont carnivore (Frey et al.,
2010), as well as undescribed sirenian material (Dugongidae) and a
The dugongid remains consist of a nearly-complete specimen at
the HLMD (Schöggl and Micklich, 2012), an isolated forelimb at the
SMNS, and skull and rib fragments at GEOW, and have been referred
to the early Oligocene species “Halitherium cf. schinzii”based on the
occurrence of this taxon in other early Oligocene coastal deposits from
Central Europe, especially Germany (e.g., Domning and Pervesler,
2001; Kaup, 1838; Voss, 2008).
The chiropteran specimen housed at the SMNS consists of a par-
tial skeleton preserved as part and counterpart, including a left den
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx 15
tary and a right maxillary fragment. There are no accessory cuspules
on the lingual talon of the upper molars, and so a referral to Leuconoe
can be ruled out. The specimen is most consistent with the vespertil-
ionid Myotis horaceki from the late Oligocene of Germany (Ziegler,
2000, 2003). A second nearly-complete specimen is housed in the
Castle of Bruchsal Museum, but was not available for study.
The creodont Apterodon rauenbergensis consists of a mandibular
fragment. Other European taxa referred to this genus have been recov-
ered from the Rupelian fissure fillings of the Quercy Phosphorites and
the Mainz and Weisselster Basins. This is the first terrestrial mammal
reported from Rauenberg.
6.1. Biotic interactions and soft-tissue preservation
There are several bivalves in the Rauenberg material (< 10% of
the total sample) bearing boreholes attributable to the ichnospecies
Sedilichnus paraboloides (Fig. 4B–C). Recent representatives of both
Naticidae and Muricidae are known to create holes comparable to
S. paraboloides (Zonneveld and Gingras, 2014), suggesting that gas-
tropods from these families are responsible for these traces. There are
no traces of predation on Rauenberg echinoderms, in spite of the pres-
ence of Cassidae (Gastropoda), perhaps not surprisingly since echino-
derms are represented only by burrowing spatangoids.
Many vertebrate fossils from Rauenberg are exceptionally pre-
served, capturing ecological interactions and soft-tissue details rarely
available for fossil taxa. The fishes are particularly informative.
Among the sharks, Carcharhias gustrowensis is known from a preg-
nant female containing eight embryos in the reproductive tract
(Hovestadt and Hovestadt-Euler, 2010), while the holotype of Triakis
kelleri is preserved as the stomach contents of a carcharhinid shark
(Hovestadt and Hovestadt-Euler, 2002). One specimen of the bird Ru-
pelornis is preserved with two shark teeth that probably stuck in the
soft tissue of the carcass and either indicate direct predation or scav-
enging (Mayr et al., 2002).
Many of the teleost fishes also preserve gastrointestinal contents.
All macroscopic gut contents identified preserve the remains of other
fishes. Piscivorous fishes in the assemblage include the aulostomids
Frauenweilerstomus and Aulostostomus, as well as Palaeogadus,
Palaeorhynchus (Fig. 7I), Propercarina sp. (Fig. 6L), trichiurids,
scombrids (e.g., Sarda,Scomberomorus: Fig. 7H) and carangids (e.g.,
Seriola,Archaeus). Some of the smallest scombrids (30 mm in total
length) preserve other fishes in the digestive tract. Particularly note-
worthy is a number of specimens of Archaeus (7 individuals) pre-
served with Oliganodon partially swallowed, emphasizing that these
two common genera encountered each other fairly often and were not
spatially or temporally segregated. A specimen of Carcharhias gus-
trowensis is interpreted as having myliobatid and chimaeroid spines
embedded in the jaw cartilages (Hovestadt and Hovestadt-Euler,
2010). In addition to gastric contents, a single specimen of Lepido-
pus, as well as several scombrids, have phosphatic cololites (intestinal
casts) preserved in the posterior abdominal region; these occasionally
contain externally visible fish scales (e.g., SMNS 87457/297).
Stomach contents consisting of a large mass of fish bones are
preserved in the fossil loon Colymboides?metzleri (Mayr, 2004b),
and the holotype of the turnicid Turnipax oechslerorum contains gas-
troliths (Mayr and Knopf, 2007a). A cheloniid turtle also preserves
unidentified intestinal contents (Alexander and Frey, 2010).
One of the cheloniids shows evidence of colonization by bal-
anomorph barnacles during its lifetime (Alexander and Frey, 2010).
Barnacles are filter-feeders that are attached to a hard substrate, which
can be either rocks or larger animals. The isolated specimens ei-
ther fell off a dead turtle, a marine mammal, such as the dugongid
“Halitherium cf. schinzii”, or driftwood in the case of the goose barna-
cles. The barnacles attached to the turtle carapace represent the oldest
record of sea-turtle fouling balanids in the fossil record, a biological
interaction that has otherwise only been recorded from the Miocene
(Harzhauser et al., 2011; Hayashi et al., 2013).
