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Elasmobranch remains are quite common in Miocene deposits and were the subject of numerous studies since the middle of the nineteenth century. Nevertheless, the taxonomic diversity of the Marine Molasse sharks, rays and skates is still largely unknown. Here, we describe 37 taxa from the lower Miocene of the Molasse Basin: 21 taxa could be identified at species level, whereas 15 taxa could only be assigned to genus and one taxon is left as order incertae sedis. The material was collected from deposits of the Auwiesholz Member of the Achen Formation (middle Burdigalian, middle Ottnangian age, ca. 17.8 Ma) exposed near Simssee, Upper Bavaria. This faunal assemblage is a mixture of shallow marine, near-coastal, pelagic and deep-water taxa. The fauna from Simssee displays different biogeographic dynamics at local and regional scales, possibly related to the intense climatic, oceanographic and tectonic events that occurred during the Eggenburgian–Ottnangian stages. The faunal relationships of the early Miocene chondrichthyan faunas from the Mediterranean Sea and Paratethys with others regions are established on the basis of qualitative (presence/absence) data. The beta diversity (Sørensen–Dice coefficient) of the Miocene Molasse elasmobranchs was used to characterize the taxonomic differentiation between localities and regions. According to our results, the fauna from Simssee shows close similarities with those from Switzerland, Austria, France and northern Germany. Faunal similarities and differences are mainly related to tectonic events and oceanographic variables (i.e. migration through seaway passages) or might represent collecting biases.
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Vol.:(0123456789)
1 3
PalZ
https://doi.org/10.1007/s12542-020-00518-7
RESEARCH PAPER
Sharks, rays andskates (Chondrichthyes, Elasmobranchii)
fromtheUpper Marine Molasse (middle Burdigalian, early
Miocene) oftheSimssee area (Bavaria, Germany), withcomments
onpalaeogeographic andecological patterns
JaimeA.Villafaña1,2 · GiuseppeMarramà1,9 · StefanieKlug3 · JürgenPollerspöck4· MarkusBalsberger5·
MarceloRivadeneira6,7,8 · JürgenKriwet1
Received: 3 November 2019 / Accepted: 27 April 2020
© The Author(s) 2020
Abstract
Elasmobranch remains are quite common in Miocene deposits and were the subject of numerous studies since the middle of the
nineteenth century. Nevertheless, the taxonomic diversity of the Marine Molasse sharks, rays and skates is still largely unknown.
Here, we describe 37 taxa from the lower Miocene of the Molasse Basin: 21 taxa could be identified at species level, whereas 15
taxa could only be assigned to genus and one taxon is left as order incertae sedis. The material was collected from deposits of the
Auwiesholz Member of the Achen Formation (middle Burdigalian, middle Ottnangian age, ca. 17.8Ma) exposed near Simssee,
Upper Bavaria. This faunal assemblage is a mixture of shallow marine, near-coastal, pelagic and deep-water taxa. The fauna from
Simssee displays different biogeographic dynamics at local and regional scales, possibly related to the intense climatic, oceano-
graphic and tectonic events that occurred during the Eggenburgian–Ottnangian stages. The faunal relationships of the early Miocene
chondrichthyan faunas from the Mediterranean Sea and Paratethys with others regions are established on the basis of qualitative
(presence/absence) data. The beta diversity (Sørensen–Dice coefficient) of the Miocene Molasse elasmobranchs was used to char-
acterize the taxonomic differentiation between localities and regions. According to our results, the fauna from Simssee shows close
similarities with those from Switzerland, Austria, France and northern Germany. Faunal similarities and differences are mainly
related to tectonic events and oceanographic variables (i.e. migration through seaway passages) or might represent collecting biases.
Keywords Chondrichthyes· Molasse· Burdigalian· Early Miocene· Paratethys· Beta diversity
Handling editor: Mike Reich.
* Jaime A. Villafaña
villafanaj88@univie.ac.at
Giuseppe Marramà
giuseppe.marrama@unito.it
Stefanie Klug
stefanie.klug@gauss.uni-goettingen.de
Jürgen Pollerspöck
juergen.pollerspoeck@shark-references.com
Markus Balsberger
mbalsberger@gmx.de
Marcelo Rivadeneira
marcelo.rivadeneira@ceza.cl
Jürgen Kriwet
juergen.kriwet@univie.ac.at
1 Department ofPalaeontology, University ofVienna,
Althanstraße 14, 1090Vienna, Austria
2 Centro de Investigación en Recursos Naturales y
Sustentabilidad, Universidad Bernardo O’Higgins, Santiago,
Chile
3 School ofScience (GAUSS), Georg–August University,
37077Göttingen, Germany
4 Bavarian State Collection ofZoology, Munich, Germany
5 Achberg 11, 83259Schleching, Germany
6 Centro de Estudios Avanzados en Zonas Áridas,
Av. Ossandon 877, Coquimbo, Chile
7 Departamento de Biología Marina, Facultad de Ciencias del
Mar, Universidad Católica del Norte, Coquimbo, Chile
8 Departamento de Biología, Universidad de La Serena,
LaSerena, Chile
9 Dipartimento di Scienze della Terra, Università degli Studi di
Torino, Via Valperga Caluso, 35, 10125Torino, Italy
J. A. Villafaña etal.
1 3
Introduction
After the Tethys Ocean had nearly completely vanished by
the end of the Eocene, an isolated Paratethys Sea developed
in the latest Eocene–earliest Oligocene related to the develop-
ment of the Alpine mountains (Baldi 1980; Rusu 1988; Rögl
1999). This island chain acted as barrier partly separating
the Paratethys from the Mediterranean Sea. The Paratethys
extended from the Rhone Valley in the east towards Inner
Asia. Marginal to the Paratethys, the Molasse Basin, which
represents a foreland basin, developed in the Oligo–Miocene
during the Alpine–Himalayan orogeny. The Molasse Basin of
southern Germany was thus part of western Paratethys during
the Miocene.
The Paratethys and Mediterranean seas experienced dramatic
changes during their development (Rögl 1999; Piller etal. 2007;
Sant etal. 2017). During the Eggenburgian (lower Burdigalian),
a broad sea passage between the Paratethys Sea and the Indo-
Pacific Ocean was open, providing optimal environmental condi-
tions for marine faunas and opportunities for widespread faunal
exchanges (Rögl 1999). Additionally, a seaway passage through
the Alpine fore-deep between the Mediterranean and Paratethys
was open. Later, during the Ottnangian (middle Burdigalian),
the sea passage into the Indo-Pacific Ocean was closed due the
collision of Africa and Arabia with the Anatolian plate. The con-
nection between the Western–Central Paratethys and the Medi-
terranean seas still persisted through the Rhine Graben, but the
eastern Paratethys was already isolated (i.e. it informed the so-
called Kotsakhurian Sea). All these events also induced changes
in sea levels, salinity and temperature (Haq etal. 1988). Studies
based on early Miocene marine invertebrates of Europe indicate
that these intense climatic and oceanographic events had impor-
tant effects on diversification patterns of organisms (Kroh 2007).
Remains of sharks, rays and skates generally are quite com-
mon in Miocene sediments of the Paratethys (Barthelt etal.
1991; Kocsis 2007; Reinecke etal. 2011; Schultz 2013; Poller-
spöck and Straube 2017; Szabó and Kocsis 2016; Underwood
and Schlögl 2013), but despite all progress accomplished in
the last decades, our understanding of Miocene elasmobranchs
taxonomic diversities and faunal relationships remains very
incomplete. Here, we document an elasmobranch assemblage
from the lower Miocene Upper Marine Molasse of the western
Paratethys and present results about their relationships with
other faunas from the early Miocene.
Materials andmethods
Data collection
A total of 466 elasmobranch specimens (including teeth
and tail spines) were recovered by screen washing and sur-
face collecting from several points along the Auwiesholz
Member of the Achen Formation in the Simssee area
(Bavaria, S. Germany, Fig.1), during several trips con-
ducted by two of the authors (JP and MB) and other
collaborators in the late 90s. Part of the material was
collected by N. Rückert-Ülkümen (Bavarian State Col-
lection of Palaeontology and Geology) near the village
of Hirnsberg in 1993. The precise stratigraphic origin of
all material, however, remains ambiguous. The Achen
Formation is of middle Ottnangian age (middle Burdiga-
lian, early Miocene, ca. 17.8Ma) and denotes the second
cycle of the Upper Marine Molasse in Bavaria (Pippèrr
etal. 2007). The depositional setting represents an inner
neritic environment basally, which deepens upwards. This
interpretation was based on the presence of foraminifera
(e.g. Ammonia) and ostracoda (e.g. Cytheretta), and ver-
tebrates as bony fishes (e.g. Rhynchoconger) were also
reported from this member. The elasmobranch fossil fauna
described herein is almost coeval to the assemblage from
the Baltringer Horizon in Baden-Württemberg, SW Ger-
many (Probst 1879).
The fossils are housed in the Bayerische Staatssa-
mmlung für Paläontologie und Geologie at Munich,
Germany, with the catalogue numbers bearing the prefix
SNSB-BSPG 2019 III.
Completeness ofthetaxonomic inventory
We estimated the completeness of Simssee taxonomic inven-
tory based on two complementary methods, assuming teeth/
spines represent different individuals. First, rarefaction was
used to estimate the impact of sampling effort (specimens)
on total taxonomic richness, where a plateau in the curve
suggests a saturation trend (Sanders and Hessler 1969;
Gotelli and Colwell 2011). Second, we estimated the true
(i.e. discoverable) taxonomic richness based on the Chao
Fig. 1 Map of the Simssee area
Sharks, rays and skates from the Upper Marine Molasse (Burdigalian, Miocene) of Bavaria, Germany
1 3
1 extrapolation index (Chao and Lee 1992; Colwell and
Coddington 1994). The Chao 1 index is a non-parametric
method that estimates the taxonomic richness in a given
locality based on the number of rare taxa (i.e. with one and
two individuals), providing a point estimator and an upper
level confidence interval (95%). The completeness of the
taxonomic inventory was calculated as the fraction between
the observed richness and the extrapolated richness (point
estimator and upper level confidence interval). This method
has been previously used in paleontological studies to esti-
mate the discoverable species richness (e.g. Rivadeneira and
Nielsen 2017).
Faunal comparison
We compared the faunal composition of Simssee with other
early Miocene elasmobranch faunas worldwide by using a
comprehensive literature dataset (see references in Table2).
Analyses were carried out at the genus level and pooling
nearby localities to reduce identification and sampling
biases. Since some localities lack precise age estimations,
we only used localities that could be assigned to the Eggen-
burgian–Ottnangian time span (i.e. lower to middle Burdiga-
lian). We used presence–absence data to estimate differences
in generic composition between Simssee and other localities
using the Sørensen–Dice similarity index. This index has
been widely recommended in the ecological and paleonto-
logical literature due to its statistical properties (Hubalek
1982; Murguía and Villaseñor 2003; Hammer and Harper
2006; Jost etal. 2011). However, the Sørensen–Dice index,
as any presence–absence similarity index, is sensitive to the
completeness of taxonomic inventories (Jost etal. 2011).
Systematic palaeontology
Class Chondrichthyes Huxley, 1880
Subclass Elasmobranchii Bonaparte, 1838
Infraclass Neoselachii Compagno, 1977
Order Hexanchiformes Buen, 1926.
Family Chlamydoselachidae Garman, 1884
Genus Chlamydoselachus Garman, 1884
Type species. Chlamydoselachus anguineus Garman, 1884
Chlamydoselachus sp.
Figure2a, b
Material. One antero-lateral tooth—SNSB-BSPG 2019
III-1.
Description. The antero-lateral tooth displays a one pre-
served long and slender cusp, which is lingually bent
(Fig.2a, b). The crown surface is mostly smooth, but faint
vertical folds are present at the base of both cusp faces. In
profile view, the lingual face is concave, whereas the labial
face is convex. Although the root is incomplete and abraded,
the small nutritive foramen is still distinguishable.
Remarks. The frilled shark Chlamydoselachus is currently
represented by two species: C. africana which occurs in the
Southeastern Atlantic, and C. anguineus, reported from the
western Indian, Eastern Atlantic and Pacific Ocean (Uyeno
etal. 1983; Ebert and Compagno 2009). The fossil record
of Chlamydoselachus extends back to the Upper Cretaceous
(Kriwet etal. 2016; Cappetta 2012). Early Miocene records
of Chlamydoselachus were reported from Austria (Pfeil
1983; Schultz 2013), Germany (Barthelt etal. 1991), and
the USA (Phillips etal. 1976).
As the single tooth is abraded and incompletely pre-
served, it is not possible to identify the specimen at spe-
cific level.
