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© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
NEW OLIGOCENE TURTLE REMAINS
OF THE OBERLEICHTERSBACH DOLINE FILLING
(LOWER FRANCONIA, GERMANY) AND
REVISION OF THE GENUS PALAEOMAUREMYS
(TESTUDINES: GEOEMYDIDAE)
[Nuevos ejemplares del Oligoceno superior de los sedimentos lacustres de la dolina
Oberleichtersbach (Baja Franconia, Baviera, Alemania)]
Hans-Volker Ka r l *,**
Elke Gr ö n i n G ***
Carsten Br a u c K M a n n ***
(*) Thüringisches Landesamt für Denkmalpflege und Archäologie. Humboldtstraße 11.
D-99423 Weimar. Correo-e: hvkarl@web.de
(**) Geoscience Centre of the University of Göttingen. Department Geobiology. Golds-
chmidtstrasse 3, D-37077 Göttingen.
(***) Institut für Geologie und Paläontologie. TU Clausthal. Leibnizstrasse 10. D-38678
Clausthal-Zellerfeld. Germany. Correo-e: elke.groening@tu-clausthal.de; carsten.brauc-
kmann@tu-clausthal.de
(FE c H a d E r E c E p c i ó n : 2012-01-26) (FE c H a d E ad Mi si ón : 2012-01-30)
BIBLID [0211-8327 (2011) 47 (2); 175-194]
ABSTRACT: New Late Oligocene (Chattian) remains of the terrapins Palaeoemys
hessiaca Schleich, 1994 and Palaeomauremys tuberculata (Portis, 1882), the soft-she-
lled turtles Allaeochelys parayrei Noulet, 1867 and Trionyx cf. triunguis Forskål,
1775, and the snapping turtle Chelydrasia decheni (H. V. Meyer, 1852) from lacus-
trine sediments of the Oberleichtersbach doline (Lower Franconia, Bavaria,
Germany) are reported. The morphological features of these five species, their
taxonomic position and their palaeobiological implications are discussed. The
new Palaeomauremys material suggests that P. mlynarskii (Hervet & Lapparent de
Broin, 2000) is a junior synonym of P. tuberculata (Portis, 1882).
Key words: Testudines, Palaeoemys hessiaca Schleich, 1994, Palaeo maur emys tuber-
culata (Portis, 1882), Allaeochelys parayrei Noulet, 1867, Trionyx cf. triunguis Forskål,
ISSN: 0211-8327 Studia Geologica Salmanticensia, 47 (2): pp. 175-194
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
176
1775, Chelydrasia decheni (H. V. Meyer, 1852), Late Oligocene, Chattian, Oberleichters-
bach, Lower Franconia, description, palaeo biology.
RESUMEN: En este artículo se estudian nuevos ejemplares del Oligoceno superior
de los sedimentos lacustres de la dolina Oberleichtersbach (Baja Franconia, Bavie-
ra, Alemania), determinados como de los galápagos Palaeoemys hessiaca Schleich,
1994 y Palaeomauremys tuberculata (Portis, 1882), las tortugas de caparazón
blando Allaeochelys parayrei Noulet, 1867 y cf. Trionyx triunguis Forskål de 1775,
y la tortuga mordedora Chelydrasia decheni (V. H. Meyer, 1852). Se discuten las
características morfológicas de estas cinco especies, su posición taxonómica y sus
implicaciones paleobiológicas. El nuevo material de Palaeomauremys sugiere que
P. mlynarskii (Hervet y Lapparent de Broin, 2000) es un sinónimo de P. tubercu-
lata (Portis, 1882).
Palabras clave: Testudines, Palaeoemys hessiaca Schleich, 1994, Palaeomauremys
tuberculata (Portis, 1882), Allaeochelys parayrei Noulet, 1867, Trionyx cf. triunguis
Forskål, 1775, Chelydrasia decheni (H. V. Meyer, 1852), Oligoceno superior, Chattien-
se, Oberleichtersbach, Baja Franconia (Alemania), descripción, paleobiología.
