ACTA PALAEONTOLOGICA ROMANIAE V. 7 (2011), P. 343-349
MIDDLE-LATE MIOCENE SNAKES FROM THE PANNONIAN BASIN
MÁRTON VENCZEL 1
Abstract. The fossil record of Astaracian–early Vallesian (MN 6–MN 9) snakes is reviewed based on information
derived from sixteen localities, situated on or adjacent to the former Pannonian Sea margins. The studied deposits
yielded both Scolecophidia and Alethinophidia (Boidae, Colubridae, Elapidae, Viperidae). Indeterminable Scolecophidia
were present in most localities, their remains consisting of trunk vertebrae only. The only Boidae recorded is a smallsized Erycinae snake (Albaneryx cf. volynicus) from Felsőtárkány 2, N-Hungary, which represents an important link
between West- and East-European members of the genus. Coluber pouchetii, a member of the “large-sized colubrines”
with still unresolved relationships, apparently was widely distributed across the European continent, whereas Coronella
miocaenica, Hierophis cf. hungaricus and Natrix cf. rudabanyaensis possibly were newcomers from the Asiatic
continent. Small-sized Elapidae, resembling the North-American genus Micrurus, were recorded on the basis of
presacral vertebrae only. Macrovipera sp., from the “Oriental vipers group” is known from the Tau and Vârciorog
localities, W-Romania, whereas the vipers from the “V. aspis group” are known from Mátraszőlős 3, Subpiatră 2/1 and
Felsőtárkány-Felnémet 2/3. The first occurrence of the “V. berus group” in Central Europe is reported from Felsőtárkány
(1, 2 and 3/10 localities). The Astaracian–early Vallesian snake faunas of the Pannonian Basin were already dominated
by modern colubrids (Natricinae and Colubrinae). Increasing faunal influence from the Asiatic continent is presumed to
have started during the early Vallesian.
Keywords: Serpentes, Miocene, Fossil record, Hungary, Romania
INTRODUCTION
The evolution of snake faunas was continuously
influenced by climate and environmental changes as well
as by tectonic rearrangement of continental plates. A
scenario for the origin and history of European Cenozoic
snakes was published by Ivanov et al. (2000), outlining the
major changes which led to the extinction of ancient taxa
and to the emergence of extant genera and species. The
modern colubrids became the main component of the
snake assemblages from the Miocene onward, while after
the so-called Miocene Climatic Optimum (18-14 Ma:
Ottnangian–Middle Badenian) (see Böhme, 2003) the nonErycinae Boidae disappeared completely from the fauna.
The aim of the present paper is to review the fossil
record of snakes from a series of Astaracian–Vallesian
microvertebrate localities within the territory of Hungary
and Romania. Sixteen localities included in this study are
situated on or adjacent to the former Pannonian Sea
margins and the resulting faunas belong to four
biostratigraphic units (MN6–MN9). Most of these sites with
snake rests were discovered in the last decade or so (Hír
et al., 1998; Gál et al., 1999, 2000; Hír, 2006; Venczel &
Hír, 2008; Hír & Kókay, 2009). The fossil remains
discussed in this paper are housed in the Municipal
Museum of Pásztó, Hungary (FF, FT, M, SH), the
Speleological Institute „Emil Racovi ǎ”, Bucharest,
Romania (TAU) and the ării Crişurilor Museum, Oradea,
Romania (SU, TAS, VA). Osteological nomenclature of
snakes and measurements of vertebrae follows Szyndlar
(1984).
Abbreviations used in the present paper: MMP:
Municipal Museum of Pásztó, Hungary; ISER:
Speleological Institute „Emil Racovi ǎ”, Bucharest,
Romania; MTC:
ării Crişurilor Museum, Oradea,
Romania; FF: Felsőtárkány-Felnémet (N-Hungary); FT:
Felsőtárkány (N-Hungary); M: Mátraszőlős (N-Hungary);
SH: Sámsonháza (N-Hungary); SU: Subpiatră (WRomania); TAS: Tăşad (W-Romania); TAU: Tau (WRomania); VA: Vârciorog (W-Romania); CL: centrum
length; CW: centrum width.
1
SYSTEMATIC PART
Suborder SERPENTES LINNAEUS, 1758
Infraorder SCOLECOPHIDIA DUMÉRIL & BIBRON, 1844
SCOLECOPHIDIA indet.
