Available online at www.sciencedirect.com
C. R. Palevol 7 (2008) 383–390
Systematic Palaeontology (Vertebrate Palaeontology)
A new Miocene fauna of snakes from eastern Siberia, Russia.
Was the snake fauna largely homogenous
in Eurasia during the Miocene?
Jean-Claude Rage a,∗ , Igor G. Danilov b
a
UMR CNRS 5143, département ‘Histoire de la Terre’, Muséum national d’histoire naturelle, CP 38,
8, rue Buffon, 75231 Paris cedex 05, France
b Zoological Institute of the Russian Academy of Sciences, Universitetskaya Emb. 1,
Saint Petersburg 199034, Russia
Received 30 January 2008; accepted after revision 20 May 2008
Available online 3 July 2008
Presented by Philippe Taquet
Abstract
Togay, a locality from Ol’khon Island, Baykal Lake, Russia, has yielded a fauna of snakes of Late Middle or early Late Miocene
age. It is located in a broad area from which no Neogene snake has been reported; therefore, it represents an important landmark. The
fauna includes a non-erycine boid, two or three colubrids, a viperid of the ‘oriental complex’ of Vipera, and perhaps another viperid.
This assemblage is astonishingly reminiscent of the snake faunas from the late Early and early Middle Miocene from western and
central Europe, it being understood that Miocene faunas are practically unknown in the geographically intermediary area. It may be
entertained whether a homogenous snake fauna inhabited Eurasia (except the southern part of the continent) during the Miocene.
To cite this article: J.-C. Rage, I.G. Danilov, C. R. Palevol 7 (2008).
© 2008 Académie des sciences. Published by Elsevier Masson SAS. All rights reserved.
Résumé
Une nouvelle faune de serpents miocènes en Sibérie orientale, Russie. La faune de serpents eurasiatique était-elle largement
homogène au Miocène ? Togay, gisement de l’île d’Ol’khon (lac Baïkal, Russie), a produit des serpents du Miocène moyen tardif
ou supérieur précoce. Il se situe au sein d’une très vaste zone qui n’avait pas fourni de serpents néogènes ; il s’agit donc d’un repère
important. La faune comprend un Boidae non Erycinae, deux ou trois Colubridae, un Viperidae du groupe des ‘vipères orientales’
et, peut-être, un autre Viperidae. De façon étonnante, cette faune rappelle celles du Miocène inférieur tardif et moyen précoce
d’Europe occidentale et centrale, étant entendu que les faunes miocènes sont pratiquement inconnues dans la zone géographiquement
intermédiaire. Ce gisement suggère donc qu’une faune peut-être homogène occupait l’Eurasie, sauf sa partie méridionale, pendant
le Miocène. Pour citer cet article : J.-C. Rage, I.G. Danilov, C. R. Palevol 7 (2008).
© 2008 Académie des sciences. Published by Elsevier Masson SAS. All rights reserved.
Keywords: Snakes; Miocene; Boidae; Colubridae; Viperidae; Eastern Russia; Palaeobiogeography
Mots clés : Serpents ; Miocène ; Boidae ; Colubridae ; Viperidae ; Russie orientale ; Paléobiogéographie
∗
Corresponding author.
E-mail address: jcrage@mnhn.fr (J.-C. Rage).
1631-0683/$ – see front matter © 2008 Académie des sciences. Published by Elsevier Masson SAS. All rights reserved.
doi:10.1016/j.crpv.2008.05.004
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1. Introduction
In Eurasia, fossil snakes from the Neogene are well
documented from western and central Europe, but they
are very poorly known from more eastern regions. East
of westernmost Russia, and aside from Ol’khon Island,
central Mongolia, northeastern China and few localities in the Zaisan Basin (see below), Neogene snakes
were reported only from southern Eurasia: Anatolia
[13,20,31], Saudi Arabia [34], Siwalik beds of Pakistan
and India [5,18], Burma [24], Thailand [17], and Japan
[6].