INTRODUCTION
From the German terrestrial and limnic deposits of Late Oligocene (Chat-
tian) age the terrapin turtle Palaeomauremys mlynarskii (Hervet & Lapparent
de Broin, 2000) was already described from Rott (North Rhine-Westphalia),
Enspel (Westerwald: Rhineland-Palatinate), and Oberleichtersbach (Bavaria:
Lower Franconia). In the meantime, additional material is available from the
Walter Heck collection (Oberleichtersbach) which enables the revision of
Emys tuberculata Portis, 1882, a species of previously uncertain status which
was assigned to Palaeomauremys Hervet, 2004 by HE r v E t (2004) and Ka r l &
WE t t l a u F E r (2010). Based upon the now known morphological features in
the new specimens, both taxa can easily be synonymized on generic level.
The genesis and topography of the Late Oligocene (Chattian) doline filling
of Oberleichtersbach near Bad Brückenau (fig. 1) is discussed in detail by Ma r -
t i n i (2008) which contribution is explicitely referred to. In the same volume
Bö H M E (2008) presented a complilation of 43 taxa of fish, amphibians and rep-
tiles. The first consist of six species in two families: five species of Cyprinidae
and the new Cobites primigenus Böhme, 2008 (Cobitidae). The amphibians
include eight species of Allocaudata and Urodela as well as six species of
Anura. The reptiles are present with one species of Diplocynodon (Crocodylia)
and 15 species of Squamata: Lacertilia, Anguimorpha, and Ophidia, the latter
with the new Texasophis hecki Böhme, 2008. Many of these species present the
most ancient record of their groups, one of them is the latest one. With this
content of fossils, this locality shows the highest diversity of ectothermic verte-
brates of the Late Oligocene world-wide. Additionally there are six species of
Testudines. One pleural of “Palaeomauremys mlynarskii” is figured (pl. 4, fig.
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
177
7a-b), the other taxa are listed without figures and catalogue numbers. Here it
is referred to under the discussion of the species in question.
This paper is supplement 180 of chapter “C. Fourth faunal change: Paleo-
gene-Neogene (P/N) boundary/V. Rise of stream terrapins”, in: Studia
Palaeocheloniologica, IV (Ka r l , 2011).
Figure 1. Geographical positions of the Palaeomauremys- localities: 1= Rochette near
Lausanne, Switzerland, type locality of Emys tuberculata Portis, 1882; 2= Trbovlje (former
Trifail) in Slovenia, type locality of Testudo riedli Hoernes, 1892; 3= Wuhrberg south of
Windischgarsten, Austria; 4= Rott, Germany, type locality of Palaeochelys mlynarskii Hervet
& Lapparent de Broin, 2000; 5= Enspel, Germany; 6= Oberleichtersbach, Germany. Among
the localities 4 and 5, see also the Geological Map of the Tertiary Westerwald, which was
taken by KA r l & We t t l A u f e r (2011) in sc h i n d l e r & Wu t t K e (2010).
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
178
SYSTEMATIC PALAEONTOLOGY
Ordo Testudines Linnaeus, 1758
Suprafamilia Testudinoidea Batsch, 1788
Familia Geoemydidae tH E o B a l d , 1868 (= Bataguridae Gray, 1869)
Subfamilia Palaeochelyinae Karl & Wettlaufer, 2011
Genus Palaeoemys Schleich, 1994
– 1994 Palaeoemys n. g. Schleich: 82-87.
– 2004 Palaeoemys Schleich, 1994 - HE r v E t : 65-70.
– 2004 Palaeoemys Schleich, 1994 - cl a u d E & to n G : 6.
– 2011 Palaeoemys Schleich, 1994 - Ka r l & WE t t l a u F E r : 179.
Type species: Palaeoemys hessiaca Schleich, 1994: 82.
Additional species: Palaeoemys occitana Hervet, 2004, Palaeoemys testu-
diniformis (Owen, 1842) emend. cl a u d E & to n G (2004).
Geographic and stratigraphic distribution:
Early Eocene (MP9 to MP 10 boundary), Saint-Papoul near Castelnaudary
(Aude, France) and Monthelon near Epernay (Marne, France: MP 19). Middle
Eocene (MP11 to MP 13), Borken (northern Hesse, Germany).