The available material consists of trunk vertebrae
(M2, M3, TAS, SU2/1, TAU, FF2/3, FT1, FT2, FT3/10),
preserving various parts of the vertebral centrum, neural
arch and synapophyses (Pl. I: Fig. 1). They collectively
document the typical morphological features of
scolecophidians, shared by the members of
Typhlopidae, Leptotyphlopidae, and Anomalepididae: the
neural arch is depressed dorso-ventrally and devoid of
neural spine; the centrum is flattened and free of haemal
keel; the subcentral foramina are present; the cotyle and
the condyle are depressed dorsoventrally; the
paracotylar foramina are lacking; the synapophyses are
undivided and have a hemispherical shape. Accordingly,
the material is not suitable for a more inclusive
assignment within scolecophidians. Presently our
continent is inhabited by the only European member of
Typhlopidae, the European worm snake (Typhlops
vermicularis) (Gasc et al., 1997).
Infraorder ALETHINOPHIDIA NOPCSA, 1923
Family BOIDAE GRAY, 1825
Albaneryx cf. volynicus was recorded from FT2,
based on one trunk vertebra and one caudal vertebra.
Both vertebrae are extremely small (Pl. I: Figs. 2, 3)
approaching in morphology those of Albaneryx
volynicus, described from the early late Miocene (MN 9)
of Gritsev (Zerova, 1989). The neural spine of the trunk
vertebra is somewhat longer than that of A. volynicus,
whereas its posterior margin is pointed similarly as in the
latter form (Zerova, 1989; Szyndlar, 1991a). The neural
spine of the caudal vertebra MMP 2009.608/2 is
distinctly higher and more widened posteriorly than that
of A. depereti (Hoffstetter & Rage, 1972), approaching
the condition seen in A. volynicus too.
Up to now, the fossil record of this genus included
ării Crişurilor Museum, B-dul Dacia 1-3, RO-410464 Oradea, Romania,e-mail: mvenczel@gmail.com
343
MÁRTON VENCZEL
Sansan (MN 6), La Grive M+L7 (MN 7), La Grive L3+L5
(MN 8), France, Kleineisenbach (Bavarian Molasse Basin),
Germany (Böhme & Ilg, 2003) and two late Miocene
localities from the territory of Eastern Europe: the late
Miocene (MN 9) of Gritsev (Albaneryx volynicus Zerova,
1989), and the late Miocene (MN 12) of Cherevichnoie
(Albaneryx sp.) (Szyndlar & Zerova, 1992).
Family COLUBRIDAE OPPEL, 1811
Within Colubridae the informal name “Natricinae” and
“Colubrinae”, used mostly by paleontologists as a
subfamilial rank, is applied in the present paper. All
Natricinae possess the hypapophysis in the precaudal
vertebrae, whereas in Colubrinae the hypapophysis of the
trunk vertebrae is reduced to a haemal keel.
“NATRICINAE”
Until now two fossil members of the genus Natrix were
listed from the studied localities: N. cf. rudabanyaensis
and N. sansanienis.
N. cf. rudabanyaensis was recorded from TAU, FF2/7,
FT2, FT3/2 and FT3/10 (Pl I: Figs: 4-9). The material
consists of a series of cranial bones and precaudal
vertebrae. The supraoccipital crest of the prootic (MMP
2009. 611/1 from FT2) is prominent; the foramen for the
maxillary ramus of the trigeminal nerve (V2) is relatively
small and of oval shape, and it is joined anteroventrally by
the foramen for the exit of the constrictor internus dorsalis
nerve (V4) (Pl. I: Fig. 4). In the parietal from M1, the dorsal
crests do not converge anteriorly the parietal posterior
margin (Pl. I: Fig. 5). The vertebrae of N. cf.
rudabanyaensis are of relatively small size (the CL of the
larger specimens reaches 4 mm) and moderately
elongated (CL/CW approaching 1.5). The relatively long
hypapophysis projects beyond the condyle and is rounded
distally. The vertebrae from the above localities approach
in morphology and size those from the type locality (Pl. I:
Figs. 6-9). In fact, this species was described from the
early Vallesian (MN 9) locality complex of Rudabánya (NHungary) on the basis of precaudal vertebrae only
(Szyndlar, 2005). Thus, the available cranial bones were
assigned with some doubt to this species. N.
rudabanyaensis was not recorded beyond the borders of
the Carpathian Basin and it may be considered an early
Vallesian invader from Asia.