Family Hexanchidae Gray, 1851
Genus Notorynchus Ayres, 1855
Type species. Notorynchus maculatus Ayres, 1855
Notorynchus primigenius (Agassiz, 1843)
Figure2c–l
Material. Two upper anterior teeth—SNSB-BSPG 2019 III-
2, SNSB-BSPG 2019 III-3; 2 upper antero-lateral teeth—
SNSB-BSPG 2019 III-4, SNSB-BSPG 2019 III-5; 12 lower
antero-lateral teeth—SNSB-BSPG 2019 III-6, SNSB-BSPG
2019 III-7 (4 teeth), SNSB-BSPG 2019 III-8 (7 teeth); 1
commissural tooth—SNSB-BSPG 2019 III-9 and 1 lower
symphyseal tooth—SNSB-BSPG 2019 III-10.
Description. The upper anterior teeth have a slender and tri-
angular cusp that is distally oriented (Fig.2c, d). The cutting
edges are smooth and do not reach the base of the crown.
The root is high and rounded with a convex outline in labial
and lingual views.
The upper antero-lateral teeth are labio-lingually com-
pressed, but narrower than the lower teeth (Fig.2e, f). The
crown is composed of a main cusp followed by a secondary
cusp that is distally oriented. The root is high and flat with
almost straight basal aspect.
The lower antero-lateral teeth are labio-lingually com-
pressed and mesio-distally elongated (Fig.2g, h). The
crown shows a comb-like shape with triangular cusps,
which are distally oriented. The most complete specimen
J. A. Villafaña etal.
1 3
bears six cusps decreasing in size distally. The mesial cut-
ting edge is slightly convex with awl-shaped coarse ser-
rations that become finer towards the base of the crown.
The root is incomplete, being low and flat with rectilinear
basal part.
The symphyseal tooth is mesio-distally compressed with
a main straight cusp flanked by two mesial and three distal
secondary cusplets (Fig.2i, j). The mesial secondary cus-
plets are mesially oriented, whereas the distal ones are dis-
tally oriented. The root is low and possibly abraded. The
commissural tooth displays a very low crown, mesio-distally
elongated (Fig.2k, l). The distal part of the crown is miss-
ing. The root is high and flat.
Fig. 2 Hexanchiformes. a, b Chlamydoselachus sp., antero-lateral
tooth. cl Notorynchus primigenius: c, d anterior tooth, e, f upper
antero-lateral tooth, g, h lower lateral tooth, i, j lower symphyseal
tooth, k, l commissural tooth. Labial a, c, e, g, i, k; lingual b, d, f, h,
j, l. Scale bar 2mm
Sharks, rays and skates from the Upper Marine Molasse (Burdigalian, Miocene) of Bavaria, Germany
1 3
Remarks The genus Notorynchus is nowadays only rep-
resented by the broadnose sevengill shark N. cepedianus,
which is a cosmopolitan species occurring in warm tem-
perate to subtropical seas, but which is absent in the North
Atlantic and Mediterranean Sea (Compagno etal. 1989; Last
and Stevens 2009). having teeth described here similar to N.
primigenius from the early Miocene of Germany (Reinecke
etal. 2011), we determine them as belonging to this species,
whose fossil record ranges from the Oligocene to the Mio-
cene (Cappetta 2012). Early Miocene records of N. primi-
genius were also reported from Austria (Schultz 2013), Ger-
many (Probst 1879 as Notidanus primigenius; Lutzeier 1922;
von Ihering 1927; Barthelt etal. 1991; Scholz and Bienert
1992; Baier etal. 2004; Höltke 2014; Sach 2016), France
(Cappetta 1970, 1973, as Hexanchus primigenius), Hungary
(Kordos and Solt 1984, as Hexanchus primigenius; Kocsis
2007), Slovakia (Holec etal. 1995), Switzerland (Leriche
1927; Fischli 1930, as Notidanus primigenius; Bolliger etal.
1995) and the USA (Kent 2018).
Order Squaliformes Goodrich, 1909
Family Squalidae Bonaparte, 1838
Genus Squalus Linnaeus, 1758
Type species. Squalus acanthias Linnaeus, 1758
Squalus sp.
Figure3a, b
Material. Two lower antero-lateral teeth—SNSB-BSPG
2019 III-11, SNSB-BSPG 2019 III-12.
Description. These teeth show a broad, triangular and
strongly distally bent cusp with a rounded apex (Fig.3a, b).
The distal heel is high with a convex outline. The distal and
mesial cutting edges are completely smooth. In labial view,
a long apron is present. In lingual view, the uvula is abraded
but still distinguishable. The root is short.
Remarks. The genus Squalus is currently represented by 35
species worldwide distributed, including the Mediterranean
Sea (Pollerspöck and Straube 2019). Squalus species inhabit
the upper continental and insular slopes at tropical to tem-
perate latitudes. The fossil record of Squalus extends back to
the Upper Cretaceous. Early Miocene records were reported
from Austria (Schultz 2013), Chile (Suarez etal. 2006; Vil-
lafaña etal. 2019), France (Ledoux 1972), Hungary (Kocsis
2007), Germany (Probst 1879; von Ihering 1927; Barthelt
etal. 1991; Reinecke etal. 2008; Pollerspöck and Beaury
2014; Pollerspöck and Straube 2017), India (Mondal etal.
2009), Slovakia (Holec etal. 1995), Switzerland (Bolliger
etal. 1995) and the USA (Emry and Eshelman 1998; Purdy
etal. 2001; Kent 2018).
Fig. 3 Squaliformes. a, b Squalus sp., antero-lateral tooth, c, d Centrophorus granulosus, lower antero-lateral tooth, e, f Deania sp., lower
antero-lateraol tooth, g, h Isistius triangulus, lower anterior tooth. Labial a, c, e, g.; lingual b, d, f, h. Scale bar 1mm
J. A. Villafaña etal.
1 3
The teeth reported here differ from those previously
described from Germany (Reinecke etal. 2011; Poller-
spöck and Straube 2017). In those specimens, the apex of
the crown is very acute and the apron is narrower. However,
differences in the material described here could be related
also to a taphonomic effect. Therefore, as the teeth are very
abraded and lack diagnostic characters, our specimens can
be only identified at the genus level.
Family Centrophoridae Bleeker, 1859
Genus Centrophorus Müller and Henle, 1837
Type species. Squalus granulosus Bloch and Schneider, 1801
Centrophorus granulosus (Bloch and Schneider, 1801)
Figure3c, d
Material. Eleven lower antero-lateral teeth—SNSB-BSPG
2019 III-13, SNSB-BSPG 2019 III-14 (2 teeth), SNSB-
BSPG 2019 III-15 (8 teeth); and 3 upper lateral teeth—
SNSB-BSPG 2019 III-16, SNSB-BSPG 2019 III-17 (2
teeth).
Description. The lower antero-lateral teeth are labio-lin-
gually compressed with a broad and distally inclined cusp
(Fig.3c, d). The mesial edge is faintly sigmoidal and ser-
rated, whereas the distal edge is slightly convex and smooth.
The distal heel is notched and convex without any serrations.
The apron is long and broader at its base with a rounded
end. On the lingual face, a short uvula with a deep infun-
dibulum is present just below its lower extremity. The distal
part of the root is larger than the mesial one and displays
some foramina. In labial view, the mesial part of the root
displays a large foramen and has a concave contour. In upper
lateral teeth, the crown is higher than broad. The mesial edge
is slightly sigmoidal and serrated. The distal heel is short
and strongly convex with weak serrations. The root is very
abraded and covered by sediment. Apron, uvula and root
foramina are not distinguishable.
Remarks. The genus Centrophorus is currently represented
by 13 species with global distributions (Pollerspöck and
Straube 2019). These mid- to deep-water sharks inhabit
tropical to temperate environments (Compagno etal. 2005;
White etal. 2013). The fossil record of Centrophorus
extends back into the upper Cretaceous (Cappetta 2012).
Early Miocene records of Centrophorus were reported from
Austria (Pfeil 1983; Schultz 2013), Colombia (Carrillo-
Briceño etal. 2016a), Germany (Probst 1879; Fischli 1930;
Barthelt etal. 1991; Scholz and Bienert 1992; Baier etal.
2004; Pollerspöck and Beaury 2014; Sach 2016; Pollerspöck
and Straube 2017), Switzerland (Bolliger etal. 1995; Jost
etal. 2016), Slovakia (Holec etal. 1995 as Squalus sp.) and
the USA (Phillips etal. 1976).
According to Vialle etal. (2011), the serrated mesial
cutting edge and the absence of folds on the uvula are
the diagnostic characters that separate C. granulosus from
other species. Thereby, these characters that are present in
our material allow us to identify it at species level.
Genus Deania Jordan and Snyder, 1902
Type species. Deania eglantina Jordan and Snyder, 1902
Deania sp.
Figure3e, f
Material. One lower tooth—SNSB-BSPG 2019 III-18.
Description. The lower tooth is labio-lingually compressed
with a short, narrow and distally oriented cusp (Fig.3e, f).
The cutting edges are completely smooth. The mesial cut-
ting edge is convex in its lower part and slightly straight in
its upper part. The distal cutting edge is slightly convex.
The distal heel is convex and serrated. The apron is short
and broad, reaching the base of the root. In lingual view,
the uvula is short. The root is short, abraded and covered
with sediment; thus, the infundibulum and foramina are not
distinguished.
Remarks. The genus Deania is currently represented by four
species (D. calcea, D. hystricosa, D. profundorum and D.
quadrispinosa) occurring in the Atlantic, Indian, and Pacific
oceans, but not in the Mediterranean Sea (Compagno 1984a;
Akhilesh etal. 2010). The fossil record of Deania extends
back into the lower Paleocene (Cappetta 2012). Early Mio-
cene records were only reported from Austria (Pfeil 1983;
Schultz 2013; Pollerspöck etal. 2018), Germany (Poller-
spöck and Straube 2017) and Switzerland (Bolliger etal.
1995). However, according to Reinecke etal. (2011), Deania
also occurrs in different localities of the early Miocene in the
Paratethys and Mediterranean seas.
Considering the narrow cusp and the smooth mesial
cutting edge, the tooth described here can be unambigu-
ously identified as belonging to the genus Deania. How-
ever, due to the lack of diagnostic characters and the small
number of specimens available for this study, it is not pos-
sible to identify it at specific level.
Family Dalatiidae Gray, 1851
Genus Isistius Gill, 1864
Type species. Scymnus brasiliensis Quoy and Gaimard, 1824
Sharks, rays and skates from the Upper Marine Molasse (Burdigalian, Miocene) of Bavaria, Germany
1 3
Isistius triangulus (Probst, 1879)
Figure3g, h
Material. Two lower antero-lateral teeth—SNSB-BSPG
2019 III-19, SNSB-BSPG 2019 III-20.
Description. The crowns of the two antero-lateral teeth are
abraded and the root is incomplete (Fig.3g, h). The crown
is labio-lingually compressed and triangular. The cutting
edges are smooth and almost straight. The crown slightly
overhangs the root distally and mesially. The root is high,
flat and covered with sediment; thus, the median foramina
cannot be distinguished.
Remarks. The genus Isistius is currently represented by
three species: I. brasiliensis, I. labialis and I. plutodus
(de Figueiredo and de Carvalho 2018). The cookie cut-
ter shark (I. brasiliensis) has a wide geographic distri-
bution in tropical and subtropical environments, whereas
the largetooth cookie cutter shark (I. plutodus) has been
reported from the Atlantic and Northwest Pacific oceans
(Compagno 1984a; Compagno etal. 2005). The fossil
record of I. triangulus ranges from the early Miocene to
the early Pliocene (Cappetta 2012). Early Miocene records
are from Austria (Schultz 2013), France (Cappetta 1970),
Germany (Probst 1879, as Scymnus triangulus; von Iher-
ing 1927; Barthelt etal. 1991; Pollerspöck and Beaury
2014; Sach 2016; Pollerspöck and Straube 2017), Hungary
(Kocsis 2007), Portugal (Antunes and Jonet 1970), Slo-
vakia (Holec etal. 1995) and Switzerland (Leriche 1927;
Jost etal. 2016).
According to Laurito (1997, 1999), the fossil species I.
triangulus can be distinguished from the extant I. brasilien-
sis and I. plutodus by its different crown shape. In I. triangu-
lus, the crown edges form an equilateral triangle (Carrillo-
Briceño etal. 2014; Pérez and Marks 2017), whereas it is
isosceles in I. brasiliensis and I. plutodus.
Order Squatiniformes Buen, 1926
Family Squatinidae Bonaparte, 1838
Genus Squatina Dumeril, 1806
Type species. Squalus squatina Linnaeus, 1758
Squatina sp.