This paper reports on a snake fauna from the Miocene
of the Togay locality, Ol’khon Island, Baykal Lake, Russia. Ivanjev and Khosatzky [7] briefly listed fossils from
the locality and assigned, without any comments, all
snake remains to Coluber. However, our study shows
that this fauna includes several taxa of snakes and represents a small but significant assemblage from the
late Middle or early Late Miocene. Togay is located
in the middle of a broad area from which no Neogene snake has been studied. Consequently, it is an
important landmark. The closest locality from which
Neogene snakes were reported is Builstyn Khudang
(Late Miocene, central Mongolia) (Fig. 1), which yielded
two erycines, two colubrines and one natricine [3]. Two
other Neogene localities of the area are more distant:
Zaisan Basin (Kazakhstan) and Ertemte (Inner Mongolia, northeastern China). Chkhikvadze [4] listed, but
did not describe snakes from the Zaisan Basin (see
below), whereas Schlosser [23] only reported an inde-
Fig. 1. Neogene localities bearing snakes in Eurasia (M: Miocene, P:
Pliocene). Grey area: part of Europe from which Neogene snakes are
well-documented.
Gisements d’Eurasie ayant fourni des serpents néogènes (M : Miocène,
P : Pliocène). Zone grisée : partie de l’Europe où les serpents néogènes
sont fréquents.
terminate snake from the latest Miocene (MN 13) of
Ertemte.
2. The locality
Togay is situated in the Togay Bay of Ol’khon Island,
Baykal Lake, Russia (Fig. 1 in [7]). Snakes come from
the lower part of the Khalagai Formation, whose age
is estimated as late Middle or early Late Miocene
[10,11].
3. Systematic account
The fauna is not rich, but it comprises four, perhaps
six species that belong to the Boidae, Colubridae s.l., and
Viperidae. The material was collected by N.A. Logachev
in 1958 and it is preserved in the Zoological Institute of
the Russian Academy of Sciences, palaeoherpetological
collection (ZIN PH), Saint Petersburg, Russia.
3.1. Boidae
Boinae
Indeterminate genus and species (Fig. 2)
Material. One trunk vertebra (ZIN PH 1/101).
Description
Only one trunk vertebra is referred to the Boidae.
It displays the characteristic vertebral morphology of
the family: vertebra short, wide and massive; prezygapophyseal processes small, not projecting laterally;
hypapophysis replaced by a haemal keel in the nonanterior part of the vertebral column. The presence of
well-marked subcentral grooves suggests that the vertebra likely comes from the posterior trunk region. Other
noticeable characters are the relatively vaulted neural arch, clearly inclined zygapophyses, well-developed
neural spine, slightly thickened dorsal border of the neural spine, and the presence of two paracotylar foramina
on either side of the cotyle.
Discussion
The Boidae are generally subdivided into three
groups: Boinae, Erycinae and Pythoninae; however, the
pythonines are sometimes regarded as a family distantly
related to the boine-erycine assemblage [35,37]. Irrespective of the precise relationships between these three
groups, it should be noted that the overall vertebral
morphology of the three taxa is very homogenous. Erycinae are distinguished from the two other groups on the
basis of the peculiar morphology of their caudal vertebrae; however, their trunk vertebrae may be useful here
because their neural arch is never vaulted. The latter
feature shows that the vertebra from Togay cannot be
J.-C. Rage, I.G. Danilov / C. R. Palevol 7 (2008) 383–390
385
Fig. 2. Boidae, indeterminate genus and species, posterior trunk vertebra (ZIN PH 1/101) in (A) dorsal, (B) lateral, (C) ventral, (D) anterior, and
(E) posterior views. Scale bar: 5 mm.
Boidae, genre et espèce indéterminés, vertèbre dorsale postérieure (ZIN PH 1/101) en vues dorsale (A), latérale (B), ventrale (C), antérieure (D)
et postérieure (E). Échelle: 5 mm.
referred to the Erycinae. The distinction between boines
and pythonines is difficult [22,32]. However, the presence of paracotylar foramina clearly argues against an
assignment to pythonines [32,33]. In addition, the lack
of a vertical ridge on the anterior face of the zygosphene
and the clearly inclined zygapophyses are consistent with
the boine morphology [22].
Extant Boinae occur in tropical Americas, Madagascar and islands of the West Pacific. Extinct Boinae were
reported from the Tertiary of Europe and the Americas [14,32]. The specimen from Togay differs from all
American boines from the Tertiary [14]; they are not
considered here. It clearly differs from the European
Palaeogene boines Palaeopython and Paleryx in having
paracotylar foramina, zygapophyses clearly inclined on
the horizontal, and the neural spine shorter anteroposteriorly. The overall morphology of the vertebra does
not markedly differ from that of Bavarioboa (Oligocene
and Miocene of Europe); more specifically, the presence of two paracotylar foramina on each side and the
thickened dorsal border of the neural spine are somewhat reminiscent of Bavarioboa hermi from the Early
Miocene of Europe [32]. However, the boine from Togay
is distinguished from Bavarioboa by its anteroposteriorly shorter neural spine and by the concave anterior
border of the zygosphene. Finally, although not markedly
different from Bavarioboa, the boine from Togay cannot
be referred to one of the known Boinae, but it cannot be
described as a new taxon because it is represented by a
single vertebra.