The leaf remains are often strongly infected by fungi. The cuticles
show both hyphae and fructifications. Often the stomata are masked
by fungi that obviously invaded the leaves (Fig. 3E). Fungal infection
and growth started shortly prior to or after abscission, during the trans-
port of the leaf but prior to deposition.
Several of the fishes preserve color patterns (e.g., Microcanthus
sp.: Micklich and Parin, 1996), and even more common is preservation
of retinal pigments (Figs. 6M, 7C–E, K). Some of the keratin plates
in both the SMNK cheloniids are preserved as a bacterial crust (see
Martill, 1987; Wuttke, 1996). Except for those of the procellariiform
Rupelornis, most bird skeletons are strongly dissociated, which is es-
pecially true for small land birds. Although soft-tissue preservation
appears to be rare, a specimen of Rupelornis (SMNS 85947/1) shows
preservation of feathers; an isolated feather is also known (SMNS un-
Numerous authors have postulated a marine connection during
the Rupelian between the Paleogene North Sea and the Paratethys
to the South via the Upper Rhine Graben (but see Berger, 1996,
who questioned the existence of a southern connection). The south-
ern connection is hypothesized to have passed through either the
Bresse and Rhône Valleys and the Raurachian Depression (Doebl and
Teichmüller, 1979; Sittler, 1992; Weiler, 1952, 1956) or via a narrow
straight along the uplifting Alps (Büchi, 1983; Spiegel et al., 2007).
Berger et al. (2005a, 2005b) identified three transgressive events dur-
ing the Rupelian and did not exclude the existence of a marine con-
nection between the North Sea, the Upper Rhine Graben, and the
Paratethys during the deposition of the Hochberg Member (nanno-
plankton zones NP 21–23) (Fig. 1D). Based on the lateral continu-
ity of the Froidefontaine Subgroup, Sissingh (2003) also concluded
an intermittent North-South marine connection existed during the Ru-
pelian, via the Hessian Depression to the North. The existence of a
North-South marine connection during the early Oligocene has been
supported based on calcareous nannoplankton (Martini, 1982; Martini
and Müller, 1971), fossil fishes (Micklich, 1998), and sedimentologi-
cal markers (Kuhlemann and Kempf, 2002).
Consistent with a connection to the North Sea, the calcareous
nannoplankton flora corresponds to the Zone NP 23 of Northern Ger-
many, but in the Rauenberg section Reticulofenestra dictyoda and R.
scissura are less abundant. The foraminiferal fauna agrees with the
fauna in the Hochberg Member of the Bodenheim Formation of the
Mainz Basin. The dinoflagellate Tectatodinium pellitum is also found
in the Bodenheim (Hochberg and Rosenberg Members), Alzey, and
Stadecken Formations of the Mainz Basin (Pross, 1997).
The molluscan species and genera identified at Rauenberg are all
known from coeval and neighboring deposits in Germany (Mainz
Basin) and Belgium (e.g., Marquet, 2010; Neuffer, 1973, 1984). The
molluscan fauna of Rauenberg appears to be much less diverse than
those of coeval localities, however this may be biased by the small
number of specimens recovered.
16 Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx
The paleobiogeography of the fish fauna of the Wiesloch-Rauen-
berg Tertiary block was first considered by Weiler (1931). More mod-
ern revisions suggest that, aside from some northern elements, the
Rauenberg fish fauna is very similar to that of localities in the eastern
Paratethys (Micklich and Parin, 1996; Pharisat and Micklich, 1998).
Centriscids generally are typical for the Red Sea and Indo-Pacific;
Leiognathoides [“Equula”]altapinna is described from the lower
Oligocene (Rupelian) of Allschwil in Switzerland (Weiler in Hess
and Weiler, 1955), as well as from the lower Oligocene of the North
Caucasus (Bannikov and Parin, 1997); Anenchelum angustum,Palae-
orhynchus glarisianus and P. zitteli are reported from the lower
Oligocene of Canton Glarus, Switzerland (Furrer and Leu, 1998;
Monsch and Bannikov, 2012); Syngnathus incompletus,Repropca
sabbai,Propercarina rebeli, and P. pietschmanni are known from
the Rupelian of Suslăneşti-Muscel in the Romanian Carpathians
(Bannikov, 1991; Cosmovici, 1887; Přikryl et al., 2014).
Oligosphenopsis gracilis,Abadzhekia marinae,A. tarletskovi,Auxides
cernegurae,Pinichthys pulcher, and Rybapina caucasica are typical
elements of the upper Eocene (Kuma Horizon), the lower Oligocene
(Psheka Horizon), as well as for the Chattian of the northern Lower
Caucasus (Bannikov, 1988, 1995; Bannikov and Parin, 1997; Monsch
and Bannikov, 2012).
As with the fishes, the cheloniids show paratethyan affinities: “Ch-
elonia”gwinneri is known only from the Upper Rhine Graben lo-
cality of Flörsheim (Wegner, 1918), whereas Glarichelys knorri is
known from Glarus, Switzerland and several localities in the Eastern
Paratethys (Winnica, Ukraine; Caspian Sea region: Aslanova et al.,
1979; Gray, 1831, Młynarsky, 1959).
