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© 2013 International Society of Zoological Sciences, Institute of Zoology/
Chinese Academy of Sciences and Wiley Publishing Asia Pty Ltd
Integrative Zoology 2014; 9: 517–530 doi: 10.1111/1749-4877.12082
ORIGINAL ARTICLE
First Asian record of Panthera (Leo) fossilis (Mammalia,
Carnivora, Felidae) in the Early Pleistocene of Western Siberia,
Russia
Marina V. SOTNIKOVA1 and Irina V. FORONOVA2
1Geological Institute of Russian Academy of Sciences, Moscow, Russia and 2V.S. Sobolev Institute of Geology and Mineralogy,
Siberian Branch of Russian Academy of Sciences, Novosibirsk, Russia
Abstract
A lion-like pantherine felid is described as Panthera (Leo) fossilis from the late Early Pleistocene sediments of
the Kuznetsk Basin (Western Siberia, Russia). The nd of P. fossilis rst recorded in Asia considerably extends
the current notion of the eastward expansion of the most ancient lions. The Siberian lion is geologically the old-
est form and is dimensionally among the largest members of the group of fossil lions on the Eurasian conti-
nent. Although known by mandibular remains only, it is readily distinguished from Panthera (Leo) spelaea by a
heavy built mandibular corpus with rectangular prole in the cheek teeth area, a deep, well-outlined and narrow
anterior section of the masseteric fossa, and a large р4 supported by a big unreduced anterior root. The Siberian
lion shares these features with the European Middle Pleistocene P. fossilis and the American Late Pleistocene P.
(Leo) atrox, which suggests their close relationship. P. atrox originated from P. fossilis and was isolated in North
America south of the Late Pleistocene ice sheets. This explains why the American lion has retained more primi-
tive features than the coeval Eurasian cave lion P. (L.) spelaea.
Key words: Early Pleistocene, evolution, Panthera fossilis, relationships, Western Siberia
Correspondence: Marina V. Sotnikova, Geological Institute of
Russian Academy of Sciences, Pyzhewsky 7, 119017 Moscow,
Russia.
Email: sotnikmarina@yandex.ru
INTRODUCTION
Fossil members of the Felidae lion group Panthera
(Leo) spp. have a complex history, with numerous spe-
cies and subspecies described. Lions originated in Afri-
ca and dispersed northward into Europe during the early
Middle Pleistocene, 0.75–0.68 Ma (Hemmer 2011). In
the Late Pleistocene, this group of pantherine cats was
widespread in the Holarctic, ranging from Africa and
across Europe and Asia to America (Vereshchagin 1971;
Hemmer 1974; Kurtén 1985; Turner & Antón 1997; Ya-
maguchi et al. 2004; Barnett et al. 2009).
The Middle Pleistocene distribution of the oldest
member of this group of Felidae, Panthera (Leo) foss-
ilis (von Reichenau, 1906), has long been known, but
only in Europe (von Reichenau 1906; Freudenberg
1914; Dietrich 1968; Kurtén 1968; Schütt 1969; Hem-
mer 1974, 2011; Kurtén & Poulianos 1977; Schütt &
Hemmer 1978; Sala 1990; Wolsan 1993; Argant et al.
2007; Cuenca-Bescós & García 2007; Hankó & Korsós
2007; Barycka 2008; Marciszak & Stefaniak 2010; Sab-
ol 2011a).
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M. V. Sotnikova and I. V. Foronova
© 2013 International Society of Zoological Sciences, Institute of Zoology/
Chinese Academy of Sciences and Wiley Publishing Asia Pty Ltd
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Fossil evidence documenting the occurrence of
early lions in the Middle Pleistocene of Asia is very
poor. Outside Asian Russia, the only large Panthera
(Leo) youngi (Pei, 1934) from the Peking Man site of
Zhoukoudian 1, northeastern China, dated at 0.69–0.42
Ma (Qiu 2006), was referred to the lion group of pan-
therine cats (Pei 1934; Harington 1969).
Numerous finds of fossil lions in Asian Russia are
known from the papers by Riabinin (1919), Gromo-
va (1932), Vangengeim (1961), Vereshchagin (1971),
Alexeeva (1980), Foronova (1982, 1999, 2001), Barysh-
nikov and Boeskorov (2001), Sotnikova and Nikolskiy
(2006), Baryshnikov and Petrova (2008) and Ovodov
and Tarasov (2009). However, the material presented in
these works mainly belongs to the Late Pleistocene Pan-
thera (Leo) spelaea (Goldfuss, 1810). Middle Pleisto-
cene large lion-like felids in Russia have been reported
occasionally but these nds have been associated neither
with the European P. fossilis nor with Asian P. youngi
(Vereshchagin 1971). The assumption of the presence of
P. fossilis in the Asian part of Russia was rst made by
Baryshnikov and Boeskorov (2001) based on finds of
very large lions’ limb bones in the Middle Pleistocene
localities along the Volga and Lower Tunguska Rivers.
However, the rst clear evidence of a giant lion similar
to the P. fossilis–P. youngi group in Asia is a mandible
described by Foronova (1998, 2001, 2005) as Panthera
sp. from the Early Pleistocene of the Kuznetsk Basin
in Western Siberia. The mandibular size of this felid is
close to that of the largest forms of fossil pantherine cats
and in some parameters (Lp3-m1) even exceeds them.
Consequently, the Siberian form is geologically the old-
est and metrically the largest member of the fossil lions
in Eurasia.
The objective of the present paper is to provide a de-
tailed description and comparison of the Siberian spec-
imen with the similar-sized taxa. This study will also
help to expand the existing knowledge of the morphol-
ogy and the stratigraphic and geographic distribution of
the oldest members of the lion group in Asia, as well as
to clarify their relationship with European P. fossilis and
American P. (Leo) atrox (Leidy, 1853) distributed south
of the Late Pleistocene ice sheets.