Original diagnosis: scHlEicH (1994: 82). With the present specimen the
stratigraphic distribution is extended to the Late Oligocene and thus to the
late Palaeogene.
3.1.1. Palaeoemys cf. hessiaca Schleich, 1994
– 2008 Emydidae indet. Böhme: 168.
New material: OLB 2-1, rigth epiplastron in ventral view (plate 4, figs. 1-2);
OLB 2-2, rigth pleural I remain (plate 4, figs. 3-4); OLB 2-3, pleural remain (pla-
te 4, figs. 5-6); OLB 4-1, phalanx III (plate 4, fig. 7); OLB 4-2, scapula remain
(plate 4, fig. 8); OLB 2-4, left hyoplastron remain (plate 4, figs. 9-10); OLB 2-4,
left hyoplastron remain (plate 4, figs. 9-10).
Discussion: The hitherto single record of Palaeoemys. Further specimens
possibly belonging to this taxon are too fragmentary for a definite assignment.
The ventral and visceral course of the gular suture, the furrow of the skin
margin and the extremely week gular bulge exactly resemble the known mor-
phology of this genus (see HE r v E t , 2004; sc H l E i c H , 1994), but due to the pre-
servation the specimen is only tentatively assigned to P. hessiaca.
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
179
Plate 1. Palaeomauremys tuberculata (Portis, 1882) from Oberleichtersbach, ex Coll.
Walter Heck, carapace remains: 1= OLB 1-1, anterior carapace remain in dorsal view,
2= in visceral view; 3= OLB 1-2, left pleural II in dorsal view, 4= in visceral view; 5= OLB
1-3, pygal in dorsal view; 6-8= OLB 1-4, OLB 1-5, OLB 1-6, bridge peripherals in dorsal
view. Scale bar 3 cm. Photo Brigitte Stefan, design Heike Künzel (TLDA).
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
180
3.2. Genus Palaeomauremys Hervet, 2004
Selected synonymy:
– 1882 Emys - po r t i s : 19-20.
– 2000 Palaeochelys Schleich 1994 - HE r v E t & la p p a r E n t d E Br o i n : 563.
– 2004 Palaeomauremys n. gen. HE r v E t : 54-56.
– 2011 Palaeomauremys Hervet, 2004 - Ka r l & WE t t l a u F E r : 175.
Type species: Emys tuberculata Portis, 1882.
Original description: po r t i s (1882: 19-20, pl. 4).
Type locality: Rochette near Lausanne (Switzerland).
Type stratum: Subalpine Molasse (Molasse d’eau douce inférieure, USM),
Late Oligocene: MP 28-30 (En G E s s E r , 1990), MP 29 (En G E s s E r & Mö d d E n , 1997).
Holotype: MGL 8901 (= po r t i s , 1882, pl. 4), anterior fragment of carapace
+ fragments of plastron (pl. 76, figs. E, F, G and H).
Emended description by HE r v E t (2004): “Le repli ventral de la cervicale est
de longueur moyenne sur la face interne de la nucale, il est très large (Pl. 76,
fig. G). Le repli ventral des M1 est assez long et il à présence de deux légers
bourrelets transversaux sur la face interne de la nucale. La face interne de la PL1
montre l’insertion du pilier axillaire sur environ un tiers de la larger de la pleurale
(Pl. 76, fig. H); il y a un léger bourrelet osseux sur la PL1, médialement à cette
insertion”. For the generic diagnosis of the carapace: see HE r v E t (2003, 2004),
in particular map 4.11 and chapter 4 in the paper in 2004.
Remarks: Most important synonym for the region in question is Palaeochelys
mlynarskii (Hervet & Lapparent de Broin, 2000) from the Late Oligocene (MP
30: Bö H M E & la n G , 1991; Mö r s , 1995) of the lignites of Rott near Bonn (North
Rhine-Westphalia, Germany; see: HE r v E t & la p p a r E n t d E Br o i n , 2000; HE r v E t
2003 and 2004, with generic diagnosis). For the Enspel specimen Exemplar:
see Ka r l & WE t t l a u F E r (2011). A small species with a carapace length of ori-
ginally only 125 mm, but the present material suggests a length of more than
200 mm.