3.2. Colubridae s.l.
The Colubridae, as they have been long understood,
make up an enormous, non-monophyletic assemblage.
Despite this phylogenetic structure, morphological features are homogenous and the systematics of colubrids
based on such characters has been so far unsuccessful.
Many attempts at a subdivision of the traditional Colubridae into several monophyletic families have been
made, the most recent ones being based on molecular
data [38]. However, palaeontology is at a loss to distinguish such subgroups. In view of these difficulties, and
for convenience, palaeontologists retain the traditional
understanding of this assemblage that is here referred
to as ‘Colubridae s.l.’ On the basis of vertebrae, it is
only possible to clearly distinguish two morphological
groups, the ‘colubrines’ and the ‘natricines’, that is
an artificial subdivision. In ‘colubrines’, mid- and
posterior trunk vertebrae lack hypapophyses, whereas
in ‘natricines’, hypapophyses are present in the entire
trunk region. Togay has yielded only ‘colubrines’; two
or three species are present. It is worth mentioning that
among the extant colubrids, present in the area around
the Baykal Lake and potentially on Ol’khon Island, are
two species [1]: Elaphe dione (a ‘colubrine’) and Natrix
natrix (a ‘natricine’).
3.2.1. Coluber s.l.
As used in palaeontology, the genus name Coluber
does not correspond to the genus in its present taxo-
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J.-C. Rage, I.G. Danilov / C. R. Palevol 7 (2008) 383–390
Fig. 3. (A–C) Colubridae s.l., unnamed species A, mid-trunk vertebra (ZIN PH 2/101), anterior border of neural spine reconstructed from ZIN PH
3/101. (D–F) Viperidae,? Vipera (‘oriental complex’), mid-trunk vertebra (ZIN PH 34/101) (A, D: anterior views; B, E: ventral views; C, F: lateral
views). Each scale bar: 5 mm.
(A–C) Colubridae s.l., espèce A non nommée, vertèbre dorsale moyenne (ZIN PH 2/101), bord antérieur de la neurépine basé sur ZIN PH 3/101.
(D–F) Viperidae, ? Vipera « groupe des vipères orientales », vertèbre dorsale moyenne (ZIN PH 34/101) (A, D : faces antérieures ; B, E : faces
ventrales ; C, F : faces latérales). Échelles : 5 mm.
nomic meaning. Vertebrae of several living and extinct
colubrids show a homogenous morphology that renders
identification difficult or impossible at the genus level.
More specifically, in several living genera, the vertebral
morphology is similar to that of Coluber. Consequently,
extinct species belonging to this assemblage have been
referred symbolically to the genus Coluber s.l., it being
understood that the relationships between these snakes
remain unknown within this complex.
3.2.1.1. Unnamed species A (Fig. 3A–C).
M
. aterial. 25 trunk vertebrae (ZIN PH 2/101 to 26/101).
Description
This snake is a large colubrine; the length of the centrum ranges from above 7 to 4.9 mm. The vertebrae are
comparatively high and short. The neural arch is moderately vaulted; moreover, in posterior view, the posterior
borders of the neural arch are nearly straight to weakly
convex. Epizygapophyseal spines are present on most
vertebrae; apparently, their presence does not depend
on the size of the vertebrae. The neural spine is often
incomplete, but some remnants show that it was relatively high. The anterior border of the zygosphene is
approximately straight between the two lateral lobes;
however, in several vertebrae, a weakly convex median
lobe is present, the lobe being indented by a median
notch (which results in the presence of two lobes between
the lateral lobes: Fig. 3B). In most vertebrae, the cotyle
and condyle are slightly compressed laterally, which is a
somewhat unusual feature. The prezygapophyseal processes are well developed, strong, but short. Paracotylar
foramina are present. Astonishingly, the presence of two
foramina on each side represents the most frequent condition; moreover, foramina may be as numerous as four
on one side and three on the other side. The ventral face
of the centrum is triangular and longer than wide. It bears
a well-defined haemal keel that is spatulate, except in two
vertebrae in which it is gladiate. The para- and diapophyseal areas are distinct from each other. The diapophysis is
clearly shifted posteriorly with regard to the parapophysis. Two other features are worth noting. A sinuous,
blunt and vertical ridge occurs on either side, below the
postzygapophyseal facet; the smaller vertebrae lack it.