MATERIALS AND METHODS
This study is based on a detailed morphological
analysis of mandibular and dental features of the pan-
therine felid from Kuznetsk Basin and large Pleistocene
lion-like cats of Europe and North America. For com-
parison we used collections of P. spelaea from the V.S.
Sobolev Institute of Geology and Mineralogy, Siberi-
an Branch of Russian Academy of Sciences, Novosi-
birsk (IGM SB RAS), the Geological Institute, Russian
Academy of Science, Moscow (GIN RAS) and the Zo-
ological Institute, Russian Academy of Science, St. Pe-
tersburg (ZIN RAS). Additional morphological and di-
mensional information was obtained from the literature.
In this study, we used a system of standard measure-
ments of lower jaws and teeth adopted by many authors
(Merriam & Stock 1932; Schütt 1969; Vereshchagin
1971; Baryshnikov & Boeskorov 2001).
The analysis shows that the terms used to describe
the premolar cusps of Felidae do not match in various
authors. For instance, the posterior additional cusp on
р3–4 is referred to as metaconid in Barycka (2008, p.
28, g. 6), whereas other paleontologists call this cusp
hypoconid (Hankó & Korsós 2007, g. 1; Sabol 2011b).
Therefore, we accept the system of dental elements used
by US researchers, in particular by Merriam and Stock
(1932), to describe non-carnassial teeth of felids, where
the premolar cusps are designated as anterior, posterior,
cingular and principal (or main) premolar cusps.
The dental information of the mandible from the
Kuznetsk Basin was obtained only on the basis of alve-
olar measurements. Considering the fact that in P. spe-
laea the alveolar length of cheek teeth is very close to
the dental length (with an error no more than 1.0–1.5
mm), we believed it possible to compare the alveolar
length of our nding with the teeth parameters of other
fossil lion-like pantherine felids.
The studied material is housed at IGM SB RAS, No-
vosibirsk. All measurements are in millimeters.
RESULTS
Remarks on the geological setting of Siberian
Panthera
Hemmer (2011), referring to Foronova (1998), ques-
tions the correctness of the stratigraphic position of the
mandible of a large pantherine cat derived from the
Sagarlyk Formation in the Bachatsk Quarry of Kuznetsk
Basin. In this regard, he considers the geological evi-
dence for the occurrence of a large fossil lion in the Si-
berian Early Pleistocene to be unreliable. The original
paper (Foronova 1998, p. 358) includes brief informa-
tion on 2 fragments of Panthera sp. from the Kuznetsk
Basin, that is, a very large mandible IGM-519 from the
Sagarlyk Formation (Fm) of the Bachatsk Quarry, which
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Early Pleistocene Panthera from Siberia
© 2013 International Society of Zoological Sciences, Institute of Zoology/
Chinese Academy of Sciences and Wiley Publishing Asia Pty Ltd
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we describe in this article, and a maxilla (No. 3608)
found in deposits of the younger Sergeevo Fm in the
Mokhovo Quarry. In the caption to gure 5 (p. 359) the
analyzed mandible IGM-519 from the Bachatsk Quar-
ry is incorrectly numbered 3608. This number belongs
to the maxilla from the Mokhovo Quarry, but its image
is missing in gure 5. In addition, in gure 3 (p. 357),
which shows numerous fossiliferous sections from dif-
ferent quarries, the section of the Bachatsk Quarry that
contained mandible IGM-519 is absent. The inaccura-
cies made in this publication have created the impres-
sion that the mandible from the Bachatsk Quarry was
found in an obscure geological position. We hope that
the following information on the geological and strati-
graphic position of the material described and on the
associated fauna completely eliminates the misunder-
standing.
The Kuznetsk Basin is the largest intermontane de-
pression located in the south-eastern part of Western Si-
beria (Fig. 1). In the Late Cenozoic an almost contin-
uous sequence of Quaternary sediments, over 150 m
thick, ranging from the beginning of the Early Pleis-
tocene to the Holocene, was accumulated in this area.
These alternating deluvial–proluvial and subaqueous se-
quences are recovered in large open coal pits, which
provide comprehensive biostratigraphic characteristics
of the Quaternary sediments of the region. The Early
Pleistocene sediments are represented upward from the
base by deposits of the Mokhovo, Sagarlyk, Sergeevo
and the lowermost part of the Kedrovka Fm with fair-
ly complete paleontological and magnetostratigraphic
characteristics. On the basis of these data in the compos-
ite stratotype section of the region, the major chronos-
tratigraphic boundaries at 0.8 and 1.8 Ma are recognized
and reliably characterized, and the paleomagnetic mark-
er events (i.e. the Brunhes/Matuyama boundary and Ja-
ramillo Subchron) are recorded (Foronova 1998, 1999,
2001, 2005) (Fig. 2).
The Panthera mandible IGM-519 was found in the
Bachatsk Quarry, in the section represented by sedi-
ments of 2 extremely diachronous and lithologically-dif-
ferent subaqueous formations. Very compact gray–green
sandy clays of Sagarlyk Fm are overlain with a signi-
cant hiatus by the lacustrine–alluvial gray–blue plastic
clays of the Kedrovka Fm. The subaerial deposits of the
Sergeevo Fm separating these subaqueous sequences in
the complete sections are missing in this outcrop.
The fossil lion find was revealed by one of the au-
thors (IF) in an open part of a large bone-bearing lay-
er in the upper part of the Sagarlyk Fm. Bones and teeth
of Equus aff. suessenbornensis Wüst, 1901 and Equus
sanmeniensis Teilhard de Chardin and Piveteau, 1930
were also found in the same layer, as well as a molar of
Archidiskodon sp. similar to Archidiskodon tokunagai
(Matsumoto) Teilhard and Trassaert, 1937 from the Ear-
Figure 1 Location of the Kuznetsk Basin (A) and Kurtak ar-
cheological area (B).