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
181
Figure 2. Reconstruction of Palaeomauremys tuberculata (Portis, 1882) from
Oberleichtersbach: 1= anterior carapace region based on OLB 1-1 (plate 1, fig. 1);
2= anterior half plastron based on OLB 1-9 (plate 2, fig. 7); 3= caudal lobus of plastron
based on OLB 1-12 (plate 2, fig. 10). Carapace plates (left side): nuchal = nu, neurals
= n, pleurals = pl, peripherals = pe. Carapace scutes (rigth side): cervical = ce, centrals
= c, laterals = l, marginals = m. Plastron plates (left side): epiplastrals = epi, entoplastron
= ento, hyoplastron = hyo, xiphiplastron = xiphi. Plastron scutes (rigth side): gulars = gu,
humeral = hu, pectorals = pec, abdominals = ab, femorals = fe, annals = an.
Graphic H.-V. KA r l & Heike Kü n z e l (TLDA).
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
182
Plate 2. Palaeomauremys tuberculata (Portis, 1882) from Oberleichtersbach, ex Coll. Walter
Heck, plastron remains: 1= OLB 1-10, rigth epiplastron in ventral view, 2= in visceral view;
3= OLB 1-11, left epiplastron in ventral view, 4= in visceral view; 5= OLB 1-12, left
xiphiplastron in visceral view, 6= in ventral view; 7= OLB 1-9, rigth anterior plastral lobe in
ventral view, 8= in visceral view; 9= OLB 1-13, rigth xiphiplastron remain in ventral view,
10= in visceral view. Scale bar 3 cm. Photo Brigitte Stefan, design Heike Künzel (TLDA).
3.2.1. Palaeomauremys tuberculata (Portis, 1882)
Selected synonymy:
– 1882 Emys tuberculata n. sp. Portis: 19-20; pl. 4.
– 1892 Testudo Riedli nov. form. Hoernes: 243 (synonyms in Ka r l , 1994/95).
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
183
– 2000 Palaeochelys mlynarskii n. sp. Hervet & Lapparent de Broin: 563-569;
figs. 1-3.
– 2004 Palaeomauremys «tuberculata» (Portis, 1882) - HE r v E t : 163.
– 2011 Palaeochelys mlynarskii Hervet & Lapparent de Broin, 2000 - Ka r l
& WE t t l a u F E r : 175-177.
– 2011 Palaeomauremys “tuberculata” (Portis, 1882) - Ka r l & WE t t l a u F E r : 175.
Geographic and stratigraphic distribution: (i) Germany: Top of Late Oli-
gocene (MP 30), Rott near Bonn (North Rhine-Westphalia); Enspel Maar (Rhi-
neland-Palatinate: Westerwald); doline filling at Oberleichtersbach (Bavaria:
Lower Franconia; Ka r l & WE t t l a u F E r , 2011).
(ii) Slovenia: Trbovlje (former Trifail).
New material: OLB 1-1, anterior carapace remain (plate 1, figs. 1-2); OLB
1-2, left pleural II (plate 1, figs. 3-4); OLB 1-3, pygal (plate 1, fig. 5); OLB 1-4,
OLB 1-5, OLB 1-6, bridge peripherals (plate 1, figs. 6-8); OLB 1-9, rigth ante-
rior plastral lobe (plate 2, figs. 7-8); OLB 1-10, rigth epiplastron (plate 2, figs.
1-2); OLB 1-11, left epiplastron (plate 2, figs. 3-4); OLB 1-12, left xiphiplas-
tron; OLB 1-13, rigth xiphiplastron remain (plate 2, figs. 9-10).