Figure4a, b
Material. Four lateral teeth—SNSB-BSPG 2019 III-21,
SNSB-BSPG 2019 III-22 (two teeth), SNSB-BSPG 2019
III-23.
Description. The lateral teeth are broader than high (Fig.4a,
b). The crown is rather low, slender and distally inclined.
The cutting edges are smooth. The mesial and distal heels
are elongated and low with sharp cutting edges. In labial
view, the apron is short and basally rounded. The mesial
and distal crown–root boundaries are slightly concave. The
lingual protuberance is massive with a well-developed fora-
men at its extremity. In profile view, the lingual crown face
is slightly concave, whereas the labial crown face is convex.
The root is high and broad in lingual view. In labial view,
the root face is slightly straight basally.
Remarks. The genus Squatina is currently represented by 24
species with global distributions in temperate and tropical
seas (Pollerspöck and Straube, 2019). In the Mediterranean
Sea, Squatina is represented by the angel shark (S. squatina),
the sawback angelshark (S. aculeata) (Serena 2005; Ebert
and Stehmann 2013) and the smoothback angelshark (S. ocu-
lata) (Ergüden etal. 2019). The genus extends back to the
Early Cretaceous (Klug and Kriwet 2013). Early Miocene
records of Squatina were reported from Austria (Brzobohatý
and Schultz 1971; Schultz 2013), Chile (Suarez etal. 2006;
Villafaña etal. 2019), France (Cappetta 1970, 1973; Canevet
2011), Germany (Probst 1879; Lutzeier 1922; von Ihering
1927; Barthelt etal. 1991; Scholz and Bienert 1992; Baier
etal. 2004; Reinecke etal. 2011; Pollerspöck and Beaury
2014; Sach 2016), Hungary (Kordos and Solt 1984; Kocsis
Fig. 4 Squatiniformes. a, b Squatina sp., lateral tooth, Pristiophori-
formes. ce Pristiophorus sp., rostral spine. Labial: a; lingual: b; dor-
sal: c; ventral: d; anterior: e. Scale bar 2mm
J. A. Villafaña etal.
1 3
Sharks, rays and skates from the Upper Marine Molasse (Burdigalian, Miocene) of Bavaria, Germany
1 3
2007), Peru (Landini etal. 2019), Portugal (Antunes and
Jonet 1970), Switzerland (Leriche 1927; Fischli 1930; Bol-
liger etal. 1995), Slovakia (Holec etal. 1995) and the USA
(Phillips etal. 1976; Case 1980).
Order Pristiophoriformes Berg, 1958
Family Pristiophoridae Bleeker, 1859
Genus Pristiophorus Müller and Henle, 1837
Type species. Pristis cirratus Latham, 1794
Pristiophorus sp.
Figure4c–e
Material. Two rostral spines—SNSB-BSPG 2019 III-24,
SNSB-BSPG 2019 III-25.
Description. The rostral spines are abraded and the basal
peduncle is missing (Fig.4c–e). The enameloid cap is long,
slender and dorso-ventrally flattened with smooth cutting
edges. The enameloid is smooth and devoid of any ornamen-
tation. The spine is slightly bent towards the rear.
Remarks. The genus Pristiophorus is currently represented
by seven species that are distributed in temperate and sub-
tropical regions (Compagno 2001). This genus is present
in the western Pacific, western Central Atlantic, and Indian
oceans, but not in the Mediterranean Sea (Compagno 1998;
Last and Stevens 2009; Yearsley etal. 2008). The fossil
record of Pristiophorus extends back to the lower Creta-
ceous (Cappetta 2012). Early Miocene records were reported
from Australia (Fitzgerald 2004), Austria (Schultz 2013),
Chile (Suarez etal. 2006; Villafaña etal. 2019), Colombia
(Carrillo-Briceño etal. 2016a), Germany (von Ihering 1927;
Barthelt etal. 1991; Reinecke etal. 2011; Pollerspöck and
Beaury 2014; Sach 2016; Pollerspöck and Straube 2017),
Slovakia (Underwood and Schlögl 2013) and Switzerland
(Fischli 1930; Jost etal. 2016).
The rostral spine reported here can be unambigu-
ously identified as belonging to the genus Pristiophorus.
According to Underwood and Schlögl (2013) and Engel-
brecht etal. (2017), Pristiophorus species erected on the
sole basis of rostral teeth should be considered as nomina
dubia because of the lack of specific diagnostic charac-
ters. Thereby, the oral teeth should be used as comparative
material instead of rostral spines. For this reason, identi-
fication at specific level of the single Pristiophorus spine
reported here is not possible.
Order Lamniformes Berg, 1937
Family Mitsukurinidae Jordan, 1898
Genus Mitsukurina Jordan, 1898
Type species. Mitsukurina owstoni Jordan, 1898
Mitsukurina lineata (Probst, 1879)
Figure5a, b
Material. Two upper anterior teeth—SNSB-BSPG 2019 III-
26, SNSB-BSPG 2019 III-27.
Description. The teeth have a high, slender and in profile
view slightly sigmoidal cusp with a broad base (Fig.5a,
b). In labial view, the crown surface is almost completely
smooth with only few short folds at its base. In lingual view,
strong longitudinal folds extend from its base to the middle
part of the cusp. Both faces of the crown are slightly convex
transversely. The root is missing.
Remarks. The genus Mitsukurina is represented today only
by the goblin shark M. owstoni, occurring in the Atlantic,
Pacific and western Indian oceans, but absent in the south-
eastern Pacific and Mediterranean Sea (Last and Stevens
1994; Compagno 2001). The fossil record of Mitsukurina
lineata dates back to the early Miocene of Europe (Cappetta
2012).
Early Miocene records of this species are from Austria
(Schultz 2013), France (Joleaud 1912; Cappetta 1975),
Germany (Probst 1879, as Lamna lineata; Lutzeier 1922;
Barthelt etal. 1991; Scholz and Bienert 1992; Baier etal.
2004; Höltke 2014; Pollerspöck and Beaury 2014; Sach
2016), Hungary (Kocsis 2007), Slovakia (Holec etal.
1995) and Switzerland (Bolliger etal. 1995; Jost etal.
2016).
Family Alopiidae Bonaparte, 1838
Genus Alopias Rafinesque, 1810
Type species. Alopias macrourus Rafinesque, 1810
Alopias exigua (Probst, 1879)
Figure5c, d
Material. Five lower lateral teeth—SNSB-BSPG 2019 III-
28, SNSB-BSPG 2019 III-29.
Fig. 5 Lamniformes. ad Mitsukurina lineata, a, b upper lateral
tooth, Alopias exigua, c, d lower lateral teeth, e, f Araloselachus
cuspidatus, g, l Carcharias acutissima, g, h upper lateral tooth, i, j
lower anterior tooth, k, l lower latero-posterior tooth, mp Odon-
taspis molassica, qt Carcharodon hastalis, q, r upper lateral, lower.
Labial: a, c, e, g, i, l, n; lingual: b, d, f, h, j, k, m. Scale bar 2mm
J. A. Villafaña etal.
1 3
Description. The teeth have a slender, acute and distally
inclined cusp with a broad base (Fig.5c, d). The mesial cut-
ting edge is rather straight, whereas the distal one is strongly
concave. Both cutting edges are sharp and reach the base
of the crown. The labial crown face is almost flat, whereas
the lingual face is convex. Both faces are devoid of any
ornamentation. In labial view, the crown/root boundary is
straight. A narrow and rather straight crown neck separates
the labial crown face from the root. The root lobes are short
and well divergent with rounded extremities.
Remarks. The genus Alopias is currently represented by
three species: A. pelagicus, A. superciliosus, and A. vulpi-
nus. The pelagic thresher (A. pelagicus) is distributed in
the Indo-Pacific and eastern Pacific. The bigeye thresher (A.
superciliosus) and the thresher (A. vulpinus) have a wide
distribution in tropical and temperate oceans, including the
Mediterranean Sea (Last and Stevens 1994; Compagno etal.
2005). The fossil record of A. exigua ranges from the early
Oligocene to the middle Miocene (Cappetta 2012). Early
Miocene records of A. exigua were reported from Austria
(Brzobohatý and Schultz 1971), France (Cappetta 1970),
Germany (von Ihering 1927; Barthelt etal. 1991; Baier etal.
2004; Reinecke etal. 2011; Höltke 2014; Pollerspöck and
Beaury 2014; Sach 2016) and Hungary (Kordos and Solt
1984; Kocsis 2007).
The extinct A. exigua can be distinguished from the
extant species by its narrower cusp and lower root. The
teeth described here match perfectly with those reported
from the early Miocene of northern Germany by Reinecke
etal. (2011).
Family Odontaspididae Müller and Henle, 1838
Genus Araloselachus Glikman, 1964
Type species. Araloselachus agespensis Glikman, 1964
Araloselachus cuspidatus (Agassiz, 1843)
Figure5e, f
Material. Six upper antero-lateral teeth—SNSB-BSPG 2019
III-30, SNSB-BSPG 2019 III-31, SNSB-BSPG 2019 III-32
(four teeth); and one anterior tooth—SNSB-BSPG 2019
III-33.
Description. The upper antero-lateral teeth have a triangular
and distally inclined cusp (Fig.5e, f). The mesial cutting
edge is slightly concave at its base and convex at its upper
part. In profile view, the crown is rather straight. The distal
cutting edge is concave at its base and straight in its medial
and upper parts. A pair of low lateral cusplets are present,
which are triangular and devoid of any ornamentation. The
root is low with short and well-separated lobes.
The anterior tooth (not figured) is very abraded and part
of the apex is missing. The crown is triangular, high and
robust. In profile view, the cusp also is straight. The labial
crown/root boundary is concave and the labial face over-
hangs the root in its medial part. The enameloid surface is
completely smooth on both crown faces. The lateral cusplets
are broken. The root is high with well-separated and long
lobes.
Remarks. A. cuspidatus is common in Oligocene and Mio-
cene fossiliferous sites (Cappetta 2012). Early Miocene
records are from Australia (Pledge 1967), Austria (Brzobo-
hatý and Schultz 1971), France (Cappetta 1970, 1973;
Goedert etal. 2017), Germany (Barthelt etal. 1991, as
Synodontaspis cuspidata; Sach and Heizmann 2001; Sach
2016, as Carcharias cuspidatus), Hungary (Kocsis 2007, as
Carcharias cuspidatus), Italy (Marsili etal. 2007), Portugal
(Zbyszewski 1949), Slovakia (Holec etal. 1995), Switzer-
land (Bolliger etal. 1995, as Carcharias cuspidatus) and the
USA (Case 1980; Kent 2018).
According to Cappetta (2012), the genus Araloselachus
has sufficient morphological characters to be separated from
other odontaspid genera. In A. cuspidatus, features of the
anterior teeth are used to separate the species from other
odontaspids. In this species, the teeth are more robust and
stronger, the crown is straight in profile view and the ename-
loid surface is completely smooth. The teeth reported herein
are similar to those described from the middle Miocene of
Hungary (Szabó and Kocsis 2016).
Genus Carcharias Rafinesque, 1810
Type species. Carcharias taurus Rafinesque, 1810
Carcharias acutissimus (Agassiz, 1843)
Figure5g–l
Material. 44 anterior teeth—SNSB-BSPG 2019 III-34,
SNSB-BSPG 2019 III-35 (4 teeth), SNSB-BSPG 2019 III-36
(39 teeth); 31 upper lateral teeth—SNSB-BSPG 2019 III-37,
SNSB-BSPG 2019 III-38 (5 teeth), SNSB-BSPG 2019 III-39
(25 teeth); and 20 lower lateral teeth—SNSB-BSPG 2019
III-40, SNSB-BSPG 2019 III-41 (5 teeth), SNSB-BSPG
2019 III-42 (14 teeth).
Description. The anterior teeth show an elongated and
slender cusp with a strong sigmoidal profile (Fig.5g, h).
The cutting edges are smooth and do not reach the base of
the crown. The crown/root boundary is strongly concave in
labial view. One to two pairs of sharp lateral cusplets are pre-
sent, which are lingually bent. The root is high with two long
Sharks, rays and skates from the Upper Marine Molasse (Burdigalian, Miocene) of Bavaria, Germany
1 3
and well-separated lobes. The lingual protuberance displays
a well-developed nutritive groove.
In the upper lateral teeth, the crown is triangular and
distally inclined (Fig.5i, j). In profile view, the crown
is straight. In lingual view, the enameloid surface shows
weakly vertical folds at its base. The crown/root boundary
is slightly concave in labial view.