A marked, uneven tuberosity is present on each side of
most vertebrae, close to the lateral foramen and dorsal to
the paradiapophysis. Its size and shape are variable and
its presence does not appear to be connected to the size
of the vertebrae. Only six vertebrae lack it.
Discussion
This snake clearly belongs to the so-called ‘largesized colubrines’ [27,29]. This informal group includes
extinct species referred to the genera Coluber, Elaphe,
and Malpolon which are still extant. Among these large
species, the vertebral morphology of species A from
Togay most approaches that of Coluber pouchetii, C.
dolnicensis and C. caspioides. C. pouchetii was reported
from the Early (MN 4) and Middle (MN 6) Miocene of
France [16] and perhaps from the Late Miocene (MN
J.-C. Rage, I.G. Danilov / C. R. Palevol 7 (2008) 383–390
9) of Hungary [29]. C. dolnicensis is known only from
the Early Miocene (MN 3 and 4) of the Czech Republic [9,25]. C. caspioides was recovered from the Early
Miocene of the Czech Republic (MN 3 [9]), Germany
(MN 4 [33]) and Austria (MN 4 [29]), and from the
Middle Miocene (MN7 + 8) of France [29]. In addition,
a younger colubrine from the Pliocene of the Balearic
Islands belongs to this assemblage, but it cannot be identified at the species level [2]. Aside from the large size,
four characters are common to C. pouchetii, C. dolnicensis, C. caspioides, and species A from Togay: (1)
neural arch weakly depressed to moderately vaulted, (2)
posterior borders of the neural arch nearly straight to
weakly convex in posterior view, (3) neural spine not low,
(4) anterior border of the zygosphene generally indented
by a median notch, the border being otherwise straight,
weakly convex or weakly concave between the lateral
lobes.
Within this assemblage, species A compares more
favourably with Coluber pouchetii and C. dolnicensis.
In these three snakes, the diapophysis is shifted posteriorly and the cotyle and condyle of various vertebrae
are slightly compressed laterally. However, aside from
the presence of lateral tuberosities whose significance is
unknown, two characters distinguish species A from C.
pouchetii and C. dolnicensis. In at least a part of the vertebral column of the latter two species, the haemal keel
forms a step in the anterior portion of the centrum, while
species A lacks such a step. Moreover, both C. pouchetii
and C. dolnicensis have a single paracotylar foramen on
either side of the cotyle, which is the condition in nearly
all colubrids. Most vertebrae of species A have at least
two foramina on each side, and four foramina are even
present on one side in two vertebrae; to our knowledge,
so numerous paracotylar foramina were not reported
previously in snakes (except in the Cretaceous–Eocene
Palaeophiidae). Finally, species A from Togay appears
to be related to the ‘large-sized colubrine’ assemblage
previously reported from the Neogene of Europe, but it
probably represents a distinct, new species. The description of this new taxon is not our present objective.
3.2.1.2. ? Unnamed species B.
M
. aterial. Five trunk vertebrae (ZIN PH 27/101 to
31/101).
Description
The vertebrae are somewhat similar to those of
species A, but they are smaller (length of centrum: 5.1
to 4.2 mm). Aside from its size, this snake differs from
species A in having more vaulted neural arches and
convex posterior borders of neural arches in posterior
view. Moreover, the cotyles and condyles are not com-
387
pressed laterally. In addition, ‘colubrine B’ lacks the
lateral tuberosities and the subvertical ridges located
below the postzygapophyses. Epizygapophyseal spines
may be present or absent. At least two paracotylar foramina are present on each side, but three foramina can occur.
Discussion
Several of the characters that distinguish specimens
allocated to ‘colubrine B’ from those referred to species
A are perhaps not significant. The absence of lateral
compression of the cotyles and condyles also occurs
in various vertebrae of species A and the small number of specimens belonging to ‘colubrine B’ precludes
any conclusion. The absence of tuberosities and posterior subvertical ridges may be size related, although
the tuberosities are lacking in some large vertebrae, but
are present in some small vertebrae of species A. On the
other hand, the more vaulted neural arches and more convex posterior borders of the neural arches do not appear
to result from the smaller size. On the contrary, the posterior borders of the neural arches tend to be more convex
in large individuals than in small ones. From this, it might
be inferred that ‘colubrine B’ is distinct from species A.