Figure 2 Stratigraphy of the Early–Middle Quaternary deposits
of the Kuznetsk Basin and position of Panthera fossilis (IGM-
519).
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M. V. Sotnikova and I. V. Foronova
© 2013 International Society of Zoological Sciences, Institute of Zoology/
Chinese Academy of Sciences and Wiley Publishing Asia Pty Ltd
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ly Pleistocene of China. Previously, this peculiar an-
cient elephant was attributed by various paleontologists
either to Archidiskodon or Palaeoloxodon, but present-
ly Chinese and Japanese scientists (Wei et al. 2006) be-
lieve that it belongs to Mammuthus meridionalis (Nesti,
1825) (=Archidiskodon meridionalis according to taxon-
omy accepted in Russia).
All fossil bones from Sagarlyk Fm are very heavy
and strongly mineralized. The described mandible of
dark brown color is massive and heavy; its hollows and
cavities were also lled with dense cemented and ferru-
ginized enclosing deposits. According to these fossiliza-
tion features, it does not differ from other bones in the
Sagarlyk Fm. The biostratigraphic characteristics of this
part of the formation are supplemented with ndings in
other sections of Allophaiomys pliocaenicus Kormos,
1932, Prolagurus pannonicus Kormos, 1930, Archidis-
kodon meridionalis tamanensis Dubrovo, 1964, E. aff.
suessenbornensis, E. sanmeniensis, very large Alces aff.
latifrons (Johnson, 1874) and Bison sp. (ex gr. priscus
Bojanus, 1827).
The sediments of the upper part of the Sagarlyk Fm
bearing fossils including the described Panthera mandi-
ble, teeth of meridionaloid elephants and bones of other
mentioned forms are normally magnetized and are cor-
related with the Jaramillo Subchron of the Matuyama
Chron (Fig. 2). Consequently, the evolutionary stage
of major fossils (i.e. rodents, meridionaloid elephants,
horses and broad-fronted moose), along with the paleo-
magnetic record, permit the correlation of the Sagarlyk
fauna with the Tamanian assemblage of European Rus-
sia and with faunas of the Early Galerian of Western Eu-
rope (Foronova 1998, 1999, 2001, 2005).
The overlying deposits of the Kedrovka Fm in its
complete volume have a rather wide age range, and in
the most part are correlated with the Brunhes Chron.
The rich paleontological characteristics of this forma-
tion permit its subdivision into heterochronous fossilif-
erous layers. However, in the studied section the Sagar-
lyk Fm is overlain by only basal layers of the Kedrovka
Fm, which are reversely magnetized, and, consequently,
the accumulation of the formation started in the termi-
nal Matuyama Chron (Fig. 2). This part of the formation
contains significantly younger fauna compared to the
Sagarlyk Fm; namely, Mammuthus trogontherii, Equus
mosbachensis, Rangifer sp., Bos sp. and large ‘priscoid’
Bison.
As for fossilization, the bones of Kedrovka Fm differ
substantially from Sagarlyk bones, having lower degree
of mineralization and lighter color. This faunal assem-
blage is correlated with Tiraspolian and Viatkian faunas
of Russia, as well as with the Cromerian (excluding the
earliest one) faunas of Western Europe.
Thus, considering all available geological, paleonto-
logical and paleomagnetic data, it can be concluded that
the age of the mandible described corresponds to the
time of accumulation of the upper part of the Sagarlyk
Fm; namely, to the Jaramillo Subchron of the uppermost
Matuyama Chron (i.e. to the late Early Pleistocene).
Systematic paleontology
Order Carnivora Bowdich, 1821
Family Felidae Fischer, 1817
Subfamily Pantherinae Pocock, 1917
Genus Panthera Oken, 1816
Subgenus Leo Brehm, 1829
Panthera (Leo) fossilis (von Reichenau, 1906)
Panthera sp.: Foronova 1998: 358–359 part, figure
5(1) – No. 519; Foronova 1999: 73,77 part; Foronova
2001: 69–70 part, 196-197: table IV, gure 2; Foronova
2005: 97, gure 3. Panthera leo fossilis: Hemmer 2011,
p. 203.
Material: Horizontal ramus of the left mandible
(IGM-519) with the canine and p3 alveoli; with 2 roots
exposed inside the alveolus of p4 and a fractured crown
of m1. The anterior part of the symphyseal region is
partly destroyed; coronoid, condylar and angular pro-
cesses are missing (Fig. 3a–c).
Locality: Southeast of Western Siberia, Kuznetsk Ba-
sin, Bachatsk Quarry (geographic position: 54.28°N,
86.16°E), upper part of the Sagarlyk Fm.
Age: Late Early Pleistocene.
Taxonomic remarks
The lion affinities of large pantherine felids from
the Pleistocene of Eurasia and North America had been
clarified by numerous morphological studies and sup-
ported by the molecular data of Burger et al. (2004) and
Barnett et al. (2009).
Consequently, following the concept of Vereshchagin
(1971) we grade the lion group of pantherine cats up to
the rank of their own subgenus Leo Brehm, 1829. With-
in this subgenus, according to the latest morphologi-
cal and molecular record (Sotnikova & Nikolskiy 2006;
Barnett et al. 2009), we include 4 taxa; that is, the living
Panthera leo (Linnaeus, 1758) (Africa and South Asia)
and Pleistocene P. (L.) spelaea (Eurasia and Beringia),
P. (L.) fossilis (Eurasia), and P. (L.) atrox distributed in
North America south of the Late Pleistocene ice sheet.