The lower lateral teeth show a straight, rather low and
triangular cusp (Fig.5k, l). There are one to three pairs
of lateral cusplets, which are high and triangular in labial
view. The crown/root boundary is strongly concave. The
root is low with two short and well-separated lobes. The
lingual protuberance is bifurcated with a nutritive groove.
Remarks. The fossil record of C. acutissimus ranges from the
Oligocene to the Pliocene (Cappetta 2012). Early Miocene
records were reported from Austria (Schultz 2013), Costa
Rica (Laurito etal. 2014), France (Chevalier 1961; Cap-
petta 1970, as Odontaspis acutissima), Germany (Barthelt
etal. 1991, as Synodontaspis acutissima; Baier etal. 2004;
Höltke 2014), Hungary (Kordos and Solt 1984, as Odon-
taspis acutissima; Kocsis 2007), Italy (Marsili etal. 2007),
Slovakia (Holec etal. 1995, as Synodontaspis acutissima),
Switzerland (Leriche 1927; Fischli 1930) and the USA (Case
1980, as Odontaspis acutissima). Although teeth of C. tau-
rus share some similarities with those of C. acutissimus
(Arambourg 1952), no detailed studies about their morpho-
logical characters have been carried out so far. The teeth
described here are similar to those from the early Miocene
of Italy (Marsili etal. 2007) and the middle Miocene of
Hungary (Szabó and Kocsis 2016).
Genus Odontaspis Agassiz, 1843
Type species. Squalus ferox Risso, 1810
Odontaspis molassica (Probst, 1879)
Figure5m–p
Material. Two anterior teeth—SNSB-BSPG 2019 III-43,
SNSB-BSPG 2019 III-44; and two upper antero-lateral
teeth—SNSB-BSPG 2019 III-45, SNSB-BSPG 2019 III-46.
Description. The anterior teeth have a high and slender cusp
with a slightly sigmoidal profile (Fig.5m, n). The enam-
eloid surface is completely smooth on both cusp faces. The
two pairs of lateral cusplets are high and sharp. The cutting
edges are smooth and do not reach the base of the crown.
The lingual protuberance is very strong and it is divided
by a nutritive groove. The crown/root boundary is slightly
concave. The root is high and massive in lingual view. Both
lobes are well separated, but lack their extremities.
The upper antero-lateral tooth has a triangular cusp,
which is strongly distally oriented (Fig.5o, p). Three pairs of
lateral cusplets are present distally. The mesial cutting edge
is convex, whereas the distal one is concave. The crown/root
boundary is straight in both faces. The root is rather low and
straight. The mesial part of the crown and root is broken.
Remarks. Odontaspis molassica is not very common in the
fossil record. Early Miocene finds are from France (Cap-
petta 1970, 1973), Germany (Probst 1879; Barthelt etal.
1991; Baier etal. 2004, Sach 2016) and Portugal (Antunes
etal. 1981).
According to Reinecke etal. (2011), the anterior teeth of
O. molassica can be distinguished from Carcharias species
(e.g. C. gustrowensis) by their more slender and higher cusp.
Additionally, the crown surface in O. molassica is always
smooth on both cusps faces in lateral and anterior teeth.
Finally, the teeth described here have a slightly concave to
straight crown/root boundary in labial view. All these char-
acters were originally highlighted by Probst (1879) and later
confirmed by Bracher and Unger (2007).
Family Lamnidae Müller and Henle, 1838
Genus Carcharodon Müller and Henle, 1838
Type species. Squalus carcharias Linnaeus, 1758
Carcharodon hastalis (Agassiz, 1838)
Figure5q–t
Material. One upper lateral—SNSB-BSPG 2019 III-47; and
one lower lateral tooth—SNSB-BSPG 2019 III-48.
Description. The upper lateral tooth displays a triangular
cusp with smooth cutting edges (Fig.5q, r). The mesial cut-
ting edge is slightly convex, whereas the distal one is con-
cave. The enameloid surface is completely smooth and the
labial face is flat. The root is high with short lobes.
In the lower lateral tooth, the cusp is also triangular, but
straight and narrower than the upper tooth (Fig.5s, t). The
basal part of the mesial and distal edges is concave. The root
is high and flat with a straight basal part, probably eroded.
Remarks. The taxonomic classification of C. hastalis has
been widely debated in the last years (see Purdy etal. 2001;
Cappetta 2012; Cione etal. 2012; Ehret etal. 2012; Boesse-
necker etal. 2019). Ehret etal (2012) proposed a reconstruc-
tion of the evolutionary history of the genus Carcharodon
based on dental characters shared between the fossil and
extant species. According to these authors, the non-ser-
rated C. hastalis evolved into the semi-serrated C. hubbelli
and then to the fully serrated C.carcharias. Therefore, the
J. A. Villafaña etal.
1 3
evolutionary transition from C. hastalis to C. carcharias
occurred within a span of 6.9–5.3Ma. (Long etal. 2014;
Boessenecker etal. 2019).
The fossil record of C. hastalis ranges from the Miocene
to the Pliocene (Cappetta 2012). Early Miocene records
were reported from Argentina (Scasso and Castro 1999),
Austria (Schultz 2013), Chile (Suarez etal. 2006), Egypt
(Cook etal. 2010), France (Cappetta 1970), Germany (Bar-
thelt etal. 1991, as Isurus hastalis; Baier etal. 2004; Rei-
necke etal. 2011); Sach 2016), Hungary (Kordos and Solt
1984, as Isurus hastalis; Kocsis 2007), Italy (Marsili etal.
2007), Peru (Bianucci etal. 2018; Landini etal. 2019, as
Cosmopolitodus hastalis), Slovakia (Koch 1904, as Isu-
rus hastalis), Spain (Vicens and Rodríguez-Perea 2003, as
Isurus hastalis), Switzerland (Leriche 1927) and the USA
(Purdy 1998; Purdy etal. 2001; Kent 2018).
Order Carcharhiniformes Compagno, 1973
Family Carcharhinidae Jordan and Evermann, 1896
Genus Carcharhinus de Blainville, 1816
Type species. Carcharias melanopterus Quoy and Gaimard,
1824
Carcharhinus priscus (Agassiz, 1843)
Figure6a, d
Material. 25 upper antero-lateral teeth—SNSB-BSPG 2019
III-49, SNSB-BSPG 2019 III-50 (6 teeth), SNSB-BSPG
2019 III-51 (18 teeth); and 14 lower teeth—SNSB-BSPG
2019 III-52, SNSB-BSPG 2019 III-53 (13 teeth).
Description. The upper antero-lateral teeth have a broad and
triangular cusp, which is distally inclined (Fig.6a, b). The
cutting edges are continuously serrated along the main cusp
and on the heels. The root is high in lingual view with well–
separated root lobes. A shallow nutritive groove is present
in the lingual root protuberance.
In the lower teeth, the cusp is narrower and also distally
inclined with broad lateral heels (Fig.6c, d). The cutting
edges are faintly serrated in the main cusp and heels. In lin-
gual view, the root is mesio-distally extended with a distinct
furrow and foramen. The basal face of the root is straight
basally.
Remarks. Carcharhinus priscus is one the most common
species reported from the Neogene of the Mediterranean
Sea and Paratethys (Cappetta 2012; Szabó and Kocsis 2016),
but might represent a wastebasket taxon. This fossil species
range from the early Miocene to Pliocene according to Rei-
necke etal. (2011). Early Miocene records were reported
from Austria (Schultz 2013), Brazil (Toledo 1989), France
(Cappetta 1970, 1973), Germany (Barthelt etal. 1991;
Scholz and Bienert 1992; Baier etal. 2004; Reinecke etal.
2011; Pollerspöck and Beaury 2014; Sach 2016), Hungary
(Kocsis 2007), Pakistan (Welcomme etal. 1997), Portugal
(Antunes etal. 1981), Saudi Arabia (Thomas 1982), Slova-
kia (Holec etal. 1995), Spain (Vicens and Rodríguez-Perea
2003), Switzerland (Bolliger etal. 1995) and the USA (Case
1980; Kent 2018).
According to Maisch etal. (2018), the serrated cutting
edges and the absence of a notch separating the main cusp
and the tooth shoulders allow to separate C. priscus from
other species of Carcharhinus. The teeth reported herein
share similarities with those from the early Miocene of
Northern Germany (see Reinecke etal. 2011) and are there-
fore assigned to this species.
Carcharhinus sp.
Figure6e, f
Material. 14 upper antero-lateral teeth—SNSB-BSPG 2019
III-54, SNSB-BSPG 2019 III-55 (13 teeth); and three lower
antero-lateral teeth—SNSB-BSPG 2019 III-56.
Description. The upper antero-lateral teeth display a triangu-
lar and distally inclined cusp (Fig.6e, f). The cutting edges
are completely smooth, probably eroded. The heels are low
and slightly convex in labial view. The root is high in lingual
view with well-separated lobes. In the lower antero-lateral
teeth, the crown is rather low and upright. The cutting edges
are also smooth along the cusp and crown shoulders. The
root is low with well-separated lobes.
Remarks. The genus Carcharhinus is currently represented by
35 species with global distributions (Pollerspöck and Straube
2019). In the Mediterranean Sea, the sandbar shark (C.
plumbeus), the bignose shark (C. altimus), the copper shark
(C. brachyurus), the silky shark (C. falciformis), the spin-
ner shark (C. brevipinna), the blacktip shark (C. limbatus),
blacktip reef shark (C. melanopterus) and the dusky shark
(C. obscurus) have been reported up to now (Garibaldi and
Relini 2012; Froese and Pauly 2019). The fossil record of Car-
charhinus extends back to the middle Eocene (Kriwet 2005;
Cappetta 2012; Underwood and Gunter 2012). Early Miocene
records of Carcharhinus were widely reported from Europe
and America (Cappetta 1970, 2012; Case 1980; Barthelt etal.
1991; Suarez etal. 2006; Reinecke etal. 2011; Schultz 2013).
Genus Galeocerdo Müller and Henle, 1837
Type species. Squalus arcticus Faber, 1829
Galeocerdo aduncus Agassiz, 1843
Figure6g–j
Sharks, rays and skates from the Upper Marine Molasse (Burdigalian, Miocene) of Bavaria, Germany
1 3
Material. Eight antero-lateral teeth—SNSB-BSPG 2019 III-
57, SNSB-BSPG 2019 III-58, SNSB-BSPG 2019 III-59 (six
teeth); and one posterior tooth—SNSB-BSPG 2019 III-60.
Description. The antero-lateral teeth have a triangular and
broad cusp, which is strongly distally inclined (Fig.6g, h).
The mesial cutting edge is long and sigmoidal, whereas the
distal one is short and slightly convex. Both cutting edges
are strongly serrated from the base to the middle of the cusp,
being only faintly serrated in the apex. The distal heel is high
and strongly serrated. The root is very high in lingual view
and low in labial face.
Fig. 6 Carcharhiniformes. ad Carcharhinus priscus, a, b upper
antero-lateral tooth, c, d lower antero-lateral tooth, e, f Carcharhi-
nus sp., gj Galeocerdo aduncus, k, l Physogaleus contortus, m, n
Rhizoprionodon fischeri, o, p Isogomphodon acuaris, q, r Sphyrna
sp. Labial: a, c, e, g, i, k, m, o; lingual: b, d, f, h, j, l, n, p. Scale bar
2mm
J. A. Villafaña etal.
1 3
The posterior tooth is broader than high (Fig.6i, j). The
crown is low, triangular and strongly distally bent. The
mesial cutting edge is convex, whereas the distal one is
straight. Both cutting edges are serrated. The distal heel
is short, low and also serrated. The root is higher than the
crown in lingual view.
Remarks. Galeocerdo aduncus is common in Miocene
deposits (Cappetta 2012). Early Miocene records were
reported from Austria (Schultz 2013; Pollerspöck etal.
2018), France (Cappetta 1970), Germany (Probst 1879;
Lutzeier 1922; Barthelt etal. 1991; Scholz and Bienert 1992;
Baier etal. 2004; Reinecke etal. 2011; Sach 2016), Hungary
(Kordos and Solt 1984; Kocsis 2007), Italy (Marsili etal.
2007), Malta (Ward and Bonavia 2001), Portugal (Antunes
etal. 1981), Peru (Bianucci etal. 2018; Landini etal. 2019),
Slovakia (Holec etal. 1995), Spain (Vicens and Rodríguez-
Perea 2003; Mas 2009), Switzerland (Leriche 1927; Holec
etal. 1995) and the USA (Purdy 1998; Kent 2018).
Genus Physogaleus Cappetta, 1980
Type species. Trigonodus secundus Winkler, 1876
Physogaleus contortus (Gibbes, 1849)
Figure6k, l
Material. One antero-lateral tooth—SNSB-BSPG 2019
III-61.