Based on foraminiferal assemblages, Grimm (1994) suggested a
N-S-directed current system predominated in the Upper Rhine Valley
marine strait during deposition of the Hochberg Member. The origi-
nally northerly current switched to a southerly current during the later
Rupelian, concurrent with deposition of the Meletta Beds (Grimm,
1994; Spiegel et al., 2007). This north to south current direction is con-
sistent with the North Sea affinities of the plankton in the Hochberg
Member. An ecological filter at the southern opening has also been in-
voked to explain the discrepancy between the invertebrate and verte-
brate faunas (Spiegel et al., 2007).
Several Oligocene floras are known from marine deposits along
the western and southern coasts of ancient Europe (Fig. 1D). In the
Rhine Graben, the flora of Flörsheim (Mainz Basin), also dated to NP
23, has been recently revised (Kvaček, 2004). The flora of Rauenberg
closely resembles that of Flörsheim, both with regard to the deposi-
tional facies, floristic composition and the fact that numerous species
are known from only one or few specimens (Suppl. Tables 1–2). The
most remarkable similarities are the presence of the cycad Ceratoza-
mia floersheimensis,Sloanea, and the thus-far oldest occurrences of
Laurus abchasica and fan and feather palms in both assemblages. De-
ciduous woody taxa are diverse neither in Flörsheim nor in Rauen-
berg, but in the former locality single leaves document the presence
of Carpinus,Ostrya, and Rosa, while in the latter these have not
been detected. Only a single leaf of probable Betulaceae affinity and
a winged fruit of Betula are documented. While nearly absent from
Rauenberg, three species of Fagaceae are recorded from Flörsheim,
among which Trigonobalanopsis rhamnoides is rather common. The
differences between the sites may partly be related to more favor-
able depositional conditions for terrestrial plants in Flörsheim than in
Rauenberg. It is remarkable, however, that palm remains are more di-
verse and abundant in Rauenberg than in Flörsheim. At each site, nu-
merous cryptic taxa occur. The plant assemblage of Flörsheim is clas
sified to the floristic complex Nerchau-Flörsheim (Kvaček and
Along the northern Paratethys coast, the upper part of the Tard
Clay Formation, mainly the sites Budapest and Eger-Kiseged, Hun-
gary (Hably, 1992; Kvaček and Hably, 1998), are also dated to NP 23.
This material is less suitable for cuticle studies than that of Rauenberg
or Flörsheim, which impairs comparisons. Rauenberg (and Flörsheim)
share the occurrence of Ceratozamia floersheimensis,Daphnogene,
Engelhardia,Laurophyllum,Platanus neptuni,Sloanea, and the
conifers Pinus,Taxodium, and Tetraclinis salicornioides, as well as
the extremely low diversity of deciduous woody angiosperms, with
the flora of the Tard Clay. Rauenberg and the flora from the Tard
Clay Formation share rare records of Craigia fructifications. Rauen-
berg differs from the Hungarian flora among others by the presence,
diversity (5 species), and abundance of palm leaf fragments as well as
by the absence of taxa characteristic of Eocene and early Oligocene
floras such as Chamaecyparites hardtii.
Further east, the plant assemblage from the surroundings of Linz,
Austria (Kovar, 1982), recently dated to the earliest Miocene (Aqui-
tanian: Grunert et al., 2010), shares several taxa with Rauenberg,
such as Lauraceae (Daphnogene,Laurophyllum pseudoprinceps,L.
acutimontanum), Platanus neptuni,Engelhardia,Pinus,Tetraclinis
salicornioides, and Taxodium. Furthermore, an as-yet undescribed
feather palm also from the area of Linz and now stored in the collec-
tions of the Vienna Natural History Museum indicates similarities to
the flora from Rauenberg (J. Kovar-Eder pers. observ.). The Linz flora
differs from that of Rauenberg in the presence of several Fagaceae
species (deciduous and evergreen) and rather common twig fragments
of the conifer Cunninghamia. Fan palms are present from Linz but
their remains are rare. Though deciduous woody angiosperms from
Linz are neither common nor diverse and, moreover, poorly preserved,
Acer, ?Alnus,Fagus,Ulmus, and ?Zelkova are documented, unlike at
the Rauenberg locality.
The rich early Oligocene record from Northern Bohemia and
northeastern parts of Germany includes lowland assemblages often
connected to lignite formation, e.g., Haselbach-Serie, Weisselster
Basin near Leipzig (Mai and Walther, 1978), and volcanic environ-
ments, e.g., Kundratice (Kvaček and Walther, 1998), Bechlejovice
(Kvaček and Walther, 2004), Seifhennersdorf (Walther and Kvaček,
2007). The lowland record primarily reflects a wide range of wetland
environments ranging from fluvial to swampy habitats, while the plant
assemblages, especially from maar deposits, provide insights into the
mesophytic (i.e., zonal) vegetation. The much higher diversity of de-
ciduous taxa in the record of Northern Bohemia and adjacent regions
is remarkable in comparison to the flora of Rauenberg and other fos-
sil plant sites in Hungary and Austria along the southern and western
coasts of ancient Europe. This phenomenon may be partially related
to differences in depositional processes, i.e., longer transport possibly
connected to higher water energy may be assumed for the plant re-
mains in marine deposits. Nevertheless, differences between the fos-
sil floras from the Rhine Graben and the northern Paratethys region
on the one hand and from North Bohemia and northeastern parts of
Germany on the other hand are consistent. This implies not only a
north-south vegetational gradient (Boreal Province versus Paratethys
Province), but the strong floristic affinities of the floras from the Rhine
Graben and those from the Paratethys compared to the sites in Bo-
hemia and northeastern Germany also indicate a gradient from more
continental regions to environments close to marine coasts along the
Paratethys and Rhine Graben seas.