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Early Pleistocene Panthera from Siberia
© 2013 International Society of Zoological Sciences, Institute of Zoology/
Chinese Academy of Sciences and Wiley Publishing Asia Pty Ltd
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Description
The fragment of mandible IGM-519 has a strong and
unusually massive horizontal body, with the greatest
thickness below p4 (thickness [T] = 32.0 mm) and with
an equal-in-size depth in front of p3 and behind m1. An
almost straight mandibular ventral margin below p3-m1
is also characteristic of this specimen.
In buccal view a deep and distinctly outlined masse-
teric fossa reaches forward to the level of the space be-
tween 2 roots of m1. The anterior part of the masseteric
fossa is dorso–ventrally narrower than in P. spelaea and
P. leo. Its ventral margin only slightly crosses the mid-
line of the mandibular body in a dorso–ventral direction.
The basin between the anterior border of the masseter-
ic fossa and its deepest part, which is usually shallow in
mandibles of P. spelaea and P. leo, is very deep in the
Siberian specimen. The anterior mental foramen is bro-
ken, and the posterior one is doubled and is situated be-
low the anterior root of p3 and the middle of the c-p3
diastema. The greatest diameter of the doubled foramen
is 15.5 mm, whereas each of the foramens has the great-
est and smallest diameters of 7.1/3.5 mm and 11.0/7.7
mm, respectively. Like the ventral margin of the anteri-
or part of the masseteric fossa, the posterior mental fo-
ramen is located near the horizontal midline of the man-
dibular body (Fig. 3a).
In lingual view the posterior border of the symphyse-
al surface is nearly vertically oriented. Its most posteri-
orly projecting part does not reach the level of the mid-
dle part of the diastema between canine and p3 (Fig.
3b).
In occlusal view, the space for incisors is moderate-
ly wide; the postcanine diastema is long, with length
(L) of approximately 28.5 mm. According to the alveo-
lar structure, the lower canine had a large size and a ver-
tically-oriented root. Its estimated length and width are
approximately 31.0 and 21.0 mm. Judging by the alveo-
li, the cheek-tooth series slightly arches buccally. In the
tooth row, premolars occupy a clearly more labial posi-
tion in relation to m1, so the lingual margin of p4 roots
is displaced at 6.7 mm buccally relative to the lingual
border of the m1 root. There is also a relatively long di-
astema (L = 3.7 mm) between p4 and m1 roots. There
is no any overlap or crowding of the cheek teeth in the
mandible of Siberian specimen. The alveolar length of
p3-m1 is very large (Fig. 3c, Table 1).
The anterior root of p3 (root length directly below
a tooth crown [Lr] = 10.0 mm) is smaller than the pos-
terior root (Lr = 12.2 mm), whereas the roots of p4 are
nearly equal in size, with Lr/Lr = 13.1 and 13.2 mm, re-
spectively. The m1 crown is supported by a large anteri-
or root with Lr/root width directly below a tooth crown
(Wr) = 20.0/13.5 mm and a smaller posterior root with
Lr/Wr = 10.0/9.0 mm. The lingual protrusion of the in-
ner part of the posterior root of m1 beyond the lingual
margin of anterior root is also seen on the lower carnas-
sial in the Siberian form.
Comparison and species identication of the
Siberian form
The resemblance of the Siberian specimen to the
group of the largest fossil pantherine cats is beyond
question but its species identification requires spe-
cial discussion. The most complete diagnoses of Euro-
pean fossil lions P. fossilis and P. spelaea are reported
in Barycka (2008). The data of many authors on crani-
al and dental characteristics of the taxa are summarized
Figure 3 The mandible of Panthera fossilis (IGM-519). (a)
lateral view; (b) occlusal view; (c) lingual view. Scale bar =
50 mm.
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M. V. Sotnikova and I. V. Foronova
© 2013 International Society of Zoological Sciences, Institute of Zoology/
Chinese Academy of Sciences and Wiley Publishing Asia Pty Ltd
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in these diagnoses. Unfortunately, the mandibular fea-
tures are almost missing in them and the species and
subspecies differentiation of large lion-like felids from
the Pleistocene of Europe is mainly based on morpho-
metric analysis of dental characters (Schütt 1969; Hem-
mer 1974; Schütt & Hemmer 1978). The lack of teeth in
the Siberian specimen forced us to conduct a more de-
tailed analysis of the structure of the mandibular body
of large pantherine cats and to recognize additional fea-
tures characteristic of the fossil lions group.
One of the major characters that distinguish liv-
ing lions from tigers and jaguars is the form of mandi-
ble (Dawkins & Sanfort 1868; Merriam & Stock 1932;
Christiansen & Harris 2009). A mandible with slightly
convex ventral margin and anteriorly tapering horizon-
tal body is characteristic of living lions. Conversely, ac-
cording to Hemmer et al. (2001, 2010), the mandibles
of the Pleistocene and living jaguars demonstrate a less-
er depth of the horizontal corpus behind the molar than
in the diastema in front of p3. However, as the analysis
shows, tigers and jaguars more often retain their man-
dible shape, with a rectangular outline of the horizontal
body between p3 and m1 (Christiansen & Harris 2009).
The similar characters to a greater or lesser degree are
demonstrated by fossil Pleistocene lions of Europe and
North America (Fig. 4b,d,f,i). For instance, according
to Christiansen and Harris (2009, p. 939), a rectangu-
lar outline of the horizontal corpus of the mandible is
among the general characteristics of the American fos-
sil lion Panthera atrox. A rather straight ventral margin
of the mandibular corpus below the cheek teeth row is
also common for the group of the fossil European lions
(Barycka 2008).
This similar structure of mandible as a whole is char-
acteristic of the most ancient Villafranchian panther-
ine cats, such as Panthera toscana (Schaub, 1949), and
Panthera palaeosinensis (Zdansky, 1924), as well as of
the oldest known member of the Leo subgenus, Panthera
sp. from the Olduvai upper Bed-II in Africa (Fig. 4g).