Description. The tooth displays a slender and distally
inclined cusp (Fig.6k, l). The cutting edges are weakly ser-
rated along the cusp and distal heel. The distal heel is low
and convex. The root is high in lingual view and low in labial
view. The root/crown boundary is strongly convex in lingual
view, whereas it is slightly concave in labial view. Both root
lobes are short and their extremities are missing.
Remarks. According to Reinecke etal. (2011), the fossil
record of P. contortus ranges from the Oligocene to the
middle Miocene. Early Miocene records were reported from
Italy (Marsili etal. 2007), Germany (Reinecke etal. 2011),
Hungary (Kocsis 2007, as Galeocerdo contortus), Panama
(Pimiento etal. 2013), Peru (Bianucci etal. 2018; Landini
etal. 2019) and the USA (Case 1980, as Galeocerdo con-
tortus; Kent 2018).
Teeth of G. contortus have been confused often with
lower teeth of G. aduncus. We follow the opinion of Purdy
etal. (2001) and Reinecke etal. (2011), considering the
aduncus and contortus morphotypes as belonging to differ-
ent genera based on substantial differences between their
dental characters.
Genus Rhizoprionodon Whitley, 1929
Type species. Carcharias (Scoliodon) crenidens Klunzinger,
1880
Rhizoprionodon sp.
Figure6m, n
Material. Two antero-lateral teeth—SNSB-BSPG 2019 III-
62, SNSB-BSPG 2019 III-63.
Description. Both teeth are abraded and part of the distal
region is missing (Fig.6m, n). The cusp is triangular and
distally inclined with a flared base. The mesial cutting edge
is slightly concave, whereas the distal one is straight. The
distal heel is low and convex. The cutting edges are smooth
along the cusp and distal heel. In lingual view, the root is
high and shows a strong protuberance that is separated by a
transverse nutritive groove.
Remarks. The genus Rhizoprionodon is currently repre-
sented by seven species occurring in temperate and tropical
regions (Compagno 1984b). However, the genus is absent in
the Mediterranean Sea. The fossil record of Rhizoprionodon
extends back to the early Eocene (Cappetta 2012). Early
Miocene records were reported from Austria (Schultz 2013),
Germany (Barthelt etal. 1991; Baier etal. 2004; Reinecke
etal. 2011; Pollerspöck and Straube 2017), Malta (Ward and
Bonavia 2001), Switzerland (Bolliger etal. 1995) and the
USA (Case 1980; Kent 2018).
According to Reinecke etal. (2011), Rhizoprionodon spe-
cies show very similar dental characters and their identifi-
cation at specific level is difficult. For this reason, we only
identify our specimens at generic level.
Genus Isogomphodon Gill, 1862
Type species. Carcharias (Prionodon) oxyrhynchus Valen-
ciennes, 1839
Isogomphodon acuarius (Probst, 1879)
Figure6o, p
Material. One lower lateral tooth—SNSB-BSPG 2019
III-64.
Description. The cusp is rather high and slightly distally
inclined (Fig.6o, p). The mesial and distal heels are low
and mesio-distally extended. The cutting edges are smooth
along the cusp and heels. The enameloid surface is com-
pletely smooth in both faces. The root is high in lingual view
Sharks, rays and skates from the Upper Marine Molasse (Burdigalian, Miocene) of Bavaria, Germany
1 3
with a V-shaped basal face. The lobes are long with rounded
extremities.
Remarks. The fossil record of I. acuarius ranges from early
Miocene to the late Miocene (Cappetta 2012). Early Mio-
cene records were reported from Costa Rica (Laurito 1999),
France (Lalai, 1986), France (Cappetta 1970, as Aprionodon
acuarius), Germany (Barthelt etal. 1991; Baier etal. 2004;
Sach 2016), the USA (Case 1980), Switzerland (Bolliger
etal. 1995) and Venezuela (Carrillo-Briceño etal. 2016b).
The tooth described herein is similar to those from the early
Miocene of Venezuela (Carrillo-Briceño etal. 2016b) and
the middle Miocene of France (Vialle etal. 2011).
Family Sphyrnidae Gill, 1872
Genus Sphyrna Rafinesque, 1810
Type species. Squalus zygaena Linnaeus, 1758
Sphyrna sp.
Figure6q, r
Material. Seven lower lateral teeth—SNSB-BSPG 2019 III-
65, SNSB-BSPG 2019 III-66 (six teeth).
Description. The teeth have a low and distally inclined cusp
(Fig.6q, r). The mesial cutting edges are slightly concave
at their bases, but straight at their upper part. The distal
heel is rather high and strongly convex. The cutting edges
are smooth along the cusp and distal heel. The cusp and the
distal heel are separated by a notch. The root is high and
displays a vertical nutritive groove.
Remarks. The genus Sphyrna is currently represented by
nine species occurring in tropical and temperate seas (Com-
pagno 1984b). The scalloped hammerhead (S. lewini), the
great hammerhead (S. mokarran), the smalleye hammer-
head (S. tudes) and the smooth hammerhead (S. zygaena)
are known from the present-day Mediterranean Sea (Com-
pagno 1998). The genus Sphyrna extends back to the lower
Oligocene (Cappetta 2012). Early Miocene records are
from Austria (Schultz 1998), Colombia (Carrillo-Briceño
etal. 2016a), France (Cappetta 1970), Germany (Barthelt
etal. 1991; Reinecke etal. 2011), Hungary (Kordos and
Solt 1984; Kocsis 2007), Malta (Ward and Bonavia 2001),
Panama (Gillette 1984), Peru (Bianucci etal. 2018; Landini
etal. 2019), Portugal (Antunes etal. 1981), Switzerland
(Leriche, 1927), Venezuela (Carrillo-Briceño etal. 2016b)
and the USA (Purdy 1998).
Reinecke etal. (2011) described S. laevissima and S. inte-
gra from the early Miocene of northern Germany. In the
described lower antero-lateral teeth of S. integra, the crown
is broader and the distal heel is straight or faintly convex. In
S. laevissima, the crown is triangular and the distal heel is
oblique. Therefore, these species show dental characters that
differ from those of the specimens described here. However,
as the few teeth reported here are very abraded and only rep-
resent lower teeth, we prefer to identify them only at generic
level until more material is available.
Family Scyliorhinidae Gill, 1862
Genus Scyliorhinus de Blainville, 1816
Type species. Squalus canicula Linnaeus, 1758
Scyliorhinus fossilis (Leriche, 1927)
Figure7a–d
Material. Twelve anterior teeth—SNSB-BSPG 2019 III-67,
SNSB-BSPG 2019 III-68 (11 teeth); and 12 lateral teeth—
SNSB-BSPG 2019 III-69, SNSB-BSPG 2019 III-70 (8
teeth), SNSB-BSPG 2019 III-71 (3 teeth).
Description. The anterior teeth have a straight and slender
cusp with a broader base (Fig.7a, b). There is a pair of
small, low and slightly divergent lateral cusplets, which are
separated from the main cusp by a notch. A second, very
incipient pair of lateral cusplets are developed at the outer-
most margins of the crown. Labially, strong vertical folds
are present along the mesial and distal margins of the crown,
extending from the crown/root boundary towards the base of
the first pair of lateral cusplets. Lingually, short and closely
spaced ridges are present below the lateral cusplets reach-
ing up to the middle of the first pair of lateral cusplets and
to the apex of the second pair. The cutting edges are well
developed and continuous between the lateral cusplets and
main cusp. The root is abraded and only the well-developed
lingual protuberance is partially preserved.
In lateral teeth, the cusp is low and strongly distally
inclined (Fig.7c, d). In labial view, less pronounced folds
are present along the distal and mesial margins. The root
is well preserved, showing separated lobes with rounded
extremities in labial view. A deep vertical nutritive groove
is present on the lingual face of the root.
Remarks. Teeth of S. fossilis were reported from Miocene
deposits of the Paratethys and Mediterranean regions. This
species also occurs in the upper Marine Molasse of the
Molasse Basin (Barthelt etal. 1991; Pfeil 1991; Reinecke
etal. 2011; Schultz 2013). Additionally, it was described
from the Miocene (Aquitanian to Messinian) of Switzerland,
southern France, and Portugal (Leriche 1927; Antunes and
Jonet 1970; Cappetta 1970, 2006; Jost etal. 2016).
J. A. Villafaña etal.
1 3
The species S. joleaudi described by Cappetta (1970)
from the Miocene of southern France represents a junior
synonym of S. fossilis. Reinecke etal. (2011) reported
the presence of S. fossilis (aka S. joleaudi) from the early
Miocene of northern Germany. The teeth described here dis-
play a typical morphotype of S. fossilis with mesio-distally
expanded crown base and several, very strong vertical folds
and lateral cusplets.
Fig. 7 Carcharhiniformes. ad Scyliorhinus fossilis, a, b anterior
tooth, c, d lateral tooth, e, f Scyliorhinus sp., gj Pachyscyllium dis-
tans, g, h anterior tooth, i, j lateral tooth, k, l Chaenogaleus affinis,
upper lateral tooth, mp Paragaleus pulchellus. Labial: a, c, e, g, i, k,
m, p; lingual: b, d, f, h, j, l, n, o. Scale bar 1mm
Sharks, rays and skates from the Upper Marine Molasse (Burdigalian, Miocene) of Bavaria, Germany
1 3
Scyliorhinus sp.
Figure7e, f
Material. Seven lateral teeth—SNSB-BSPG 2019 III-72,
SNSB-BSPG 2019 III-73 (six teeth).
Description. The teeth of this catshark are abraded and
incompletely preserved (Fig.7e, f). The main cusp is tri-
angular in labial view, slightly bent distally, and lingually
curved. Basally, short vertical folds are present on the lin-
gual crown face, whereas the lingual face is smooth. The
preserved cusplets are high, triangular and straight with-
out any ornamentation. The cutting edges are completely
smooth. The cusp is separated from the lateral cusplets by
a deep notch. The root is heavily abraded; thus, the lingual
protuberance is missing.
Remarks. Scyliorhinus is a diverse genus comprising 16
extant species (Froese and Pauly 2019). They are globally
distributed in tropical to arctic waters, from the intertidal
to the deep–sea zones (Compagno 1984b). In the Medi-
terranean Sea the lesser spotted dogfish (S. canicula), the
Dumahel’s catshark (S. duhamelii) and the nursehound (S.
stellaris) have been reported (Soares etal. 2019). The fossil
record of Scyliorhinus extends back to the lower Cretaceous
(Cappetta 2012). Early Miocene records were reported from
Austria (Schultz 2013), Germany (Barthelt etal. 1991; Rei-
necke etal. 2011), Switzerland (Leriche 1927) and the USA
(Case 1980).
The material described here can be unambiguously identi-
fied as Scyliorhinus based on the typical characters of this
genus (i.e. sharp and rather slender cusp with one pair of
lateral cusplets). However, due to the very abraded condi-
tion of the teeth and their incompleteness it is not possible
to assign these teeth to any species known to date.
Genus Premontreia Cappetta, 1992
Type species. Premontreia degremonti Cappetta, 1992
Premontreia distans (Probst, 1879)
Figure7g–j
Material. Five anterior teeth—SNSB-BSPG 2019 III-74,
SNSB-BSPG 2019 III-75 (4 teeth); and 14 lateral teeth—
SNSB-BSPG 2019 III-76, SNSB-BSPG 2019 III-77 (13
teeth).
Description. In the anterior teeth (Fig.7g, h), the main cusp
is triangular and rather low in lingual view with one pair of
lateral cusplets. The main cusp and the lateral cusplets are
lingually curved. The lateral cusplets are broad and triangu-
lar. In labial view, vertical folds are present at the base of
the crown. The lingual cusp face is smooth, but faint folds
are developed on the lateral cusplets. The root is high in lin-
gual view with a lingual protuberance and shallow nutritive
groove. The root lobes below the lateral cusplets seem to
have been very narrow as far as can be ascertained.
The lateral teeth (Fig.7i, j) display an incomplete trian-
gular, broad and distally inclined cusp. The lateral cusplets
also are triangular and very broad. Some folds are present at
the base of the lateral cusplets on both faces. The labial face
of the crown overhangs the root. The crown/root boundary
is concave medially and rounded at the distal and mesial
regions. The root is slightly broader than the crown and is
heart–shaped in basal view.