As with the flora, the avifauna shows affinities to coeval coastal
localities. The procellariiform Rupelornis is known from contempora-
neous Rupelian localities in Belgium and from the early Oligocene of
France, and appears to have been an abundant seabird in the epiconti
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx 17
nental seas covering Europe in the early Oligocene. Colymboides?
metzleri was likewise reported from the early Oligocene Rupelian stra-
totype in Belgium (Mayr and Smith, 2013). The landbird faunas are
similar to Rupelian sites in Poland, which also yielded Eurotrochilus
fossils and passerines.
6.3. Paleoenvironmental reconstruction
6.3.1. Marine environment
Many sources of evidence support fully marine salinity levels at
Rauenberg during deposition of the Hochberg Member. Following
Martini (1978: 160), the calcareous nannoplankton Reticulofenstra
dictyoda is found only in sediments deposited under fully marine con-
ditions, but small specimens of Reticulofenstra are also found in lay-
ers with reduced salinity. All molluscan species recovered were fully
marine. Species that could tolerate reduced salinity (e.g., Neuffer,
1984) have not been recovered. Likewise, echinids can only live un-
der normal saline conditions (e.g., Prothero, 2013). According to the
life habits of adult individuals of extant comparative species, all fish
taxa from Rauenberg are found in fully marine habitats (Suppl. Fig.
3A). Due mainly to different drift times, Lutz (1997) suggested that
the taphocoenosis of terrestrial insects in marine sediments can be
a good indication for salinity, and proposed an index using the rel-
ative frequencies of the delicate Diptera + Hymenoptera (DH), ro-
bust Coleoptera + Heteroptera (CH), and other orders (AO). The re-
spective values for the Rauenberg locality (N.ges. = 221 = 100%;
DH = 30 = 13.6%; CH = 179 = 81%; AO = 12 = 5.4%) result in a cal-
culated salinity of S = 0.7%, similar to the present-day Baltic Sea [for-
mula: S = (100 − yDH − yCH)∗1.304‰= yAO ∗1.304‰]. While this
value may not be very reliable, mainly because of the large percentage
of undeterminable insects, the result does cast some doubt on the gen-
eral applicability of the formula.
Following deposition of the Hochberg Member, fully marine con-
ditions are hypothesized to have persisted during the early deposi-
tion of the Meletta Beds, followed by increasing brackish influence
22.214.171.124. Water temperature
As with salinity, an interpretation of subtropical-tropical condi-
tions during the Rupelian is well-supported for Rauenberg. The di-
noflagellate Tectatodinium pellitum –a neritic species –indicates
warm water conditions (after Head and Nøhr-Hansen, 1999). Among
the gastropods, both aporrhaids and batillariids also prefer warm-tem-
perate to tropical waters (Ozawa et al., 2009; Saul and Squires, 2015).
Most extant comparative fish taxa occur in subtropical and/or tropi-
cal regions and seas (Micklich and Hildebrandt, 2010; Micklich and
Parin, 1996) (Suppl. Fig. 3B). Very few of the taxa from the Unterfeld
clay pit indicate colder waters. Exceptions include cods of the genus
Palaeogadus and left and righteye flounders of the genera Oligo-
scophthalmus and Oligopleuronectes. A few others (e.g., ponyfishes
of the genus Leiognathoides) may represent tropical conditions. Con-
sistent with the other groups, cheloniids (Mrosovsky, 1980) and sire-
nians (Domning, 2001) spend most of their life in tropical and sub-
tropical climatic zones. Isotopic analysis indicates water temperatures
of ~ 13.6 °C winter/17.3 °C summer at 30–40 m depth for the Mainz
Basin, 65 km to the north (Walliser et al., 2015).
Unlike temperature and salinity, interpretations of water depth at
Rauenberg are widely variable. Based on the high percentage of ju-
venile forms as well as on the excellent preservation of the fishes,
Weiler (1966) assumed that the fish fauna of the Wiesloch-Rauen-
berg Tertiary block was indicative of a shallow, nutrient-rich bay,
not far from the shoreline. This hypothesis was accepted in several
more recent publications (Micklich, 2005; Micklich and Hildebrandt,
2010; Micklich et al., 2009). However, Grimm et al. (2002), based
on the presence of certain benthic forams, concluded that the fos-
sil-bearing sediments must have been deposited at the bottom of an
approximately 200 m-deep marginal basin of the Rhine Graben sea,
which was separated from the main seaway by a submarine ridge.