However, the presence of at least two paracotylar foramina on each side – a very peculiar character – is common
to these two snakes and argues for the referral of ‘colubrine B’ to species A. Unfortunately, the limited sample
does not permit us to establish whether ‘colubrine B’ is
distinct from species A.
3.2.1.3. Indeterminate genus and species.
M
. aterial. Two trunk vertebrae (ZIN PH 32/101 and
33/101).
Description and discussion
Two vertebrae of a small colubrine display a morphology clearly distinct from that of the Coluber assemblage.
The centrum length of the largest vertebra is 3.9 mm. The
vertebrae are elongate and depressed. The neural arch
is strongly depressed and the neural spine is low and
long. The zygosphene is wide (wider than the slightly
depressed cotyle), thin and provided with a wide median
lobe. One paracotylar foramen opens on either side of
the cotyle. The paradiapophyses are comparatively massive and they lack a constriction between the para- and
diapophyseal areas.
These vertebrae are reminiscent of the extant Eurasian
genus Coronella, the earliest species of which is reported
from the Latest Miocene (MN 13) of Europe [36]. However, the vertebrae from Togay differ from those of
Coronella in being more elongate and in having centra
growing narrower posteriorly. On the other hand, small
colubrines are so numerous and their osteology is so
poorly known that it cannot be excluded (? it is proba-
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J.-C. Rage, I.G. Danilov / C. R. Palevol 7 (2008) 383–390
ble) that the vertebral morphology of the fossil occurs
among unstudied living forms.
3.3. Viperidae
? Viperinae.
3.3.1. ? Vipera (‘oriental complex’)
Indeterminate species (Fig. 3D–F).
half of Europe from the late Early Miocene (MN 4) to
the Latest Pliocene (MN 16) and in southwesternmost
Asia (Anatolia) during the Late Miocene [31]. In Africa,
it perhaps reached the South of the continent during the
Early Miocene [15]. Daboia occurs in south Asia and it
was reported from the Early Pliocene [26] and perhaps
the Early Miocene [30] of western Europe and southern
Africa [15]. Protobothrops flavoviridis is restricted to
Ryukyu Islands (Japan). An Early Miocene fossil from
Honshu Island (Japan) would be close to this species [6].
Material. One trunk vertebra (ZIN PH 34/101).
Description. This relatively large vertebra (centrum
length: 5.2 mm) shows a combination of characters that
is typical of viperids: vertebra depressed, short and wide;
neural arch depressed; hypapophysis present. Other features that are worth mentioning are the marked slanting
of the prezygapophyseal facets, the small size of the
prezygapophyseal processes that do not strongly project
laterally, and the probable presence of epizygapophyseal
spines. Unfortunately, the hypapophysis, neural spine
and parapophyseal processes are broken away.
Discussion
The marked shortness of the vertebra, the strongly
depressed condition of the neural arch, and the relatively
large size point to the ‘oriental complex’ of species of
Vipera. This informal assemblage includes living and
extinct species, whose precise relationships are debated,
but whose osteology is homogenous [30,31].
However, the vertebrae of the extant viperine Daboia
[30] and crotaline Protobothrops (formerly Trimeresurus) flavoviridis [28] are somewhat similar to those of
the ‘oriental vipers’. Unfortunately, the main difference
between the ‘oriental vipers’ and Daboia is the markedly
higher neural spine of the latter. This character cannot
be used, since the neural spine is broken off in the fossil. On the other hand, the vertebrae of Daboia appear
to be higher and narrower than that from Togay; consequently, assignment of the fossil to this genus seems
unlikely. P. flavoviridis may be discarded on the basis
of its clearly larger cotyle and condyle, markedly more
extended zygapophyseal facets, and perhaps (as far as
may be inferred from the fossil) less anteriorly directed
parapophyses. Nevertheless, taking into account the fact
that the ‘oriental vipers’ assemblage is not well defined,
and that the feature that clearly distinguishes this assemblage from Daboia is not preserved, the referral of the
fossil from Togay to the ‘oriental complex’ of Vipera
cannot be accepted definitely.
Today, ‘oriental vipers’ inhabit southwestern Asia,
the Middle East, southeasternmost Europe, and northernmost Africa. The group was present in the southern
3.3.2. Indeterminate Viperidae.
Material. One trunk vertebra (ZIN PH 35/101).