Table 1 Comparative measurements of mandibles and lower dents of Panthera (Leo) fossilis, related Eurasian forms, and the largest
specimens of Panthera (Leo) atrox
Measurements
(mm)
Early–Middle Pleistocene Late Pleistocene
Asia Europe North America
Panthera (Leo) fossilis P. (L) atrox
12345678910 11 12
Length:
P3-m1 90.0 83.8 73.2 81.5 84.5 76.0 80.0 79.0 — 69.0–79.5 89.0 86.0
P4 30.0†32.3 25.0 — 31.3 28.1 28.5 31.0 30.3 27.3–32.8 32.3 32.1
M1 31.2†36.0 28.4 33.0‡31.3 31.0 31.0 31.0 31.1 28.3–32.9 33.7 33.9
Depth:
anteriorly/at p3 62.0 62.0 54.6 — 58.0 56.0 — /53.0 — 41.5–55.0 — —
anteriorly/at p4 59.7 — — — — — — — 57.4 — /60.7 /59.7
behind/at m1 62.0 59.0 /51.5 — 60.0 — 67.0 58.0 61.0 —
Thickness:
anteriorly to p3 31.0 — 30.8 — 26.9 — — — — — — —
at p4 32.0 27.3 — — — — — — — — — —
behind/at m1 /30.0 /26.1 — — 31.6 — — — — — 29.2 36.9
1 Panthera fossilis: IGM-519, Kuznetsk Basin, Russia (this paper); 2 Panthera sp., Kurtak archeological area, Russia; 3 Panthera
youngi, Zhoukoudian 1, China; 4 Panthera leo or ?fossilis, NHM M6165, Pakeeld, Great Britain; 5 Panthera spelaea or ?fossi-
lis, K.14.6, Aze cave, France; 6 Panthera fossilis, KP136, Petralona, Greece; 7–8 Panthera fossilis, no. 1, no 2, Mosbach, Germa-
ny; 9 Panthera fossilis, ISEZ MF/320/6803, Wierzchowska Górna, Poland; 10 Panthera fossilis, Cromer–Mindel, Europe; 11–12
Panthera atrox, 2901-3, UC14001, RLB, North America. Measurements from: Ovodov and Tarasov (2009); Pei (1934); Lewis et
al. (2010); Baryshnikov and Petrova (2008); Argant (1988); Kurtén and Poulianos (1977); von Reichenau (1906); Barycka (2008);
Merriam and Stock (1932). †Alveolar length. ‡Measurements are taken on the image. Dashes indicate missing data.
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Early Pleistocene Panthera from Siberia
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Figure 4 Mandibles of selected members of Pleistocene fossil lions group from Eurasia and North America. Scale bar = 50 mm. (a)
Panthera fossilis Kuznetsk Basin, (IGM-519); (b) P. fossilis from Solymár, Hungary (Hankó & Korsós 2007, p. 42, g. 2); (c) Pan-
thera (Leo) sp. from Kurtak archeological area, Russia (Ovodov & Tarasov 2009, p. 90, g. 1a); (d) P. fossilis, KP136, Petralona,
Greece (Kurtén & Poulianos 1977, p. 113), Figure reversed; (e) P. youngi, Zhoukoudian 1, China (Pei 1934, table 23, g. 1c); (f)
P. fossilis, Mosbach type locality, Germany (Hemmer 1974, table 12); (g) Panthera (Leo) sp., Olduvai upper Bed II, Africa (Petter
1973, table 8, g. 1); (h) P. spelaea, Indigirka Basin, Russia (Ovodov & Tarasov 2009, p. 90, g. 1b); (i) P. atrox, LACMMC 2900-
3, RLB, North America (Christiansen & Harris 2009, p. 939, g. 4), gure reversed.
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Therefore, it would be more logical to consider the com-
bination of characters listed above as a primitive state in
the pantherine evolution, and not as features, as put for-
ward by Christiansen and Harris (2009), that confirm
the relationship between Panthera atrox and jaguars.
Consequently, an almost square shape of the central part
of the mandible of Siberian form demonstrates its plesi-
omorphic nature and, together with other features listed
below, permits its attribution to the group of fossil lions.
The alveolar length of p3, p4 and m1 of the Siberi-
an lion fall well within the range observed for Eurasian
specimens of P. spelaea and P. fossilis, but the length
of the cheek teeth row exceeds all measurements cit-
ed in the literature for the largest fossil lions. The ra-
tio Lm1/Lp4 = 101.3% of the studied form is within the
range of those values (97.5%–105.5%) given by Schütt
and Hemmer (1978) for P. fossilis from Mosbach and
Mauer. The estimated size of the lower canin of IGM-
519 (L/W = nearly 31/21) is also similar to the range
given by Argant (2010, g. 29) for the European speci-
mens of P. fossilis. In addition, the Siberian lion closely
matches P. fossilis in the great thickness of the mandib-
ular body, in the deep, distinctly outlined and highly-po-
sitioned anterior portion of the masseteric fossa (Fig.
4b,d,f), in the ratio of p4/m1 length, and in the large lin-
ear size of the p4 anterior root. This set of features dis-
tinguishes the Siberian specimen from P. spelaea but
shows its similarity to P. atrox (Fig. 4h,i). However, the
American form, as shown below, is a more advanced
member of the P. fossilis–P. atrox lineage.