Main characters of the new material: Length of carapace = 200 mm. Cervi-
cal distinctly developed, visceral part larger than dorsal part; epiplastron stron-
gly vaulted, with distinctly marked geoemydaloid gular bulge; gular furrows
cutting entoplastron, humeropectoral furrows not; entoplastron 1,2 x broader
than long, rhomb-like, anteriorly markedly pointed; anal notch obtuse-angled;
pygal geoemydaloid; neurals emydoidal and geoemydaloid; three (to maybe
five) strong dorsal keels, no keel on bridge developed; bridge piers week,
inguinal piers inserting into pleural I via peripherals II and III, peripherals III
with distinct visceral moschus channel, all marginal furrows situated on peri-
pherals; typical geoemydaloid structure of areols on carapace present, surface
of carapace with coarse sculpture, partly with very deep furrows on the horn-
scutes. In fragmentary material the latter can feign the presence of terrestrial
turtles: “Several plastron and carapace remains belong to an undetermined
species of Testudo…, and a second land turtle known only by one plastron
fragment (not figured). It differs from Testudo in having rising (= geochelonid
according to sc H l E i c H 1988) sutural furrows, typical for large land turtles of the
genus Geochelone sensu lato” (Bö H M E , 2008).
Ka r l (1994/95) grouped Testudo riedli Hoernes, 1892 with Clemmydopsis
Boda, 1927. Based upon the new material from Oberleichtersbach it is almost
sure that it really belongs to Palaeomauremys. There are exactly the same mor-
phological features in the anterior part of the internal mold from Trbovlje (former
Trifail) in Slovenia and in the anterior visceral region of the carapax from Ober-
leichtersbach. Even the structure of the plastron is the same. The close stratigra-
phical age of the Slovenian and German specimens support their close relations-
hips (plate 3, figs. 1-2). According to ri J a v E c & do z E t (2002) the limno-brackish
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
184
Socka Beds (former Sotzkaschichten) are subdivided into the Lower Socka Beds,
the main coal horizon, and the Upper Socka Beds. They are overlain by the Late
Oligocene Marine Marly Clay, dacito-andesitic tuff and pyroclastic breccia. The
stratigraphical position is Egerian (= Late Oligocene: Chattian to the Oligocene/
Miocene boundary beds). For a detailed description and the synonymy of the
taxon see Ka r l (1994/95) and Gr o s s (1994). The Egerian in the Paratethys largely
corresponds with the late Chattian and the complete Aquitanian (25,8-20,3 MY)
(Ba l d i , 1975) and includes the “Gosau-Schichten”.
During railway constructions the visceral mold of a turtle carapace was
discovered in a quarry for building stones in the “Gosau-Schichten” at the
Wuhrberg S Windischgarsten (Upper Austria) and donated to the k. k. Geolo-
gischen Reichsanstalt in Vienna by Ing. Gross (Spital, Pahrn, Austria) in 1905.
The comparable characters completely correspond with those of the steinkern
from Trbovlje (former Trifail, Slovenia). Thus the Wuhrberg specimen has to
be assigned to Palaeomauremys, too (plate 3, fig. 3). With a length of 254 mm
and a width of 186 mm this carapace is the largest known.
Plate 3. Palaeomauremys tuberculata (Portis, 1882): 1= Steiermärkisches Landesmuseum
Joanneum, Graz n.º 5.908, Steinkern from Trbovlje (former Trifail) in Slovenia, holotype
of Testudo riedli Hoernes, 1892 from Gr o s s (2002) with permission by the author;
2= Geological Survey of Austria/Department of Paleontology n.º #, visceral mold of carapace
from Wuhrberg south of Windischgarsten/ Oberösterreich.
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
185
Suprafamilia Trionychoidea Fitzinger, 1826
Familia Carettochelyidae Boulenger, 1887
Subfamilia Carettochelyinae Boulenger, 1887
3.3. Genus Allaeochelys Noulet, 1867
Synonymy: See Ku H n (1964), Br o i n (1977) and lap p a r E n t d E Br o i n (2001).
Type species: Allaeochelys parayrei no u l E t , 1867 (original generic desig-
nation); originally described from Eocene deposits in southern France.
Diagnosis: See Mł y n a r s K i (1976: 73-74).
Remarks: Neither the genus name Allaeochelys nor its author no u l E t (1867)
were listed in the “Fossilium Catalogus” on “Trionychia fossilia” (Hu M M E l , 1932)
(see Ka r l et al., 2006).
Geographic and stratigraphic distribution: Early to late Middle Eocene in
southern France, Spain, England, Belgium and Germany, in Germany up to the
Late Oligocene (Ka r l et al., 2006).