Micklich and Hildebrandt (2010) favored the idea of upward trans-
port of the forams (e.g., by upwelling waters) over the downward
transport of the fishes because the excellent preservation of many
fish specimens excludes long-distance transport from coastal waters
into distal deeper ones. The fish fauna lacks typical mesopelagic
forms like bristlemouths (Gonostomatidae), lanternfishes (Myctophi-
dae), lightfishes (Phosichthyidae) and marine hatchetfishes (Sternop-
tychidae), all of which are well-represented in other fossil fish as-
sociations of similar age (e.g., Gregová, 2013). Although to be used
with caution (for discussion, see Micklich and Hildebrandt, 2010), a
comparison with presumably related extant taxa shows that the ma-
jority (over 80%) of the Rauenberg forms are indicative of shal-
low water depths (< 50 m). Only a few (~ 30%) entered depths be-
low 200 m (Suppl. Fig. 3C). Of the latter, some extant representa-
tives (e.g., Gempylidae, Trichiuridae) also approach the surface wa-
ters, either occasionally, during their diurnal migration at night, or
with upwelling waters (e.g., Nakamura and Parin, 1993). Basking
sharks (Fig. 6A) also cruise in shallow coastal waters during spring-
time. With a presumed relationship to the extant Sladenia, the new
lophiid genus and species is one of the few taxa which may indicate
a depth below 900 m. However, it is also related to the genus Caruso
from the Eocene of Monte Bolca in northern Italy (Carnevale and
Pietsch, 2012), which represents a shallow lagoonal environment (G.
Carnevale, pers. comm.). In contrast, halfbeaks of the genus Hemi-
ramphus, pipefishes of the genera Syngnathus and Microphis, and
shrimpfishes of the genus Aeoliscus (Fig. 6E) occur almost exclu-
sively in superficial water layers, and the tholichthys larvae of butter-
flyfishes (Fig. 7F) are neritic or oceanic and occur in the uppermost
water layers, probably in the neuston (Leis, 1989). The vast majority
of the Unterfeld clay pit fish fauna –almost 90% according to exca-
vations conducted in 2009 (Micklich and Hildebrandt, 2010) –con-
sists of small herrings and shrimpfishes, whose extant relatives inhabit
shallow waters. In addition, as already pointed out by Weiler (1966),
the Rauenberg teleostean fauna is dominated by postlarval, early ju-
venile and juvenile forms (Fig. 7). Most often, these stages are char-
acterized by lifestyles which differ from those of the adults. Shal-
low water habitats like seagrass meadows, mangrove swamps, coastal
lagoons and estuaries, as well as intertidal zones are well known
as nursery grounds for a large variety of fish species, even pelagic
species and those inhabiting deeper waters as adults (e.g., Cocheret
de la Mornière et al., 2002; Fujii and Noguchi, 1993; Griffiths, 2002;
Honda et al., 2013; Hutchings et al., 2002; Laegdsgaard and Johnson,
2001; Sinovčićet al., 2004). Juvenile stages of Scombridae, Trichiuri-
dae, and Stromateidae are also known from trawling samples in har-
bor areas (e.g., Charleston Harbor; N. Micklich, pers. observ.). Also,
it is important to note the presence of pregnant sharks, which also
prefer coastal waters as nursery grounds (McCandless et al., 2007).
One individual has been reported by Hovestadt and Hovestadt-Euler
(2010), and a second specimen is un
18 Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx
der preparation at the HLMD (N. Micklich, pers. observ.). These ob-
servations are strong arguments for a shallower water habitat rather
than for a deeper basin. A minimum water depth below fair weather
wave base (~ 20 m) is supported based on the development of a
dysaerobic-exaerobic benthic environment and indicators of a low-en-
ergy sedimentary regime (see below). However, the strong dominance
of beetles is typical of insect taphocoenoses from deeper water de-
posits (Lutz, 1997; Wedmann et al., 2009). In addition, sedimentolog-
ical parameters such as the presence of a distal turbidite in Bed 21 and
dolomite rhombs, which form during early diagenesis in water deeper
than 100 m in almost every bed, suggest depths greater than ~ 80 m.
According to the life habits of adults of extant comparative species,
most fishes (about 30%) show affinities for reef environments, how-
ever almost 39% have a pelagic lifestyle. Although this was certainly
different for post-settlement larvae and juveniles, it nevertheless may
indicate an offshore connection. This appears to be substantiated by
the presence of tholichthys larvae (these larvae increase in abundance
from nearshore to offshore in the Hawaiian Islands (Leis, 1989)) and
of some larger individuals of pelagic sharks and scombrids. However,
the larvae may not have actively entered the area from offshore, but
could have been occasionally washed in by currents or during storms.
The sharks and scombrids may either have been washed in as car-
casses, or actively entered the area in search of nursery grounds or
prey (Weiler, 1966).