N. sansanienis was briefly described from M1 by Gál et
al. (1999). The vertebrae are of various sizes with
relatively high neural spines and provided with very long
prezygapophyseal processes.
The remaining materials of Natricinae were assigned
either to Natrix sp. or to Natricinae indet. The presence of
Natricinae snakes, based on habitat preference of recent
forms,
indicates
the
existence
of
aquatic
paleoenvironments in the vicinity of the fossil deposits.
“COLUBRINAE”
Four genera were identified in the studied localities:
Coronella, Elaphe, Hierophis, and Texasophis.
The only remains assigned to Coronella miocaenica
are four trunk vertebrae recovered from TAU (Venczel &
Ştiucă, 2008). They closely resemble the type material
from the late Miocene (MN 13) of Polgárdi 4, Hungary
344
(Venczel, 1998), by their relatively small size, elongated
centrum, moderately flattened neural arch and by
extremely short prezygapophyseal processes (the
measurements of ISER Tt-0490/1: CL=3.45, CW=2.38,
CL/CW= 1.45) (Pl. I: Figs. 10-12). The neural spine of C.
miocaenica is moderately high (if compared to recent
Coronella), and it overhangs both anteriorly and
posteriorly without any thickening of the dorsal margin.
Texasophis cf. meini was described briefly by Gál et
al. (1999) from M1 (Pl. II: Figs. 1, 2). The available
material consists of five trunk vertebrae of relatively
small size (the CL of the largest one is 3.6 mm and it is
1.4 times longer than wide). The neural arch is
depressed and provided with a relatively long and low
neural spine. The zygosphene is narrow with straight or
crenate anterior margin, whereas the prezygapophyseal
processes are extremely short. The subcentral area is
concave ventrally and the haemal keel is prominent with
a flattened ventral surface. A taxon with a rather similar
morphology to Texasophis cf. meini (i.e. Telescopus
bolkayi), was described recently from the early Vallesian
(MN 9) locality of Rudabánya (Szyndlar, 2005).
Remains referred to Coluber (named as Coluber sp.)
were reported from the Astaracian (MN 6) site of SH by
Hír et al. (1998). Of the five vertebrae assigned to this
form, the largest one has a centrum somewhat longer
than wide (CL = 8.08 mm, CW = 7.62 mm, CL/CW =
1.06). The haemal keel is flattened and widened
posteriorly; in few of the assigned specimens it is steeply
angled behind the cotylar lip. The neural arch is
moderately vaulted, provided with a putatively high
neural spine; the anterior margin of the zygosphene is
straight; the paradiapophyses are distinctly differentiated
into parapophyseal and diapophyseal portions and are
roughly of equal length. More numerous and better
preserved material assignable to the above taxon was
recovered from the localities of M1, M2, M3, FT3/2 and
FT3/10; it consists of fragmentary basiparasphenoid,
quadrate, cervical and trunk vertebrae. In fact, the
vertebrae from the above localities are morphologically
comparable to those described under the name Coluber
pouchetii from the Astaracian (MN 6) of Sansan, France
(Augé & Rage 2000) and cf. Coluber pouchetii from the
early Vallesian (MN 9) of Rudabánya. However, part of
the material derived from the localities of M1 and M2
was already described under the name of Elaphe sp. by
Gál et al. (1999, 2000) (Pl. II: Figs. 3-5). After the latter
authors, the assignment was based mainly on the fact
that the hypapophyses of the cervical vertebrae have an
anteroventral orientation (Pl. II: Fig. 3), feature observed
only in some egg eating colubrids (e.g. Dasypeltis and
Elaphe). On the contrary, in members of Coluber such
morphological character was never observed (Szyndlar,
2005). Thus, the future taxonomic revision of the above
forms is considered necessary.
A single small sized vertebra (CL = 3.32, CW =
3.03) derived from FT1 may be assigned to Elaphe sp.
The centrum is short and wide (CL/CW = 1.09), whereas
the haemal keel is flattened and spatulate. The
zygosphene is crenate and the paradiapophyses are
markedly differentiated into parapophyseal- and
diapophyseal portions; the parapophyses are distinctly
longer than the diapophyses. All the above features are
typical for the members of Elaphe.