3.3.1. Allaeochelys parayrei Noulet, 1867
Selected synonymy:
– 1956 Anosteira crassesculpta Harrassowitz, 1922 - Gr a M a n n : 18.
– 1956 Anosteira crassesculpta = gracilis Harrassowitz, 1922 - Ku H n : 181.
– 1986 “Anosteira” aff. crassesculpta Harrassowitz, 1922 - sc H l E i c H : 285.
– 2006 Allaeochelys parayrei Noulet, 1867 (syn. Anosteira crassesculpta et
A. gracilis Harrassowitz, 1922) - Ka r l , Gr ö n i n G & Br a u c K M a n n : 51-57.
– 2007 Allaeochelys Noulet, 1867 (syn. Anosteira crassesculpta Harrassowitz,
1922) - Ka r l , 2007: 61.
New material: Left dental (adult) and right xiphiplastron (juvenile).
Brief description: Sculpture on xiphiplastron flat and reticular, alveolar
plain of dental without sculpture (without marked chewing ridges). Fossa mec-
keli open until symphysis. Labial margin with the typical porous structure of
Trionychoidea.
Familia Trionychidae Fitzinger, 1826
Subfamilia Trionychinae Fitzinger, 1826
Subtribus Trionychina Fitzinger, 1826
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
186
3.4. Genus Trionyx Geoffroy Saint Hilaire, 1809
Synonymy: See Ka r l (1998), older synonymy in Hu M M E l (1929, 1932), lo-
v E r i d G E & Wi l l i a M s (1957), WE r M u t H & ME r t E n s (1961), Ku H n (1964), and
iv E r s o n (1992).
Type species: Testudo triunguis Forskål, 1775.
Terra typica of type species: “In Nilo rarior” [= Nile River] (iv E r s o n , 1992).
Diagnosis: See Ka r l (1998).
Remarks: As pointed out by Ka r l (1999), the Eocene/Oligocene transition
is marked by a change in the Trionychidae in Central Europe. The Eocene
Nearctic Rafetoides austriacus (Peters, 1868) disappear and open wide space
for the oriental species Trionyx gregarius/triunguis. The latter becomes the do-
minant species of the Neogene Palaearctic an is now restricted to a postglacial
relict areal in Ethiopia (for the history of distribution see Ka r l , 1999). Trionyx
triunguis is also known from the Early Oligocene sequence in the Weißelster
Basin (Saxonia, Germany; Ka r l , 2007). On the other hand, in the Early Oligo-
cene “Melanien-Ton” of Borken (Hesse, Germany) Rafetoides austriacus still
occurs (Ka r l & MüllE r , 2008).
Geographic and stratigraphic distribution:
(i) Late Oligocene to Miocene: Austria, Slovenia, southern and Central Ger-
many, France, Hungary, Romania, Bohemia, Switzerland.
(ii) Miocene to Pliocene: Germany, Egypt.
(iii) Tertiary (without exact data): Núrpur in Nepal.
(iv) Prehistoric: Egypt.
(v) Recent: Southern Turkey to Africa (Senegal to Angola, Somalia and
Egypt) (iv E r s o n , 1992).
The additional species Trionyx gregarius (syn. Amyda gregaria Gilmore,
1931) was mentioned from the Early and “Middle” Oligocene Houldjin Forma-
tion at Camp Margetts, 25 miles SW Iren Babasu, Inner Mongolia (= Neimeng-
gu, northern China).
3.4.1 Trionyx cf. triunguis (Forskål, 1775)
Remarks: According to Bö H M E (2008) “the rearest turtle in Oberleichters-
bach are the soft-shell or river turtle Trionyx sp., documented by one charac-
teristically sculptured pleural fragment (not figured)“. Here only a single free
pleural cone, showing the structured transition of the proximal part to the
pleural plate.
Superfamilia Chelydroidae Gray, 1831
Familia Chelydridae Agassiz, 1857
Subfamilia Chelydropsinae Młynarski, 1980
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
187
3.5. Genus Chelydrasia Chkhikvadze, 1999
Synonymy: See Ku H n (1964), d E Br o i n (1977), dE la p p a r E n t d E Br o i n
(2001) and Hu t c H i s o n in st E y E r M a r K et al. (2008).