The lack of any identifiable aquatic insects (with the exception of
the four damselflies) and aquatic insect larvae (contra Micklich and
Hildebrandt (2010)) suggests that the marine environment was suffi-
ciently distant from the shore that no such insects could be washed in
via fluvial sources (in contrast to Frey et al. (2010), who hypothesized
that the sediments were deposited in the immediate vicinity of the
shoreline close to a river mouth based on the recovery of the terrestrial
mammal Apterodon). The occurrence of cerambycid and buprestid
beetles could indicate that the distance to the shore was not too great,
as these families are rarely recorded as flying at high altitudes or over
the open sea (Rust, 1999: 274), however only four specimens were re-
covered. Likewise the possible insect larvae (9 specimens) and three
arachnids would exclude aerial transportation and suggest a closer dis-
tance to the shore, although rafting on plants remains a possibility.
Blooms of calcareous nannoplankton, mainly of smaller calcareous
nannoplankton, occur in the section. Thin calcareous layers are formed
biogenetically during mass occurrences of calcareous nannoplankton
and benthic foraminifera due to episodic plankton blooms. The ben-
thic foraminifera show the same blooms as the calcareous nanno-
plankton. Mass layers of the foram Stilostomella ewaldi occur on the
top of the calcareous laminae because mass occurrences of benthic
foraminifera are slightly delayed relative to phytoplankton blooms.
Such a concentration of planktonic material is typical for larger es-
tuaries and upwelling zones. In the Upper Rhine Graben, an inland
sea, these blooms were probably caused by the appearance of nitrogen
(ammonia, nitrite and nitrate) and phosphorus, the latter due to rem-
ineralization of organic matter (Grimm et al., 2002).
126.96.36.199. Substrate (benthos)
We divided the foraminifera into four ecological groups, follow-
ing Jones and Charnock (1985). Forams which lived semi-infaunally
to infaunally are most abundant, free-living epibenthic forms typical
of a soft-bottom association are also present. Fixosessile forams are
rare because of the lack of hard ground. Both the wedge-shaped ir-
regular echinoids (Ova sp.) and the crab Coeloma are adapted for bur
rowing in soft sediment (McNamara and Philip, 1980), which is in
agreement with the clayey substrate at the locality. Aporrhaid and
batillariid gastropods are also most commonly found on fine-grained
sandy mudflats on continental margins (Ozawa et al., 2009; Saul and
Squires, 2015). The long spines projecting from the aperture of apor-
rhaids are considered an adaptation to avoid sinking in soft sediment
(Gründel et al., 2009).
The good preservation of the coccospheres indicates a lack of bot-
tom currents. A large portion of the bivalves (over 80%) are preserved
in life position, as are the echinids, which is another indication of a
very calm sedimentary environment with little bioturbation. Taphon-
omy of the cheloniids suggests in situ decay and disarticulation, and
also implies a calm environment with low rates of sedimentation.
Due to a highly specialized diet, the presence of the sirenian
“Halitherium”is an indirect but highly suggestive indicator of sea-
grass meadows in the region (Reich et al., 2015; Vélez-Juarbe, 2014).
However, given the rarity of sirenian remains and the absence of
seagrass macrofossils, these may have been located adjacent to the
Rauenberg locality, or “Halitherium”had a less restrictive diet than
188.8.131.52. Oxygen (benthos)
The dinoflagellate Thalassiphora pelagica is found in large quan-
tities in the section, indicating poorly oxygenated bottom water (af-
ter Pross, 1997). Some of the foram species (Bolivina beyrichi,
Bathisyphon tauriensis,Cyclammina placenta,Stilostommella
ewaldi), the possibly chemoautotrophic bivalve Thyasira benedeni, as
well as preservation of organic matter in the cheloniids are also in-
dicative of low (dysaerobic to exaerobic) oxygen conditions. How-
ever, endobenthic species are present at relatively high abundance
among the bivalves and echinoderms, suggesting long-term oxic con-
ditions for the beds in which they occur. In addition, almost 30% of the
fishes are demersal, supporting at least periodic oxygenation of bot-
tom waters. Examples include the eagle ray Weissobatis, and the flat-
fishes Oligoscophthalmus (Fig. 7L) and Oligopleuronectes (Hovestadt
et al., 2010; Sakamoto et al., 2003, 2004). Deutschenchelys, referred
to Moringuidae (spaghetti eels), shows morphological traits which in-
dicate a burrowing way of life (Prokofiev, 2012).
6.3.2. Terrestrial conditions
Numerous plant taxa, such as Ceratozamia floersheimensis, Lau-
raceae, Sloanea, and various palms, indicate a largely frost-free tem-
perature regime and warm climatic conditions. The flora of Rauen-
berg yields many small-leaved taxa (prevailing leaf size class mi-
crophyll, i.e., 2.25–20.25 cm2) which raises the question of seasonal-
ity in precipitation. Climate is estimated based on the closest mod-
ern analog for the flora of Rauenberg, which is the southern portion
of the modern distribution of evergreen sclerophyllous broad-leaved
forests (sensu Wang, 1961) in SE-Asia and on the climatic require-
ments of the most similar living relatives of taxa such as Ceratoza-
mia,Craigia,Platanus neptuni, and Sloanea (Kovar-Eder, submitted
for publication). The climate may have been comparable to condi-
tions in SE-Asia and America around 20°N latitude today, and is
estimated as follows: mean annual temperature 19–24 °C, mean an-
nual precipitation 1300–1700 mm, mean temperature of the warmest
month 28–29 °C, mean temperature of the coldest month 8–14 °C,
mean precipitation of the wettest month > 230 mm, that of the driest
month 18–38 mm, with the warm period being wetter than the cold
one. These data indicate seasonality in temperature and precipitation.