Hierophis cf. hungaricus was already described from
the late Middle Miocene (MN7+8) of Tau , Romania on
MIDDLE-LATE MIOCENE SNAKES FROM THE PANNONIAN BASIN
the basis of a quadrate and 35 trunk vertebrae (Venczel &
Ştiucă, 2008). Further material was recovered from FF2/3
and FF2/7 (Venczel & Hír, in prep.). The trunk vertebrae
from TAU are slightly larger (CL = 4.05-5.9, CW = 2.844.63, CL/CW = 1.23-1.45, n = 10) than those from FF2/3
and FF2/7 (CL = 3.02-3.1, CW = 2.07-2.1, CL/CW = 1.441.48, n = 4). The neural arch is moderately vaulted,
sometimes provided with small epizygapophyseal spines;
the zygosphene is convex dorsally with distinct crenate
anterior margin; the paradiapophyses are of minute size
with the parapophyseal and diapophyseal portions of
roughly the same length; the prezygapophyseal processes
are flattened dorsoventrally of variable length and usually
with an acute tip (Pl. II: Figs. 6, 7). The subcentral area
has a slightly concave surface, whereas the haemal keel is
flattened (more prominent in the posterior presacral
region). The material from M1, described briefly under the
name Coluber sp. by Gál et al. (1999), approaches in
morphology the remains from the above-mentioned
localities, probably representing the same taxon.
Several fragmentary vertebrae were assigned to
Colubrinae indet. or Colubridae indet., which probably may
have belonged to several small-sized colubrid genera.
Family ELAPIDAE BOIE, 1827
Micrurus gallicus was reported from SH. The material
consists of nine fragmentary presacral vertebrae with an
elongated centrum and with a somewhat depressed neural
arch. The hypapohysis is prominent and pointed distally; it
projects posteriorly below the condyle. The neural spine is
low and long, overhanging anteriorly and posteriorly. The
parapophyseal and diapophyseal portions of the
paradiapophyses are of equal length; the parapophyseal
processes are relatively short and directed forwards. The
anterior margin of the zygosphene is crenate with two
lateral lobes and with a medial larger one. Other elapid
material, consisting of few fragmentary vertebrae is known
from FT2 and FT3/2 (Venczel & Hír, in prep.).
All the above remains represent small-sized Elapidae.
The only large-sized elapid snake, genus Naja, is known
from the early Vallesian (MN 9) locality of Rudabánya
(Szyndlar, 2005).
name Macrovipera sp. from TAU (Venczel & Ştiucă,
2008). It is of relatively large size (CL = 8.18, CW = 6.64,
CL/CW = 1.23), provided with putatively high neural
spine and relatively long and straight hypapophysis (Pl.
II: Fig. 10). Morphologically the specimens approach
those described from the late Miocene (MN 11) of
Kohfidisch, Austria (Bachmayer & Szyndlar, 1985, 1987)
and Polgárdi (MN 13), Hungary (Venczel, 1994, 1998).
Vipera sp. (“V. aspis group”) was recorded from
FF2/3, FT3/10, M3 and SU2/1, the material consisting of
isolated and fragmentary presacral vertebrae. The neural
arch is depressed dorsoventrally; the neural spine in all
the specimens is broken off. The hypapophysis is
straight, long and posteroventrally oriented. The anterior
margin of the zygosphene is crenate; the
prezygapophyseal processes are extremely short, with
pointed tips. The paradiapophyses are differentiated into
diapo- and parapophyseal portions; the parapophyseal
processes with their tips obtuse distally project
anteroventrally. The centrum in a presacral vertebra
from SU2/1 is moderately elongated (CL = 4.24, CW =
2.92, CL/CW = 1.45) (Venczel, 2007), whereas that of a
larger specimen from M3 (Pl. II: Fig. 11) is comparatively
shorter (CL = 5.16, CW = 4.16, CL/CW = 1.24).
Vipera sp. (“V. berus group”) was recorded from FT1
(one vertebra), FT2 (one vertebra) and FT3/10 (three
vertebrae). All the specimens are fragmentary, but they
document collectively the most important morphological
features of the “V. berus group”. The centrum is
relatively long (about 1.66 times longer than wide in one
of the FT3/10 specimens), provided with a moderately
long hypapophysis projecting backward beyond the
condyle. The neural arch is depressed and the remnants
of the neural spine indicate that it was of low height and
overhanging posteriorly. Based on the above features,
the vertebrae may be assigned to the “Vipera berus
group” (Szyndlar, 1991b). In fact, the finds from the early
Vallesian (MN 9) of FT represent the oldest remains of
this group from the territory of Central Europe. Apart
from the FT localities, the oldest record of this group
comes from Gritsev (MN 9), Ukraine (Zerova, 1987;
Szyndlar, 1991b).