Type species: Chelydropsis minax Chkhikvadze, 1971 (?= Chelydra decheni
H. V. Meyer, 1852).
Diagnose: cH K H i K v a d z E (1999).
Geographic and stratigraphic distribution: Early Eocene to Pliocene in Ka-
zakhstan; Middle Eocene to Early Miocene in Europe (Hu t c H i s o n in st E y E r -
M a r K et al., 2008; Ka r l et al. in print).
Remarks: cH K H i K v a d z E (1999) established Chelydrasia with its original type
species Chelydropsis minax Chkhikvadze, 1971 from the Early Oligocene of the
Zajsan Basin in Eastern Kazakhstan. He also assigned Ch. sanctihenrici and all
the species of the so-called decheni-sanctihenrici group. Furthermore, he as-
sumed an uncertain taxon from Artenay (MN4) to be closely related. According
to Hu t c H i s o n in stE y E r M a r K et al. (2008) Chalydrasia now includes Ch. minax
Chkhikvadze, 1971, Ch. poena Chkhikvadze, 1971, Ch. apellanzini (Murelaga
et al., 1999), Ch. decheni (H. V. Meyer, 1852), Ch. sanctihenrici (De Broin,
1977), “Ch.” kusnetzovi (Chkhikvadze, 1985). Judging from the preservation,
the differences between Ch. minax and Ch. decheni seem to be not definitely
clarified. If Ch. minax is regarded a junior synonym of Ch. decheni the latter
becomes the type species.
3.5.1. Chelydrasia decheni (H. V. Meyer, 1852)
Synonymy: See Ku H n (1964), dE Br o i n (1977), dE la p p a r E n t d E Br o i n
(2001), Ka r l et al. (in print).
New material: OLB 3-1, left pleural I in dorsal view in situ at grey chalky
sediment (plate 4, fig. 11); OLB 3-2, pleural remains in dorsal view in situ at
reddish chalky sediment with many gastropods (plate 4, fig. 12).
Brief description: The dimensions of pleural I are superficially similar to
those of Palaeomauremys, but with a thickness with no more than 1 mm in
its proximal region it is distinctly thinner. The plate thickens towards the mar-
gin but does not even approximately reaches the dimensions of the following
one. The dorsal plain is smooth, with a few areoles only towards the lateral
margin, but without any rudiments of keels. The latter first appear on a few
smaller fragments of pleurals.
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
188
Plate 4. Palaeoemys cf. hessiaca Sc h l e i c h , 1994 from Oberleichtersbach, ex Coll. Walter
Heck: 1= OLB 2-1, rigth epiplastron in ventral view; 2= in visceral view; 3= OLB 2-2, rigth
pleural I remain in visceral view, 4= in dorsal view; 5= OLB 2-3, pleural remain in dorsal
view; 6= in visceral view; 7= OLB 4-1, phalanx III; 8= OLB 4-2, scapula remain;
9= OLB 2-4, left hyoplastron remain in ventral view, 10= in visceral view.
Chelydrasia decheni (H. V. Meyer, 1852) from Oberleichtersbach, ex Coll. Walter Heck:
11= OLB 3-1, left pleural I in dorsal view in situ at grey chalky sediment; 12= OLB 3-2,
pleural remains in dorsal view in situ at reddish chalky sediment with many gastropods.
Scale bar 3 cm. Photo Brigitte Stefan, design Heike Künzel (TLDA).
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
189
CONCLUSIONS
PALAEOBIOLOGY
The palaeobiological conditions of the extinct genus Palaeomauremys are
supposed to be very similar to those of the Recent and fossil Mauremys.
Today these Palaeochelyinae inhabit calm smaller water systems as pools, dit-
ches, or muddy brooks from the lowland up to the mountainous regions.
Recent species are omnivorous (see for example Ka r l & WE t t l a u F E r , 2011).
The most detailed climatic data, based upon plant remains, were pre-
sented by Ma i (2008). The annual average temperature varied between 10
and 16 °C, the warmest month reached about 22 to 28 °C whereas the tempe-
rature of the coldest month changed from about –3 to –8 °C. The annual pre-
cipitation ranged presumably from 630 to 1040 mm, with one or two dry
months of only 18 to 7 mm.