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx 19
In other words, the climate estimates correspond to a Cfa climate
with tendencies towards a relatively dry season in winter (Cw climate)
(transient to tropical monsoon (Am) or tropical winter-dry (Af) cli-
mate sensu Köppen, 1936, Peel et al., 2007). Rare and rather small re-
mains of charcoal indicate wildfires probably caused by lightning. It
is impossible to estimate frequency or magnitude of such events from
these minor remains.
These results differ from those derived for the flora of Flörsheim
(mean annual temperatures of 16–18 °C, mean coldest month tem-
perature 6–10 °C, mean warmest month temperature 25–28 °C, mean
sum of annual precipitation 1000–1300 mm: Pross et al., 1998). The
considerable discrepancies in the estimates may result from the fact
that the estimates for Flörsheim are mainly based on the pollen record,
which does not allow for a taxonomic resolution below the family or
Pross and Schmiedl (2002) interpreted changes in the dinoflagel-
late cysts assemblage as resulting from a long-term paleoenvironmen-
tal change from drier to more humid conditions during the deposition
of the Fischschiefer. At present, a lack of stratigraphic control pre-
vents our addressing this possibility.
184.108.40.206. Terrestrial environment
We include the palynological results of Sittler (1965), since this re-
mains the only available study of pollen from the Hochberg Member.
However, these results derive from cores from the Rhine Graben to the
west of Rauenberg and no modern revision has been attempted.
The land plants from Rauenberg represent an allochthonous assem-
blage. The plant material was transported prior to fossilization, as in-
dicated by fragmentation and fungal degradation, as well as by the
fact that predominantly robust plant parts are preserved and numerous
taxa are documented by few or single remains. Most likely only a very
limited percentage of the source flora is preserved, which hampers at-
tempts to reconstruct the coastal paleoenvironment.
The presence and relative high diversity of Pinus (6 species by
brachyblasts, 2 by cones), palms (5), and Myricaceae (4) in the macro-
fossil record may indicate a forest community (coastal pine forests)
that possibly developed on sandy soils near the coast. The pollen
record includes Pinus (2 types, together making up 8.5%) and Myri-
caceae (4 types), but no palm pollen. Daphnogene cinnamomifolia
and/or Laurophyllum pseudoprinceps possibly also occurred in this
community. Platanus neptuni may have grown in diverse habitats both
zonal and azonal, including coastal environments as well as along
streams and on river banks. Along with the robustness of its foliage,
this may explain its dominance in the plant assemblage of Rauenberg
(Suppl. Table 2). Carya and Populus may also have been elements of
riparian habitats. Pollen of Betula,Pterocarya, and Ulmus indicate a
higher diversity of deciduous taxa than the macrofossil record.
We suspect that the majority of the Lauraceae, Ceratozamia,
and others (macrofossil record) and Carpinus,Liquidambar, and Ul-
mus (pollen record) represent mesophytic forests, i.e. zonal vegeta-
tion. For species such as, e.g., Daphnogene cinnamomifolia or Lau-
rophyllum pseudoprinceps, as well as species of Craigia,Myrica,Be-
tula,Carpinus,Engelhardia,Liquidambar,Pterocarya, and Ulmus,
we assume rather wide ecological tolerance, so that they were not
necessarily bound to a specific habitat but may have flourished in
diverse environments. Both the macrofossil and pollen record lack
evidence of nearby swampy environments, including mangroves. In
these forests, the number of evergreen woody taxa distinctly exceeded
that of deciduous ones. Zoophilous woody angiosperms provided food
mainly for insects, and flowering plants with endozoo
chorous as well as dyschorous fruit vectors offered a diversified diet
for the terrestrial fauna (Suppl. Table 2). The Integrated Plant Record
Vegetation Analysis (Kovar-Eder et al., 2008), outecology of most
similar living relatives as well as the sociological analysis indicates
evergreen sclerophyllous broad-leaved forests sensu Wang (1961) as
the most likely zonal vegetation type (Kovar-Eder, submitted for
Most of the landbirds from Rauenberg are small arboreal birds,
consistent with the interpretation of a forested coastal environment.