Description and discussion
This specimen is only poorly mineralized or not mineralized at all. It is probably not a fossil, but this cannot be
demonstrated. It displays the morphology of the species
belonging to the ‘berus complex’ of Vipera [30,31], but
the osteology of the small Asian crotalines (Gloydius)
being practically unknown, this specimen is regarded a
viperid indeterminate. It should be noted that the living
Vipera berus and Gloydius halys occur in the area of
Baykal Lake and that their presence on Ol’khon Island
seems very probable [1].
4. Palaeobiogeographical relationships
Although neither rich nor very diverse, this fauna
provides palaeobiogeographic information.
4.1. Faunal affinities with Europe
Astonishingly, the overall composition of the fauna
from Togay recalls that of the late Early Miocene and
early Middle Miocene (MN 3–MN 5) of Europe: presence of a rather large non-erycine boid, of colubrids
including a large colubrine, and of a large viperid (the
latter lacking from MN 3). In Europe, the boid may be
either the pythonine Python or the boine Bavarioboa.
The European colubrids are diverse, but they always
include at least one large colubrine that may belong
to Coluber or Elaphe. The large viperid from the MN
4–MN 5 period in Europe belongs to the ‘oriental complex’ of Vipera or to Daboia. Before MN 3, the Miocene
of Europe lacks non-erycine large boids and large vipers
[8,21,32]. After MN5, non-erycine boids were definitely
ousted from Europe by colubroids.
4.2. The Asian faunas
In Asia, only six localities, or complexes of localities, previously produced faunas (i.e. more than one
J.-C. Rage, I.G. Danilov / C. R. Palevol 7 (2008) 383–390
species) of Neogene snakes: the Siwalik beds (Pakistan
and India), Builstyn Khudang (Mongolia), Li Mae Long
(Thailand), Zaisan Basin (Kazakhstan), Al Sarrar (Saudi
Arabia), all Miocene in age, and Çalta (Anatolia), which
is Pliocene. The Siwalik beds [5,18] and Li Mae Long
[17] correspond to aquatic environments and they cannot
be significantly compared to Togay and to the European
sites.
The terrestrial snakes from the Miocene of Al Sarrar and Zaisan Basin are poorly known. The overall
composition of the fauna from Al Sarrar (late Early
Miocene [34]) is similar to that of Europe and Togay, but
the boids, colubrids and viperids are small, which may
result from taphonomic conditions. As the fauna from
Togay, the Late Miocene of Builstyn Khudang includes
large colubrines, but it differs from it by the presence of
two erycines and a natricine and in lacking non-erycine
boids [3]. The localities of the Zaisan Basin collectively
yielded poor faunas that extend from the Early to the
Late Miocene [4]. Only the presence of a large viperid is
reminiscent of Europe and Togay. The fauna of Çalta
is the only assemblage of snakes described from the
Pliocene of Asia [20]; it mainly differs from Togay in
lacking non-erycine boids and viperids and in having an
elapid.
4.3. Was the Eurasian fauna homogenous during
the Miocene?
Maridet et al. [12] showed that the European fauna
of mammals was homogenous during the Miocene up
to zone MN 4. During the interval MN 5–MN 7 + 8, the
homogeneity decreased and it reached its lowest level
during MN 11 (middle Late Miocene) (Fig. 9 in [12]).
In addition, faunal dissimilarity mainly corresponds to a
north–south gradient. Unfortunately, this study was not
extended to Asia. As far as snakes are concerned, the
similarity between the faunas from Togay and Europe
raises the question of a possible faunal homogeneity over
a large part of Eurasia (southern Eurasia excluded) during the Miocene. Obviously, the earliest possible age
of Togay appears to be late Middle Miocene, while the
‘similar’ period in Europe ends by the early Middle
Miocene, but the fauna from Togay may be regarded
as a relict of a formerly homogenous Eurasian fauna.
The only intervening faunas (Zaisan Basin) neither go
against nor support this possibility. The hypothesis of
the faunal homogeneity in Eurasia during the Neogene
is indirectly supported by the fact that various snakes
(and mammals) from the Miocene of Europe and North
America are related, Asia appearing as the intervening
area for dispersals [19].
389
Acknowledgements
This study was made possible by the financial support
of a grant of the President of the Russian Federation to
the Leading Scientific Schools (NSh-119.2008.4) and
a grant of the Russian Foundation for Basic Research
(08-04-00041-A). F. de Lapparent (MNHN, Paris) and
M. Venczel (Muzeul Tarii Crisurilor, Oradea) provided
useful comments.
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