DISCUSSION
Origin and European occurrence of Panthera
fossilis
The early history of lion-sized pantherine cats is as-
sociated with the African continent where a nd of Pan-
thera sp. with features of jaguars, leopards and, to a
lesser extent, lions is known from the Pliocene fau-
na (approximately 3.5 Ma) of the Laetoli site (Hemmer
2011). The specimen from Laetoli was commonly con-
sidered an ancestor of the lion group of pantherine felids
(Turner & Antón 1997; Werdelin & Lewis 2005), but
Hemmer et al. (2001) consider it to be a stem species
for the whole group of felids mentioned above. Сurrent-
ly, the mandible No. 1273 from the Olduvai upper Bed-
II dated at approximately 1.4–1.2 Ma is believed to be
the most ancient form of fossil lions. According to Hem-
mer et al. (2010), only this specimen actually shares
apomorphic dental features with the lion group of felids.
Other African nds previously assigned to lion-like fe-
lids and known from sediments dated at 1.87–1.12 Ma
can be classied only as the ancestral forms belonging
to the lion stem group (Hemmer 2011).
Despite the fact that most researchers consider the
Olduvai specimen to be a possible ancestor of Panthera
(Leo), little information is available on the morpholo-
gy of this mandible. Petter (1973) notes a straight ven-
tral margin and a massive mandibular body, as well as
incisors, with narrow and long cross-section, differing
in shape from those of P. leo. Vereshchagin (1971) point
out the strong development of the posterior basal cin-
gulum on р4 and a lion type of wide coronoid process
in specimen no. 1273. Hemmer (2011) mentions a lion
condition of P3 in this specimen and showed the simi-
larity in size and dental characters of the Olduvai man-
dible to that of P. fossilis from the Mosbach type locali-
ty in Germany (Hemmer 1974, tab. X1).
According to our analysis, the Olduvai specimen
(Fig. 4g) has a straight ventral margin of horizontal ra-
mus and equal depth of the latter in front of p3 and be-
hind m1, as well as a long p4 with relatively short main
cusp, strong anterior and posterior cusps and developed
posterior basal cingulum. The large anterior part of p4
supported by a strong root is also among these features.
These characters, both plesiomorphic (mandubular out-
line) and apomorphic (reduced p3 and complicated p4),
the African form shares with P. fossilis. Thus, the fea-
tures revealed in P. fossilis and the Olduvai lion indicate
their close relationship. This conrms the view of previ-
ous researchers that Panthera sp. from the Olduvai up-
per Bed-II is the ancestor of Eurasian fossil lions.
Lion-like pantherine felids dispersed to Europe from
Africa in the rst half of the Middle Pleistocene. Finds
of the oldest European lions dated in the time span
from the Cromer Complex (early Middle Pleistocene,
about 0.78 Ma) to the Holstein interglacial (mid-Mid-
dle Pleistocene, about 0.45 Ma) are rare. Nowadays all
of them are referred to P. fossilis that was described by
von Reichenau (1906) based on well-preserved fossils
from Mosbach and Mauer sands. Other most import-
ant ndings of P. fossilis of the rst half of the Middle
Pleistocene are also known from the localities West-
bury-sub-Mendip in Great Britain, Isernia la Pine-
ta in Italy, Château in France, Vértesszólós in Hungary,
and from Cromerian beds of Petralona Cave in Greece
(Freudenberg 1914; Dietrich 1968; Schütt 1969; Kurtén
& Poulianos 1977; Sala 1990; Argant et al. 2007; Hankó
& Korsós 2007; Hemmer 2011).
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Early Pleistocene Panthera from Siberia
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The earliest occurrence of P. fossilis in the European
mainland was established on the basis of a single upper
carnassial derived from the Isernia la Pineta site in Ita-
ly, and was dated at about 0.61 Ma (Sala 1990; Hemmer
2011). However, to date, the oldest known nding of a
lion-like cat in Europe is, apparently, a specimen from
Pakeeld in Suffolk, Great Britain, dated within MIS 17
around 0.68 Ma, or even MIS 19 around 0.75 Ma (Lew-
is et al. 2010). A poorly presented mandibular frag-
ment from Pakeeld was described as P. leo by Lewis et
al. (2010), but a distinctly labial position of premolars
in relation to the molar, spaced p4 and m1, as well as a
massive mandibular body beneath the diastema c1-p3
and large dental size as a whole do not eliminate a pos-
sible assignment of specimen M-6165 from Pakeeld to
P. fossilis.
Recent research indicated that major findings of P.
fossilis and similar forms were obtained from the Middle
Pleistocene sediments of Europe. Large lion-like felids
appeared not earlier than MIS 19–17 (0.75–0.68 Ma)
and were relatively rare forms in the early Middle Pleis-
tocene faunas, but by the end of the Middle Pleistocene
they occupied a prominent place in European faunal as-
semblages (for a summary see Argant et al. 2007; Bary-
cka 2008; Marciszak & Stefaniak 2010; Sabol 2011a).
At the same time, it should be noted that among the Eu-
ropean Pleistocene specimens there are forms of lions
with the so-called ‘transitional’ or ‘intermediate’ fea-
tures (Hemmer 1974; Schütt & Hemmer 1978; Argant
et al. 2007). These forms occurred in the latest Middle–
earliest Late Pleistocene of Europe. For instance, among
them are atypical forms with mixed characters of P. fos-
silis and P. spelaea from the Austrian site Repolusthöhle
and abundant lion remains from the Biśnik Cave (Schütt
& Hemmer 1978; Marciszak & Stefaniak 2010; Hem-
mer 2011).