Only one presumed predominantly terrestrial taxon has been found,
the buttonquail Turnipax, which is likely to have been a granivorous
bird. Idiornithidae, extinct relatives of the South American seriemas
which are very abundant in early Oligocene localities of the Quercy
fissure fillings in France, are notably absent from Rauenberg (Mayr,
2009b). These terrestrial birds are assumed to have been denizens of
open areas, and their absence in Rauenberg can be attributed to the
absence of suitable open habitats. Among the insects, the absence
of Blattodea and Neuroptera, as well as the rarity of Orthoptera and
Auchenorrhyncha weakly supports the interpretation of a warm-tem-
perate and humid forest rather than an arid savannah habitat near
the coast. However, no proxies for warm climate (e.g., Mantodea or
Isoptera) could be identified among the entomofauna, with a putative
earlier record of Mantodea (Monninger and Frey, 2010) having been
Based on its elongated bill and presumed hovering capabilities, the
hummingbird Eurotrochilus was probably nectarivorous, like its ex-
tant relatives. The presence of this taxon in Rauenberg implies the oc-
currence of ornithophilous plants that evolved bird-pollinating mor-
phologies (see Suppl. Table 2). Oligocolius was frugivorous, and the
trogon and the piciform Rupelramphastoides were either insectivorous
and/or frugivorous. The presence of these taxa, as well as humming-
birds, is consistent with the interpretation of relatively mild winters,
although we cannot exclude the possibility that these birds may have
Although the Rupelian assemblage from Rauenberg is composed
predominantly of marine taxa, 95 of 302 taxa currently identified
from the locality are terrestrial (31% total, or 48% of taxa represented
by macrofossils). Based on new results, we interpret the Rauenberg
assemblage as representing a fully marine assemblage deposited in
a moderately shallow, low-energy tropical-subtropical environment.
Productivity was high, and seafloor anoxia was intermittently de-
veloped. There is no evidence for long-term brackish influence or
mangrove swamps, and no direct evidence for the development of
seagrass meadows. On land, warm, frost-free but winter-dry condi-
tions permitted the development of prevailingly evergreen sclerophyl-
lous broad-leaved forests and near-coastal pine and palm-rich forests
on sandy soils. The marine invertebrate fauna shows more northerly
affinities, whereas the vertebrate fauna is distinctly Paratethyan. The
Rauenberg locality thus provides a detailed snapshot of Rupelian ma-
rine and terrestrial diversity and paleoenvironments spanning a pe-
riod of rapid environmental change. Comprehensive faunal lists, such
as the one presented here, provide important baseline data for fur-
ther comparisons of diversity or ecological structure between localities
from different time slices, as well as between coeval localities from
different latitudes or environments.
In spite of exceptional preservation allowing detailed insights into
e.g., trophic relationships, some questions remain unanswered. Avail-
able material suffers from sampling bias: historical collections fo-
cused on spectacular and decorative fossils and as such certain pale
20 Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2016) xxx-xxx
oecologically important groups may be absent or, at minimum, un-
derrepresented. A lack of stratigraphic control does not allow an as-
sessment of changing conditions at the locality, for instance involving
faunal change through time, the existence of temporary brackish con-
ditions or the relative predominance of oxic vs. anoxic benthic con-
ditions. In addition, the amount of absolute time, and hence changes
in sedimentation rate, encompassed by the locality is unknown. New
bed-by-bed excavations at the locality, initiated in 2014, promise to
address these issues. During the first year of the project > 5400 speci-
mens were collected, and accessioning and preparation are ongoing.
Thanks to the Klaus Tschira Stiftung and the municipality of
Rauenberg, who financially and logistically supported our work at the
Unterfeld clay pit. S. Staudt, K.-L. Metzger, and many unnamed stu-
dent assistants helped with excavations, and H. and A. Oechsler col-
lected a significant amount of material. Thanks to M. Keller and C.
Knopf for donation of fossils, K. Weißfor the preparation of some
bird specimens, W. Munk (SMNK) for information about the insect
material preserved at SMNK, M.C. Grimm for discussion, A. Köthe
and T. Schindler for collaboration on micropaleontology, and L. Het-
terscheid (WUR) for help in tracking down old literature. Thanks
to A. F. Bannikov (Paleontological Institute of the Russian Acad-
emy of Sciences, Moscow), G. Carnevale (Dipartimento di Scienze
della Terra, Universitàdegli Studi di Torino), and K. A. Monsch
(Naturalis, Leiden) for valuable comments regarding the fish fauna.
We are indebted to J. Dunlop (MNB) and P. Selden (Univ. Kansas)
for information on one arachnid specimen, and to J. Skartveit (NLA
Høgskolen, Bergen) for determination of the bibionid dipteres. Thanks
to M. Harzhauser and A. Kroh (Vienna) for discussions about balanids
and echinoderms. R.B.S. was supported by doctoral grant from CNPq
(Conselho Nacional de “Desenvolvimento”Científico e Tecnológico,
Brazil) (process #245575/2012-0). This paper is dedicated to the late
Frank Broghammer (former mayor of Rauenberg) and Klaus Tschira
(founder of the Klaus Tschira Foundation GmbH). Without their help
and engagement the Unterfeld excavations would not have been pos-
sible. M. Voss and an anonymous reviewer provided comments that
improved the manuscript.
Appendix A. Supplementary data
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