Comparable Recent conditions are known only from a few Chinese wea-
ther stations, as for example Enshi (= Engshih) in Hubei (= Hupeh) as well as
Tai Shan and Jinan (= Tsinan) in Shandong (= Schantung). For the time inter-
val between 23.5 and 25 million years ago Mai (2008) proposed a new and
separate “floral complex Oberleichtersbach” which is a particular Carpino-
Ostryon (MMF) assemblage including semihumid species, typical floral ele-
ments of dry summers and a great number of annual plants of open places.
According to Bö H M E (2008), the fauna suggests the existence of a large,
shallow, permanent and oxygen-rich lake with intensively structured shore and
an afflux. The faunal climatic conditions were considered to be subtropical and
humid which is not completely in accordance with the palaeobotanical data.
SYSTEMATICS AND PALAEOGEOGRAPHY
In contrast to other authors, cl a u d E & to n G (2004) synonymized a large
number of generic names of the Palaeochelyinae with Palaeoemys which is
regarded as a taxon different from Mauremys. In particular they assign
Palaeoemys to the so-called Malayemys complex, including Malayemys,
Geoclemys, Palaeoemys, and ?Borkenia. On the other hand, Mauremys is allo-
cated to the Melanochelys complex, including Melanochelys, Mauremys, Saca-
lia, Cuora, Cistoclemmys, Notochelys, Heosemys, Hieremys, Cyclemys, Leuco-
cephalon, Ocadia, Chinemys, Palaeochelys, ?Siebenrockiella and ?Geoemyda.
These different views implicate a hotspot on the European spit during the
Paleogene as it was already pointed out by Ka r l (1998, 1999) and da n i l o v
(2005) for the Trionychidae. Likewise this seems to be true for the
Palaeochelyinae. Within the whole relationship distinct tendencies can be
recognized for splitting into the generic complex Mauremys/Ocadia, the Pty-
chogastrinae and the Geoemydinae, still combined with large overlapping
(see for example KH o z a t s K y & Ml y n a r s K i , 1966). Surely this problem cannot
H.-v. Kar l , E. Gr ö n i n G & c. Br a u c K M a n n
New oligocene turtle remains of the Oberleichtersbach doline filling (Lower Franconia, Germany)
and revision of the genus Palaeomauremys (Testudines: Geoemydidae)
© Ediciones Universidad de Salamanca Stud. Geol. Salmant., 47 (2), 2011: pp. 175-194
190
yet finally dissolved since Br o p H y et al. (2006) show twelve cases of interge-
neric hybridisation, where the following genera are connected by genetic
overlapping: Mauremys x Chinemys, Mauremys x Cuora, Mauremys x Cycle-
mys, Mauremys x Heosemys, Mauremys x Sacalia, Cuora x Geoemyda und
Cuora x Sacalia, i.e. the complete complex with its fossil roots in the
Palaearctic. Indeed, even recently often different “hybrid species” occur in
open nature, as for example Chinemys megacephala, Mauremys pritchardi,
Ocadia glyphistoma, Ocadia philippeni, Mauremys iversoni and others. The
latter is even breeded in Chinese turtle farms as a hybrid of Mauremys mutica
x Cuora trifasciata and was already discussed under taxonomic respect
(FonG & cHEn, 2010; parHaM & cH i , 2001). Consequently, Fritz & Havaš
(2007) regarded Chinemys, Sacalia und Ocadia etc. as synonyms of Maure-
mys as a conclusive result of a genetic analysis of single individuals, just in
contrast to the comparison of morphologically studied series. Once more, this
clearly shows that precipitate “results” based only on genetic analyses are not
sufficient as they are not compatible with morphological conclusions.
ACKNOWLEDGEMENTS
Unser herzlicher Dank gilt Frau Dr. Irene Zorn, Geologische Bundesanstalt
Wien, Dr. Martin Gross, Steiermärkisches Landesmuseum Joanneum Graz und
Herrn Walter Heck, Oberleichtersbach für die freundliche Bereitstellung des
Materials aus ihren Sammlungen.
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