The relationships between P. spelaea and P. fossi-
lis have not been fully elucidated. Most scientists con-
sider them as chronosubspecies of P. spelaea (Baryc-
ka 2008; Argant 2010; Marciszak & Stefaniak 2010) or
P. leo (Schütt & Hemmer 1978). Meanwhile, Sotniko-
va and Nikolsky (2006), based on the cranial characters,
believe that P. spelaea and P. fossilis are different spe-
cies. The latter record agrees well with the results of cla-
distic analysis of pantherine felids from the Pleistocene
of Europe conducted by Hankó and Korsós (2007). The
result of the study by Hankó and Korsós gave reason to
believe that P. spelaea was not a direct descendant of
P. fossilis, but represents a separate more advanced lin-
eage. These conclusions agree to some extent with the
scenario by Sabol (2011a), according to which in the
Middle Pleistocene (approximately in MIS 6) cave li-
ons were separated from hypothetical ancient local pop-
ulations of lion-like felids that penetrated into the Al-
pine regions. Initially P. spelaea became widespread in
the mountain areas of Europe and only in the Late Pleis-
tocene dispersed throughout Eurasia. According to this
scenario, relict populations of P. fossilis could have been
retained on the European plains by the last glacial time;
that is, until the time of global distribution of cave lions
in Eurasia (Sabol 2011a).
Such an interpretation of relationships between P.
fossilis and P. spelaea suggests that ‘intermediate’ forms
could be advanced members of the P. fossilis lineage
rather than being transitional from one species to anoth-
er. However, to conrm this hypothesis a detailed mor-
phological revision of the well-stratied latest Middle–
earliest Late Pleistocene nds of fossil lions is required.
Fossil Pleistocene pantherine felids of Asian
mainland
According to Qiu (2006), the oldest Asian remains of
Panthera sp. dated at 3.6–2.6 Ma were found in China
in the Late Pliocene deposits of the Yushe Basin; how-
ever, this material was not described in detail and its at-
tribution to the Panthera genus is not reliably justied.
Further development of the genus in this region can
be traced by nds of felids close in size to jaguar-like
members of the Panthera genus. These finds were re-
ported within faunal assemblages of Henan (Lok. 39),
Yangguo and Longdan as P. palaeosinensis (Zdansky
1924; Qiu 2006). Faunas of these sites are correlated
with mammalian unit NCMQ1 of the Chinese Quaterna-
ry continental scale and are dated within 2.6–1.3 Ma (Qiu
2006).
Panthera palaeosinensis was initially regarded as the
ancestral tiger (Hemmer 1967). Subsequently, it was
considered as the most primitive member of the lion or
leopard clade and even as an ancestor of the genus Pan-
thera (Mazák 2010). Analysis of P. palaeosinensis fea-
tures is beyond the scope of the present work, but it
should be noted that its mandibular morphology is clos-
er to that of the Early–Middle Pleistocene jaguar-like
felids than to the lion group of pantherine cats. Conse-
quently, the analysis of fossil data leaves no doubt that
the origin of the lion group of felids is associated with
the African continent, and the assumption of Mazák
(2010) about their Asian roots is not supported by factu-
al data.
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Information on fossil lions in China is conned to the
nding of a lion-sized mandible in Locality 1 of Zhouk-
oudian (ZKD) (Pei 1934). Views on the age of the
ZKD-1 fauna are quite variable but modern research-
ers support an age estimate of the main fossiliferous
layers of ZKD 1 within the time span of 0.69–0.42 Ma
(Qiu 2006). Pei (1934) created a new species P. youngi
for the ZKD-1 pantherine felid and noted its afnities to
both the cave lion and the American P. atrox. Hemmer
(1974) has also supported the attribution of this form
to the lion group of fossil felids. Moreover, Harrington
(1969) believed that the American fossil lion, the Eur-
asian cave lion and P. youngi are conspecific. Exam-
ination of the pictures figured by Pei (1934, fig. 40b,
pl. XXIII-1a–c) showed that the mandible from ZKD-
1, similarly to most specimens of P. fossilis, have mas-
sive mandidular body with a straight ventral margin.
Its fourth premolar also resembles P. fossilis in bear-
ing a strong anterior cusp supported by a large anterior
root, and a well-developed posterior cingular area lack-
ing a distinct cingular cusp. At the same time, a relative-
ly short p4 in P. youngi indicates its more derived posi-
tion compared to most European specimens of P. fossilis
(Table 1, Fig. 4e).
As noted above, there are few reports indicating the
Early or Middle Pleistocene occurrence of fossil lions
in Asian Russia. A skull of a giant pantherine cat (with
condylobasal length [CBL] = 422 mm and zygomatic
breadth [ZB] = 312 mm) was found together with Hys-
trix vinogradovi Argyropolo, 1941 and Ursus thibetanus
permjak Baryshnikov, 2001 in the Mokhnevskaya Cave,
Middle Ural Mountains, in the deposits correlated with
a warm stage of the late Middle Pleistocene (MIS 7) or
the last interglacial (MIS 5e) (Baryshnikov 2001, 2003).
Unfortunately, the skull was not available for detailed
study, as it is part of a private collection.
A mandible of a very large pantherine felid with
mandibular depth of 62.0 mm and Lm1 = 36.0 mm was
found on the beach in the area of the Berezhekovo local-
ity, on the left bank of the Yenisei River, Kurtak archeo-
logical area, near Krasnoyarsk (Fig. 1). This nd can be
presumably associated with the Middle Pleistocene sed-
iments exposed in the Berezhekovo section (Krukover
& Chekha 1999). The Kurtak specimen was described
as Panthera sp. (Ovodov & Tarasov 2009). It is similar
in size to the largest members of the lion group of the
Pleistocene felids (Table 1, Fig. 4c). This mandible mor-
phologically agrees with that of P. fossilis in having a
strong and massive horizontal ramus with straight low-
er border, narrow and well-outlined anterior part of the
masseteric fossa, and p4 with posterior cingular basin
bearing a ridge-like cingular cusp. However, unlike the
conditions observed in P. fossilis, the Kurtak specimen
has p4 signicantly shorter than m1, as also is seen in P.
youngi from ZKD-1.