CONCLUDING REMARKS
Family VIPERIDAE OPPEL, 1811
Within vipers usually three groups or complexes are
distinguished based on the size and morphology of
presacral vertebrae: the “Oriental vipers group”, the
“Vipera aspis group” and the “Vipera berus group”
(Szyndlar, 2005).
A large-sized maxilla and few vertebrae, representing
the “Oriental vipers group” (Macrovipera), were discovered
recently from the Astaracian (MN 7+8) locality of VA1 (Pl.
II: Figs. 8, 9). The maxilla is distinctly larger when
compared to European vipers (e.g. to V. ammodytes); it
preserves two tooth positions (Pl. II: Figs. 9, 10). The
dorsal process is relatively high and curved medially. In
posterior view, the fossa for the ectopterygoid articulation
is relatively deep, delimited dorsally by a transversal crest.
The foramina for the dental canal are situated on the
medial side of the bone at the base of the dorsal process
(two foramina in a deep common fossa). A bony crest is
developed on the anterior side of the dorsal process. On
the lateral side of the latter, near its apex, a small foramen
is observed. A presacral vertebra was described under the
The Astaracian–early Vallesian snake faunas from the
Pannonian basin are represented by taxa from at least
five different groups (Scolecophidia, Boidae, Colubridae,
Elapidae, and Viperidae). The faunas exhibit a relatively
low local diversity (1-7 taxa/locality), determined by the
fact that all the studied localities were deposited in
marshy-lacustrine areas, yielding taxa with near-water
habitat preference.
The composition of snake faunas illustrates modern
patterns, dominated by colubrid snakes. A series of
widely-distributed taxa were distinguished (e.g. Coluber
pouchetii, Natrix sansaniensis, Texasophis cf. meini),
indicating important faunal links to Western Europe. The
assemblages from younger localities (MN 9) exhibit a
comparatively more diverse fauna, which points to an
increasing influence from Asia. The latter fact is marked by
the record of Albaneryx cf. volynicus, the appearance of new
Natricinae (N. rudabanyaensis) and Colubrinae (Coronella
miocaenica, Hierophis cf. hungaricus, Telescopus bolkay)
and by the first record of the “V. berus group”.
345
MÁRTON VENCZEL
Acknowledgement
The author is grateful to the anonymous reviewers who
provided helpful comments on the manuscript and
improved the English.
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MIDDLE-LATE MIOCENE SNAKES FROM THE PANNONIAN BASIN
PLATES
Plate I
Fig.1: SCOLECOPHIDIA indet.: trunk vertebra from FT2. Figs. 2, 3: Albaneryx cf. volynicus: trunk vertebra from FT2 (2);
caudal vertebra from FT2 (3). Figs. 4-9: Natrix cf. rudabanyaensis: prootic from FT2 (4); parietal from M1 (5);
presacral vertebra from M3 (6), presacral vertebra from FT2 (7) presacral vertebra from TAU (8, 9); Figs. 10-12:
Coronella miocaenica: trunk vertebra from TAU.
1, 2, 5, 9, 10 – dorsal views; 3 – anterior view; 4, 6-8, 11 – lateral views; 12 – ventral view. Scale = 1 mm (Figs. 1-4);
scale = 2 mm (Figs. 5-12).
Plate II
Figs. 1, 2: Texasophis cf. bohemiacus: trunk vertebra from M1. Figs. 3-5: Coluber pouchetii: fragmentary cervical
vertebra from M1 (3); trunk vertebra from M3 (4, 5). Figs. 6, 7: Hierophis cf. hungaricus: trunk vertebra from TAU.
Figs. 8-10: Macrovipera sp.: maxilla from VA (8, 9); presacral vertebra from TAU (10). Fig. 11: Vipera sp. (“Vipera
aspis group”): presacral vertebra from M3. Fig. 12: Vipera sp. (“Vipera berus group”): presacral vertebra from FT3/10.
1, 3, 10, 11, 12 – lateral views; 2, 5, 7 – ventral views; 4, 6 – dorsal views; 8 – posterior view; 9 – anterior view. Scale
= 1 mm (Figs. 1-3); scale = 2 mm (Figs. 4-12).
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MÁRTON VENCZEL
PLATE I
348
MIDDLE-LATE MIOCENE SNAKES FROM THE PANNONIAN BASIN
PLATE II
349