Remarks. Premontreia distans is very common in the Oli-
gocene and Miocene of the North Sea basin and adjacent
regions (Antunes etal. 1981; Lienau 1987; Haye etal. 2008;
Reinecke etal. 2008). This taxon was originally allocated to
Scyliorhinus (Joleaud 1912). However, Cappetta (2006) and
Reinecke etal. (2008) placed this species into the extinct
scyliorhinid taxon Premontreia. Early Miocene records were
reported from France (Cappetta 1970, 1973, as Scyliorhinus
distans), Germany (Probst 1879, as Scyllium distans; von
Ihering 1927; Barthelt etal. 1991, as Scyliorhinus distans;
Sachs 2016), Portugal (Antunes etal. 1981, as Scyliorhinus
distans), Switzerland (Bolliger etal. 1995, as Scyliorhinus
distans; Jost etal. 2016) and the USA (Case 1980). We fol-
lowed the opinion of Reinecke etal (2011), considering P.
distans as valid species based on its diagnostic characters
(i.e. labial ridges at the base of the crown and the convex
curvature of the mesial cutting edge). Therefore, the material
described here can be unambiguously identified at species
level based on the presence of those characters.
Family Hemigaleidae Hasse, 1879
Genus Chaenogaleus Gill, 1862
Type species. Chaenogaleus macrostoma (Bleeker, 1852)
Chaenogaleus affinis (Probst, 1879)
Figure7k, l
Material. 13 upper antero-lateral teeth—SNSB-BSPG 2019
III-78, SNSB-BSPG 2019 III-79 (2 teeth), SNSB-BSPG
2019 III-80 (10 teeth).
Description. The teeth show a high, broad and distally
inclined cusp (Fig.7k, l). The mesial cutting edge is convex
or slightly sigmoidal in some teeth, whereas the distal cut-
ting edges are straight or convex. The distal heel shows two
to four serrations decreasing in size towards the rear. The
enameloid surface displays weak folds at the base of the
J. A. Villafaña etal.
1 3
lingual and labial faces. The root is low and slightly wider
than the crown. It shows a well-marked lingual protuberance,
which is divided by a nutritive groove.
Remarks. The fossil species C. affinis ranges from the early
Miocene to late Miocene (Cappetta 2012). Early Miocene
records were reported from Austria (Schultz 2013), France
(Cappetta 1970, as Galeorhinus affinis), Germany (Probst
1878, as Galeus affinis; von Ihering 1927; Barthelt etal.
1991; Reinecke etal. 2011; Pollerspöck and Beaury 2014;
Sach 2016), Switzerland (Fischli 1930, as Galeus affinis;
Bolliger etal. 1995; Jost etal. 2016) and the USA (Case
1980, Galeorhinus affinis). According to Herman etal.
(2001), the teeth of the only extant species C. macrostoma
have more elongated and slender cusps in upper antero-
lateral than the fossil representative C. affinis. The teeth
reported here also bear dental characters observed in the
material from early Miocene of Northern Germany and
middle Miocene of Czech Republic (Schultz etal. 2010;
Reinecke etal. 2011).
Genus Paragaleus Budker, 1935
Type species. Paragaleus gruveli Budker, 1935
Paragaleus pulchellus (Jonet, 1966)
Figure7m–p
Material. One lower anterior—SNSB-BSPG 2019 III-81;
and 30 lower lateral teeth—SNSB-BSPG 2019 III-82,
SNSB-BSPG 2019 III-83 (19 teeth).
Description. The lower anterior tooth has a slender and erect
cusp (Fig.7m, n). The mesial cutting edge is concave at its
base and straight at its upper part. The distal heel is short
with two sharp cusplets, which are distally inclined. The
enameloid surface is completely smooth. The root is incom-
plete; however, the lingual protuberance is well preserved.
The lower lateral tooth has a long, slender and strongly
distally inclined cusp (Fig.7o, p). The mesial cutting edge is
concave, whereas the distal one is slightly convex. The distal
heel is short and features three acute and distally oriented
cusplets. The root is incomplete mesially and distally. The
lingual protuberance is abraded, but nonetheless preserves
a medial foramen.
Remarks. The fossil record of P. pulchellus ranges from the
early to late Miocene (Cappetta 2012). Reinecke etal. (2011)
also indicated the possible presence of P. pulchellus in the
early Miocene of northern Germany. The taxonomic assign-
ment of P. pulchellus has been debated for many years. Jonet
(1966) erected Galeorhinus pulchellus based on the teeth
from the late Miocene of Portugal. Later, Cappetta (1970)
included this species within the genus Paragaleus. Addition-
ally, Barthelt etal. (1991) considered Galeorhinus tenuis as
synonym of P. pulchellus, a view we follow in the present
study. Early Miocene records of P. pulchellus were reported
from Austria (Schultz 2013), France (Cappetta 1970), Ger-
many (Reinecke etal. 2011), Portugal (Antunes etal. 1981)
and Venezuela (Aguilera and de Aguilera 2004).
Based on the high similarities between our teeth and
those from the early Miocene of northern Germany, we
identify the teeth described here as belonging to the species
P. pulchellus.
Superorder Batomorphii Cappetta, 1980
Order Myliobatiformes Compagno, 1973
Family Aetobatidae Agassiz, 1858
Genus Aetobatus de Blainville, 1816
Type species. Raja aquila Linnaeus, 1758
Aetobatus sp.
Figure8a, b
Material. Six lower symphysial teeth—SNSB-BSPG 2019
III-84, SNSB-BSPG 2019 III-85, SNSB-BSPG 2019 III-86
(four teeth).
Description. The symphyseal teeth are transversely elon-
gated and V-shaped in occlusal view (Fig.8a, b). The lateral
edges of the symphyseal teeth form an obtuse angle. The
teeth are labio-lingually thicker in the central region than
in the lateral region. The crown surface is very abraded in
all the examined teeth. The root vascularization is of the
polyaulacorhizous type, with laminae and shallow nutritive
grooves.
Remarks. Aetobatus is currently represented by five species:
A. flagellum, A. laticeps, A. narinari, A. narutobiei and A.
oceallatus, living in tropical and warm-temperate seas (Last
etal. 2016). This genus is absent in the Mediterranean Sea.
Its fossil record extends back to the upper Paleocene (Cap-
petta 2012). Early Miocene records were reported from Aus-
tria (Schultz 2013), Egypt (Cook etal. 2010), France (Cap-
petta 1970, 1973; Goedert etal. 2017), Germany (Barthelt
etal. 1991; Baier etal. 2004; Reinecke etal. 2011; Sach
2016), Panamá (Gillette 1984), Portugal (Zbyszewski 1949),
Slovakia (Holect etal. 1995), Switzerland (Leriche 1927;
Bolliger etal. 1995) and the USA (Purdy 1998).
Our material displays the typical shape of Aetobatus
teeth, i.e. V-shaped symphyseal teeth without any lat-
eral teeth. Although Reinecke etal. (2011) described A.
arcuatus from the early Miocene of northern Germany,
its diagnostic characters are not very clear. According
Sharks, rays and skates from the Upper Marine Molasse (Burdigalian, Miocene) of Bavaria, Germany
1 3
Fig. 8 Myliobatiformes a, b Aetobatus sp., c, d Myliobatis sp., e, f
Rhinoptera sp., gj Taeniurops cavernosus, k, l Dasyatis probsti, m,
n Dasyatis rugosa, o, p Dasyatis strangulata, q, r Dasyatis sp., s, t
Myliobatiformes indet. Basal: b; dorsal: s; lingual: c, h, j, l, n; oclus-
sal: a, d, e, p, r; profile: f, g, i, k, m, o, q; dorsal: s; ventral: t. Scale
bar 1mm
J. A. Villafaña etal.
1 3
to Hovestadt and Hovestadt-Euler (2013), identification
at the species level of isolated teeth only is not possible
due the high intraspecific variability of dental characters.
Therefore, we abstain from assigning these teeth to any
species.
Family Myliobatidae Bonaparte, 1838
Genus Myliobatis Cuvier, 1817
Type species. Raja aquila Linnaeus, 1758
Myliobatis sp.
Figure8c, d
Material. Ten symphyseal teeth—SNSB-BSPG 2019 III-87,
SNSB-BSPG 2019 III-88, SNSB-BSPG 2019 III-89 (eight
teeth).
Description. The symphyseal teeth are very abraded and
some of them are broken (Fig.8c, d). The crown is trans-
versely elongated with straight labial and lingual margins
in occlusal view. The teeth are four to five times wider than
long. All specimens have a hexagonal outline. The occlusal
surface of the symphyseal teeth is smooth. The root is
abraded, but it displays the typical polyaulacorhizous vas-
cularization type.
Remarks. The genus Myliobatis is currently represented by
11 globally distributed species occurring in temperate and
tropical seas (Last etal. 2016). The common eagle ray (M.
aquila) is the only species recorded from the Mediterranean
Sea (McEachran and Séret 1990). Reliable fossils of Myliob-
atis extend back to the Late Cretaceous (Claeson etal. 2010;
Cappetta 2012). Early Miocene records were reported from
Austria (Schultz 2013), Chile (Suarez etal. 2006), France
(Cappetta 1970), Germany (Barthelt etal. 1991; Scholz and
Bienert 1992; Baier etal. 2004; Reinecke etal. 2011; Poller-
spöck and Beaury 2014; Sach 2016), Hungary (Kordos and
Solt 1984), Panamá (Gillette 1984), Portugal (Antunes etal.
1981), Spain (Vicens and Rodríguez-Perea 2003), Switzer-
land (Leriche 1927; Bolliger etal. 1995), the the USA (Case
1980) and Venezuela (Aguilera and de Aguilera 2004). As
in Aetobatus, taxonomic identification of extinct Myliobatis
species only based on isolated teeth is extremely difficult
due the high dental variation within the genus (Hovestadt
and Hovestadt-Euler 2013).
Family Rhinopteridae Jordan and Evermann, 1896
Genus Rhinoptera Cuvier, 1829
Type species. Myliobatis marginata Geoffroy Saint Hilaire,
1817
Rhinoptera sp.
Figure8e, f
Material. Four symphyseal teeth—SNSB-BSPG 2019 III-
90, SNSB-BSPG 2019 III-91 (three teeth); and three lateral
teeth—SNSB-BSPG 2019 III-92, SNSB-BSPG 2019 III-93
(two teeth).
Description. The symphyseal teeth are broader than long
with a hexagonal contour (Fig.8e, f). In occlusal view, the
teeth are straight or slightly arched lingually. The crown
surface is smooth. The root shows a polyaulacorhizid vas-
cularization type with numerous parallel laminae and nutri-
tive grooves. The most complete specimen has 12 laminae
in basal view. In profile view, the root is slightly displaced
lingually and lingually separated from the crown by a bulge.
The lateral tooth has a regular hexagonal outline, being less
transversely enlarged than the symphyseal teeth. The crown
surface is completely smooth.
Remarks. The genus Rhinoptera is currently known by
eight species distributed in temperate and tropical oceans
(Last etal. 2016). The Lusitanian crownose ray (R. margi-
nata) is the only species reported from the Mediterranean
Sea (McEachran and Séret 1990). The fossil record of Rhi-
noptera extends back into the upper Palaeocene (Cappetta
2012). Early Miocene records are from Austria (Schultz
2013), Brazil (Távora etal. 2010), France (Cappetta 1970;
Goedert etal. 2017), Germany (Lutzeier 1922; Barthelt
etal. 1991; Baier etal. 2004; Sach 2016), India (Mondal
etal. 2009), Panamá (Gillette 1984), Portugal (Zbyszewski
1949; Antunes etal. 1981), Switzerland (Leriche 1927; Fis-
chli 1930; Bolliger etal. 1995), the USA (Case 1980) and
Venezuela (Aguilera and de Aguilera 2004).
According to Herman etal. (2000) and Cappetta (2012),
the symphyseal teeth of Rhinoptera are longer, but less broad
transversely than Aetobatus and Myliobatis. Considering
these dental characters, we can assign these teeth unambigu-
ously to the genus Rhinoptera. However, due to the lack of
diagnostic characters, identification at specific level remains
difficult.
Family Dasyatidae Jordan, 1888
Genus Taeniurops Garman, 1913
Type species. Taeniura meyeni Müller and Henle, 1841
Sharks, rays and skates from the Upper Marine Molasse (Burdigalian, Miocene) of Bavaria, Germany
1 3
Taeniurops cavernosus (Probst, 1877)
Figure8g–j
Material. 23 female teeth—SNSB-BSPG 2019 III-94,
SNSB-BSPG 2019 III-95 (22 teeth); and ten male teeth—
SNSB-BSPG 2019 III-96, SNSB-BSPG 2019 III-97 (8
teeth), SNSB-BSPG 2019 III-98.
Description. The female teeth have a rather high and acute
crown, which is lingually inclined in profile view (Fig.8g,
h). The crown shows a labial and lingual visor divided by a
sharp transverse crest. This crest displays some folds along
the distal and mesial edges. The lower region of the labial
visor is slightly convex and exhibits a reticulate ornamenta-
tion whereas the upper region is concave and smooth. The
lingual visor is concave in profile view with a smooth sur-
face. The root is high with two separated lobes.
Male teeth display a strong cuspidate and lingually ori-
ented cusp (Fig.8i, j). The labial visor is long with a slightly
convex and ornamented lower region whereas the upper
region is depressed and smooth. The transversal crest also
is folded in its distal and mesial edges. The root is rather low
and directed lingually with two short lobes.
Remarks. The fossil record of T. cavernosus ranges from the
lower to the middle Miocene (Reinecke etal. 2011; Cappetta
2012). Early Miocene records were only reported from Ger-
many (Probst 1877, as Raja cavernosa; Barthelt etal. 1991,
as Dasyatis cavernosa; Reinecke etal. 2011; Sach 2016),
Portugal (Antunes etal. 1981, as Dasyatis cavernosa), Swit-
zerland (Fischli 1930, as Trygon cavernosus; Bolliger etal.
1995) and the USA (Case 1980). According to Cappetta
(2013), the genus Taeniurops has been confused very often
with Dasyatis. However, Taeniurops shows a distinctively
depression in the labial visor which is borderer by a sharp
crest, thus differing from the condition seen in Dasyatis.
Male and female teeth reported here resemble the material
described from the early Miocene of northern Germany
(Reinecke etal. 2011).
Genus Dasyatis Rafinesque, 1810
Type species. Dasyatis ujo Rafinesque, 1810
Dasyatis probsti Cappetta, 1970
Figure8k, l
Material. Twelve male teeth—SNSB-BSPG 2019 III-99,
SNSB-BSPG 2019 III-100 (11 teeth).
Description. The teeth have a cuspidate crown, which is
lingually inclined (Fig.8k, l). The labial visor is smooth
and slightly convex; however, its medial region is deeply
depressed. The lingual visor is also smooth and concave
in profile view. The transversal crest is faintly folded. The
labial margin is convex and weakly ornamented. The root is
rather high and lingually oriented with two well-separated
lobes.
Remarks. Dasyatis probsti ranges from the early to the mid-
dle Miocene (Reinecke etal. 2011). Early Miocene records
were reported from France (Cappetta 1970, 1973), Germany
(Reinecke etal. 2011; Pollerspöck and Beaury 2014) and
Switzerland (Bolliger etal. 1995). Dasyatis probsti can be
distinguished from T. cavernosus and D. rugosa by its deep
depression on the labial visor and the weakly ornamented
labial margin of the crown.
Dasyatis rugosa Probst, 1877
Figure8m, n
Material. 33 female teeth—SNSB-BSPG 2019 III-101,
SNSB-BSPG 2019 III-102 (29 teeth), SNSB-BSPG 2019
III-103 (3 teeth).
Description. The teeth show a rather low and lingually
oriented crown (Fig.8m, n). The labial visor is convex in
profile view and strongly ornamented. The lingual visor is
concave in profile view with a smooth surface. In occlusal
view, the labial visor is angular. The root is rather high and
directed lingually with two massive lobes.
Remarks. The fossil record of D. rugosa ranges from the
Oligocene to the middle Miocene (Reinecke etal. 2011).
Early Miocene records were reported from Austria (Schultz
1998), France (Cappetta 1970, 1973), Germany (Probst
1877, as Raja rugosa; Barthelt etal. 1991; Reinecke etal.
2011; Pollerspöck and Beaury 2014; Sach 2016), Portugal
(Antunes etal. 1981), Switzerland (Bolliger etal. 1995; Jost
etal. 2016) and the USA (Kent 2018).
The material described here shows the diagnostic char-
acters of teeth of D. rugosa, i.e. a strongly ornamented
labial visor and a labial visor that appears as angular in
occlusal view. These dental characters were also observed
in teeth from the early Miocene of northern Germany (Rei-
necke etal. 2011).
Dasyatis strangulata (Probst, 1877)
Figure8o, p
Material. Two female teeth—SNSB-BSPG 2019 III-104,
SNSB-BSPG 2019 III-105.
Description. The female teeth show a bulging and lingually
directed crown (Fig.8o, p). The transverse ridge is roughly
J. A. Villafaña etal.
1 3
pronounced, separating the labial and lingual visors. The
labial visor is almost flat with a weakly reticulated surface,
whereas the lingual visor is short, smooth and slightly con-
vex in profile view. The labial margin of the crown is very
thick and convex. The root is very low with two short lobes.
Remarks. Teeth of D. strangulata are very rare in the fossil
record. This species ranges from the early Miocene to the
Pliocene. Early Miocene records were only reported from
Germany (Probst 1877, as Raja strangulata; Reinecke etal.
2011). D. strangulata can be distinguished from other spe-
cies of Dasyatis and Taeniurops by the bulging crown shape
and the absence of a labial depression.
Dasyatis sp.
Figure8q, r
Material. One female tooth—SNSB-BSPG 2019 III-106.
Description. The single tooth displays a rather bulging
and lingually oriented cusp (Fig.8q, r). The labial visor is
weakly reticulated, whereas the lingual visor is completely
smooth. In profile view, the labial visor is strongly convex
and the lingual visor is concave in its medial region. In
occlusal view, the crown displays a semicircular outline. The
root is low with two short well-separated lobes.
Remarks. According to Last etal. (2016), Dasyatis is cur-
rently represented by five species with a global distribution.
Of these, three (i.e. the marbled stingray D. marmorata, the
common stingray D. pastinaca and the Tortonese’s stingray
D. tortonesi) are currently present in the Mediterranean Sea
(Cowley and Compagno 1993). The fossil record of Dasyatis
extends back into the early Cretaceous, although most of the
species might belong to different genera (Underwood etal
1999; Cappetta 2012). Early Miocene records were reported
from Austria (Schultz 2013), France (Cappetta 1970), Ger-
many (Barthelt etal. 1991; Reinecke etal. 2011), the USA
(Purdy 1998) and Venezuela (Aguilera and de Aguilera
2004).
Myliobatiformes indet.
Figure8s, t
Material. Six incomplete tail spines—SNSB-BSPG 2019
III-107, SNSB-BSPG 2019 III-108 (three incomplete tail
spines), SNSB-BSPG 2019 III-109 (two incomplete tail
spines).
Description. The six tail spines are incomplete, missing their
distal and proximal portions (Fig.8s, t). They are dorso-
ventrally flattened, narrow and elongated. In dorsal view,
an antero-posteriorly directed central groove and additional
irregularly shaped grooves are observed. In ventral view,
a central ridge is present, but weakly pronounced. Both
sides of the tail spines display flat denticles that project
latero-proximally.
Remarks. Living stingrays (Myliobatiformes) are repre-
sented by ten families (Last etal. 2016). Early Miocene
records of tail spines were reported from northern Germany
(Reinecke etal. 2011), Venezuela (Carrillo-Briceño etal.
2016b) and Peru (Bianucci etal. 2018; Landini etal. 2019).
According to the recent review by Hovestadt and
Hovestadt-Euler (2013), in general, there are no unambigu-
ous characters that might be useful to distinguish tail spines
at genus or family level. Moreover, their morphology could
also vary ontogenetically. Therefore, we prefer to keep their
identification to a higher taxonomic level.
Order Rajiformes Berg, 1937
Family Rajidae de Blainville, 1816
Genus Raja Linnaeus, 1758
Type species. Raja miraletus Linnaeus, 1758
Raja sp.
Figure9a, d
Material. 17 female teeth—SNSB-BSPG 2019 III-110,
SNSB-BSPG 2019 III-111 (10 teeth), SNSB-BSPG 2019
III-112 (six teeth); and 15 male teeth—SNSB-BSPG 2019
III-113, SNSB-BSPG 2019 III-114 (13 teeth), SNSB-BSPG
2019 III-115.
Description. Most of the teeth are very abraded and in
some of them the apex is missing. The male teeth display a
very high and cuspidate crown, which is lingually oriented
(Fig.9a, b). The enameloid surface is completely smooth
on both the labial and lingual faces. The cutting edges are
smooth along the mesial and distal edges of the cusp, and
they do not reach the base of the cusp. The base of the crown
displays a rounded rim with an oval shape in occlusal view.
The root is low and mesio-distally expanded with two short
but wide lobes. The median furrow is narrow and shallow.
The female teeth display a rounded and low crown, which
is lingually oriented (Fig.9c, d). The cutting edges are mostly
smooth. They do not reach the basal rim of the crown. Some
teeth display some ridges on the transverse crest in the distal
and mesial edges. In profile view, the labial crown face is
strongly convex whereas the lingual face is slightly convex.
The root is low with two wide and short lobes.
Remarks. The genus Raja is currently represented by
16 globally distributed species (Last etal. 2016). In the
Sharks, rays and skates from the Upper Marine Molasse (Burdigalian, Miocene) of Bavaria, Germany
1 3
Mediterranean Sea, nine species have been reported (e.g. R.
asterias and R. clavata) up to now (Serena 2005). The fos-
sil record of Raja extends back into the upper Cretaceous,
although most of the geologically oldest species probably
do not belong to the living genus (Cappetta 2012). Early
Miocene records were reported from Austria (Schultz 2013),
France (Cappetta 1970, 1973), Germany (Barthelt etal.
1991; Reinecke etal. 2011; Pollerspöck and Beaury 2014),
Hungary (Kordos and Solt 1984), India (Sahni and Mehrotra
1981), Portugal (Antunes etal. 1981), Switzerland (Fischli
1930; Bolliger etal. 1995) and the USA (Purdy 1998; Kent
2018). Reinecke etal. (2011) described the species Raja
cecilae and Raja holsatica from the early Miocene of north-
ern Germany. However, their diagnostic characters were not
described in detail. Although our material shows the general
morphology of Raja, these dental characters are different
from those described from Germany. Due to the poor pres-
ervation state, we prefer to identify them at the genus-level.
Order Rhinopristiformes Naylor etal., 2012
Family Rhinidae Müller and Henle, 1841
Genus Rhynchobatus Müller and Henle, 1837
Type species. Rhinobatus laevis (Bloch and Schneider, 1801)
Rhynchobatus sp.
Figure9e, f
Material. Five teeth—SNSB-BSPG 2019 III-116, SNSB-
BSPG 2019 III-117 (four teeth).
Description. The teeth have a globular crown, which is
wider than long (Fig.9e, f). The crown is divided into three
regions: labial, central and lingual faces. In profile view,
the labial crown face is strongly convex. The central crown
face is slightly depressed and weakly separated from the
labial face by a transverse crest. The lingual face is oblique
and slightly depressed. The crown surface is ornamented
by granules around the labial and lingual faces not reach-
ing the basal margins. The lingual uvula is wide and quite
short. The root is very short, oriented lingually and divided
by two lobes.
Remarks. The genus Rhynchobatus is currently represented
by eight species that inhabit the Indian, western Pacific
and eastern Atlantic oceans, being in turn absent in the
Mediterranean Sea (Last etal. 2016). The fossil record of
Fig. 9 Rajiformes. ad Raja sp., a, b male tooth, c, d female tooth, Rhinopristiformes, e, f Rhynchobatus sp. Lingual: a, c, e; profile: b, d, f.
Scale bar 0.5mm
J. A. Villafaña etal.
1 3
Rhynchobatus extends back into the lower Eocene (Cappetta
2012). Early Miocene records were reported from France
(Cappetta 1970, 1973), Germany (Barthelt etal. 1991; Rei-
necke etal. 2011; Sach 2016), Malta (Ward and Bonavia
2001), Portugal (Antunes etal. 1981), Switzerland (Bolliger
etal. 1995), the USA (Case 1980; Kent 2018) and Venezuela
(Aguilera and de Aguilera 2004).
The teeth described herein display the typical characters
of the genus Rhynchobatus, i.e. the oral face is divided into
three regions, the enameloid is granular and the uvula is
wide. A number of species of Rhynchobatus and R. pris-
tinus have been reported from the early Miocene of Ger-
many (Barthelt etal. 1991; Schultz 2013). However, the
diagnostic characters of this species are not clear, thus,
we prefer to identify it at generic level until more material
is available.
Discussion
Taxonomic composition ofSimssee fauna
Elasmobranch remains are quite common in the Marine
Molasse Basin of southern Germany and highlight a diver-
sified cartilaginous fish fauna (Barthelt etal. 1991; Poller-
spöck and Straube 2017; this study). Sharks, rays and
skates are well represented in the fossiliferous deposits of
the Achen Formation in the Simssee area. However, most
of the remains are too incomplete or abraded to allow an
unambiguous identification at the species level (e.g. Mylio-
batis and Scyliorhinus). The elasmobranch fauna from the
early Miocene of Simssee/Bavaria is represented by 37 taxa
(Figs.2, 3, 4, 5, 6, 7, 8, and 9), including 26 sharks (70%,
26 out of 37 species) and 11 batoids (30%, 11 out of 37)
(Table1). The asymptotic trend of the rarefaction curve
(Fig.10) suggests that the taxonomic inventory is largely
complete. In addition, the Chao 1 non-parametric estimator
suggests that the completeness of the inventory would be no
less than 89% (Fig.10): indeed, the upper level confidence
interval (95%) of the Chao 1 extrapolation index suggests
that the total inventory would be ca. 41 taxa, i.e. 4 taxa more
than those that have been observed. The sharks are mainly
represented by members of the orders Carcharhiniformes
(46%, 12 out of 26 shark species) and Lamniformes (23%,
6 of 26), whereas for batoids, the order Myliobatiformes
is the most dominant group (82%, 9 of 11). At the fam-
ily level, the carcharhinids and the dasyatids are the most
diverse groups of sharks and batoids, respectively. At lower
taxonomic levels, 31 genera and 20 species of early Miocene
elasmobranchs were described. All the taxa described herein
were previously reported from other early Miocene localities
of Germany (e.g. Barthelt etal. 1991; Reinecke etal. 2011;
Pollerspöck and Straube 2017). Significantly, we provided
the first records of the shark species Paragaleus pulchel-
lus and Physogaleus contortus from southern Germany.
Additionally, we confirmed the presence of the rare species
Dasyatis strangulata from Germany.
Ecological traits oftheidentied taxa
Identifiable cartilaginous fishes from the Simssee area
comprise taxa that are common elements in marine sedi-
ments of Miocene age throughout Europe (Cappetta 1970,
1973; Antunes etal. 1981; Kocsis 2007; Marsili etal. 2007;
Reinecke etal. 2011; Schultz 2013). All the reported taxa
are nektonic or nektobenthic organisms and some of them
are able to migrate over long distances (e.g. Alopias, Car-
charhinus and Squalus) (McFarlane and King 2003; Car-
tamil etal. 2010; Conrath and Musick 2010). Most of the
elasmobranchs reported here are inhabitants of shallow,
nearshore and littoral marine waters in warm climatic zones,
according to our current knowledge about their fossil dis-
tribution, and in comparison, with modern representatives
(e.g. Aetobatus and Chaenogaleus) (Last etal. 2016; Froese
and Pauly 2019). This is consistent with the current distri-
bution of cartilaginous species in the Mediterranean Sea,
which are mostly distributed on the continental shelf (Fro-
ese and Pauly 2019; Ramírez-Amaro etal. 2020). However,
purported deep-water sharks are also present in the Simssee
area (i.e. Chlamydoselachus, Centrophorus, Deania, Isis-
tius, and Mitsukurina). Nowadays, only few deep-water
sharks can be found below 1000m in the Mediterranean
Sea (e.g. Centrophorus granulosus and Hexanchus griseus)
(Sion etal. 2004). Therefore, the presence of fossil and their
extant representatives indicates that the sediments of the
Simssee area were deposited in shallow-to-deep shelf envi-
ronments. According to Kroh (2007), the echinoderm fauna
from the early Miocene of the Central Paratethys inhabited
shallow and deep-water environments. The co-presence of
elasmobranchs inhabiting shallow and deep environments
was also reported from other lower Miocene, Pliocene and
Pleistocene European localities (Kocsis 2007; Marsili 2007;
Fulgosi etal. 2009; Reinecke etal. 2011; Pollerspöck and
Straube 2017).
Paleobiogeographic dynamics
The elasmobranch fauna described here experienced paleo-
biogeographic changes from the early Miocene to the recent
(Table1). At genus level, 10% of the recognised genera (3
out of 31) are globally extinct (Araloselachus, Physogaleus
and Premontreia). Comparing the presence of the survived
genera in the Mediterranean Sea today, two biogeographic
dynamics are observed. Fifty percent of the living genera (14
out of 28) are absent in the Mediterranean Sea (Aetobatus,
Sharks, rays and skates from the Upper Marine Molasse (Burdigalian, Miocene) of Bavaria, Germany
1 3
Chaeonagelus, Chlamydoselachus, Carcharodon, Deania,
Galeocerdo, Isitius, Isogomphodon, Notorynchus, Para-
galeus, Pristiophorus, Rhizoprionodon, Rhynchobatus and
Taeniurops), whereas 50% (14 of 28) are still present being
represented by at least one species. For instance, the gen-
era Squalus and Myliobatis are currently represented in the
Mediterranean Sea by the picked dogfish shark S. acanthias
and the common eagle ray M. aquila (Compagno 1988; Last
Table 1 Global status and present-day distribution in the Mediterranean Sea of chondrichthyans from Simsee area
Superorder Order Family Taxa Global status Present-day distribution in the Mediter-
ranean
Batomorphii Myliobatiformes Aetobatidae Aetobatus sp.Living Absent
Squalomorphii Lamniformes Alopiidae Alopias exigua A. superciliosus, A. vulpinus
Squalomorphii Lamniformes Odontaspididae Araloselachus cuspidatus † Absent
Squalomorphii Carcharhiniformes Carcharhinidae Carcharhinus priscus C. altimus, C. brachyurus, C. brevipinna,
C. falciformis, C. limbatus, C. melanop-
terus, C. obscurus, C. plumbeus
Squalomorphii Carcharhiniformes Carcharhinidae Carcharhinus sp. Living C. altimus, C. brachyurus, C. brevipinna,
C. falciformis, C. limbatus, C. melanop-
terus, C. obscurus, C. plumbeus
Squalomorphii Lamniformes Odontaspididae Carcharias acutissimus C. taurus
Squalomorphii Squaliformes Centrophoridae Centrophorus granulosus Living C. granulosus, C. uyato
Squalomorphii Carcharhiniformes Hemigaleidae Chaenogaleus affinis † Absent
Squalomorphii Hexanchiformes Chlamydoselachidae Chlamydoselachus sp. Living Absent
Squalomorphii Lamniformes Lamnidae Carcharodon hastalis † Absent
Batomorphii Myliobatiformes Dasyatidae Dasyatis probsti D. marmorata, D. pastinaca, D. tortonesi
Batomorphii Myliobatiformes Dasyatidae Dasyatis rugosa D. marmorata, D. pastinaca, D. tortonesi
Batomorphii Myliobatiformes Dasyatidae Dasyatis sp. Living D. marmorata, D. pastinaca, D. tortonesi
Batomorphii Myliobatiformes Dasyatidae Dasyatis strangulata D. marmorata, D. pastinaca, D. tortonesi
Squalomorphii Squaliformes Centrophoridae Deania sp. Living Absent
Squalomorphii Carcharhiniformes Carcharhinidae Galeocerdo aduncus † Absent
Squalomorphii Squaliformes Dalatiidae Isistius triangulus † Absent
Squalomorphii Carcharhiniformes Carcharhinidae Isogomphodon acuarius † Absent
Squalomorphii Lamniformes Mitsukurinidae Mitsukurina lineata M. owstoni
Batomorphii Myliobatiformes Myliobatidae Myliobatis sp. Living M. aquila
Squalomorphii Hexanchiformes Hexanchidae Notorynchus primigenius † Absent
Squalomorphii Lamniformes Odontaspididae Odontaspis molassica O. ferox
Squalomorphii Carcharhiniformes Hemigaleidae Paragaleus pulchellus † Absent
Squalomorphii Carcharhiniformes Carcharhinidae Physogaleus contortus † Absent
Squalomorphii Carcharhiniformes Scyliorhinidae Premontreia distans † Absent
Squalomorphii Pristiophoriformes Pristiophoridae Pristiophorus sp. Living Absent
Batomorphii Rajiformes Rajidae Raja sp. Living R. africana, R. asterias, R. brachyura, R.
clavata, R. miraletus, R. montagui, R.
polystigma, R. radula, R. undulata
Batomorphii Myliobatiformes Myliobatidae Rhinoptera sp. Living R. marginata
Squalomorphii Carcharhiniformes Carcharhinidae Rhizoprionodon sp. Living Absent
Batomorphii Rhinopristiformes Rhinidae Rhynchobatus sp. Living Absent
Squalomorphii Carcharhiniformes Scyliorhinidae Scyliorhinus fossilis S. canicula, S. duhamelii, S. stellaris
Squalomorphii Carcharhiniformes Scyliorhinidae Scyliorhinus sp. Living S. canicula, S. duhamelii, S. stellaris
Squalomorphii Carcharhiniformes Sphyrnidae Sphyrna sp. Living S. lewini, S. mokarran, S. tudes, S.
zygaena
Squalomorphii Squaliformes Squalidae Squalus sp. Living S. acanthias
Squalomorphii Squatiniformes Squatinidae Squatina sp. Living S. aculeata
Batomorphii Myliobatiformes Dasyatidae Taeniurops cavernosus † Absent
J. A. Villafaña etal.
1 3
etal. 2016). At the species level, all the recognized species
but one (i.e. Centrophorus granulosus) are globally extinct.
Although most of the recognized species have disappeared
from the Mediterranean Sea, different biogeographic dynam-
ics are observed by considering replacement by congeneric
species. Fifty-five percent (11 of 20) of the extinct species
were not replaced by congeneric species living today in the
Mediterranean Sea. On the contrary, 45% (9 of 20) of the
extinct species have been substituted by at least one conge-
neric species living in the present-day Mediterranean Sea.
For instance, Alopias exigua is globally extinct, but the
extant thresher shark Alopias vulpinus currently occurs in
the Mediterranean Sea (Compagno etal. 2005). All these
biogeographic dynamics could have been influenced by
intense tectonic, climatic and oceanographic events dur-
ing the early Miocene of Europe (Rögl 1999). According to
Kroh (2007), the climatic and oceanographic changes (i.e.
drop of temperature and sea level changes) were the major
factors controlling the distribution of echinoderm faunas
during the early Miocene of the Central Paratethys, and may
have also affected elasmobranchs. This idea was previously
hypothesized as a possible cause of chondrichthyan distri-
butional changes in South American localities during the
Neogene (Long 1993; Cione etal. 2007; Carrillo-Briceño
etal. 2013; Villafaña 2015; Partarrieu etal. 2018; Villafaña
and Rivadeneira, 2014, 2018; Villafaña etal. 2019).
Faunal comparison duringtheearly Miocene
According to our faunal comparison, the fauna from the
Simssee area was more similar to closely adjacent localities
in Europe rather than to other localities (Fig.11 and Table2).
The most similar faunas are from Switzerland (76%), Austria
(71%), France (62%), North Germany (59%) and Portugal
(59%). These high similarities could be related to the shorter
distances and connection between the localities. During the
Ottnangian, the western and Central Paratethys were con-
nected through the Rhine Graben (Rögl 1999). According to
Kocsis (2007), the presence of deep-water sharks such as Mit-
sukurina and Isistius should be the evidence of large and con-
nected open water surfaces, with deeper sea basin during the
Eggenburgian–Ottnangian stages. According to Pollerspöck
and Straube (2017), the diversity of fishes from the Paratethys
was shaped by immigration of taxa from other marine ecosys-
tems and favoured by oceanographic variables such as salinity
and oxygen contents. Additionally, dispersal of some sharks
could be also possible into freshwater environment (Koc-
sis etal. 2007). Therefore, the seaway passages could have
favoured the connection of elasmobranch faunas from differ-
ent localities and explain their similarities. On the contrary,
the faunas with the lowest similarities were from Colombia
(20%), Panamá (23%), Spain (24%) and Brazil (24%), pos-
sibly reflecting to the long distances between the localities.
Despite the apparent connection of Italy, Malta, and Spain
with southern Germany during the Ottnangian (Rögl 1999),
their lower similarity could be affected by sampling biases (i.e.
incompleteness of the taxonomic inventories). However, areas
where sampling is expected to be high (e.g. the Pacific coast
of USA, Australia) also show low similarity values, due to the
relatively low generic richness. In the case of India (29%), the
seaway passage between the Indo-Pacific and the Paratethys
was closed during the Ottnangian, probably explaining its
lower similarity to the fauna from Simssee.
Fig. 10 Rarefaction curve and taxonomic richness of the chondrich-
thyan fauna from Simssee
Fig. 11 Paleogeographic distribution and faunal similarity between
the Simssee fauna and others early Miocene faunas. Simssee (a),
Northern Germany (b), Switzerland (c), Austria (d), Slovakia (e),
France (f), Portugal (g), Italy (h), Malta (i) and Spain (j). Map based
on Rögl (1999)