The analysis of other findings presumably associat-
ed with the Middle Pleistocene sediments in the terri-
tory eastward from the Ural Mountains does not con-
rm their similarity with P. fossilis. For instance, Sabol
(2011a), referring to Vereshchagin (1971) and Sotniko-
va (2006), reports finds of fossil lions in the Middle
Pleistocene of Siberia, whereas Sotnikova (2006) only
notes the occurrence of jaguar-like members of pan-
therine cats in Transbaikalia. As for the ndings listed
in Vereshchagin (1971), a well-stratied material is de-
rived only from Late Paleolithic sites, whereas the ages
of older specimens require a detailed examination. Sev-
eral mandibles of Panthera spelaea were described from
the Adycha (MM6880) and Kolyma (ZIN 29405, GIN
833-104) basins from supposedly Middle Pleistocene
sediments (Baryshnikov & Boeskorov 2001). Howev-
er, according to the authors (pp. 8–9), this material was
found on the beach and its morphology is typical for P.
spelaea.
Thus, apart from the mandible from Kuznetsk Basin,
distribution of lions with features resembling P. fossilis
in the Pleistocene of Asia is conrmed by the presence
of P. youngi in ZKD-1 and the findings of very large
forms in the Kurtak archeological area and in the Mid-
dle Ural Mountains (Mokhnevskaya Cave). Compared
to the European nds, the Asian specimens are very few.
Nevertheless, keeping in mind the earliest nding of a
fossil lion in Kuznetsk Basin, we can assume that the
expansion of the oldest lions from Africa to Eurasia via
the Asian continent could also take place. New infor-
mation on the Early Pleistocene occurrence of P. fossilis
in central Siberia located quite close to the territory of
Beringia also supports the idea that this group of felids
could invade America much earlier than the cave lion P.
spelaea.
American Pleistocene lion Panthera atrox and its
relationship with Eurasian pantherine felids
It has long been thought that all the Pleistocene li-
on-like pantherine felids of North America belong to 1
species, P. atrox (Whitmore & Forster 1967; Harington
1969). Subsequently, Vereshchagin (1971) assumed that
Beringian lions from north-eastern Siberia, Alaska and
the Yukon territory are P. spelaea rather than P. atrox.
Kurtén (1985) placed the Beringian (Alaska/Yukon) li-
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Early Pleistocene Panthera from Siberia
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Figure 5 Late Pleistocene occurrence of Panthera spelaea and
Panthera atrox in North America (modied after Barnett et al.
2009, g. 1).
ons in P. spelaea and the southern North American form
in P. atrox. This opinion was later confirmed by mor-
phological and molecular records (Baryshnikov & Boe-
skorov 2001; Sotnikova & Nikolsky 2006; Barnett et al.
2009). To date, only nds of pantherine cats south of the
Late Pleistocene ice sheets are regarded as P. atrox (Fig.
5).
Despite the fact that some authors point to the sim-
ilarity of P. atrox to jaguars and even to tigers (sum-
marized in Christiansen & Harris 2009), most research
based on morphological data infers their attribution to
the group of fossil lions (Harington 1969; Vereshchagin
1971; Hemmer 1974; Kurtén & Anderson 1980; Barysh-
nikov & Boeskorov 2001; Sotnikova & Nikolsky 2006).
Nevertheless, Christiansen and Harris (2009) attempt-
ed to prove that P. atrox was, indeed, a species separate
from the other great cats and descended from an early
jaguar. The situation has become clear after recent stud-
ies of mitochondrial DNA of a living and Late Pleisto-
cene lions, P. spelaea and P. atrox, undertaken by Bar-
nett et al. (2009). The authors completely disprove the
postulated link between P. atrox and jaguar and show
that the ancient DNA sequences of the studied Pleisto-
cene forms strictly group with living lions.
Although the inclusion of P. atrox as a separate spe-
cies in the lion group of felids is currently beyond ques-
tion, its origin and relationships to other fossil lions are
still the subject of debate. Most authors believe that the
American lion descended from the ancient (Yamagu-
chi et al. 2004) or Beringian (Barnett et al. 2009; Hem-
mer 2011) populations of cave lions; others consider
it as being derived from the Chinese P. youngi (Har-
ington 1969) or from a hypothetical member of the
Asian lion-like group of felids Panthera ‘leo’ spp. (Sab-
ol 2011a).
However, results of the present study suggest a dif-
ferent scenario for the evolution of the American lion.
As is known, the largest forms of P. atrox were derived
from the Late Pleistocene asphalt beds of the Rancho
La Brea in the southern part of North America. On aver-
age, these lions are larger than P. spelaea, and their size
is comparable to that of P. fossilis. Despite the appar-
ent similarity in size of P. atrox and P. fossilis, morpho-
logical comparison of these taxa has never been done
and their relationship has not been discussed. Exclusion
of P. fossilis from comparative analysis has likely re-
sulted from the lack of its ndings in the territories of
Central Asia and Siberia intermediate between Europe
and America, and also from the fact that there is a long
break between the existence of P. fossilis in the early
Middle Pleistocene in Europe and the first occurrence
of fossil lions in America (Fairbanks, Alaska) during the
Illinoian glaciation. Our data from Siberia ll, to some
extent, a geographic gap in the Eurasian history of the
most ancient group of lions.
Examination of the Siberian nding revealed a close
afnity between Eurasian samples of P. fossilis and the
population of P. atrox from the collection of Rancho
La Brea, USA, described by Merriam and Stock (1932)
(RLB). Judging from the descriptions by Merriam and
Stock (1932) and Christiansen and Harris (2009), we
can summarize that the American lion closely resem-
bles P. fossilis in the nearly straight ventral mandibu-
lar outline, the narrow and very deep anterior part of the
masseteric fossa, the rectangular prole of the horizon-
tal ramus, the large p4 with well-developed addition-
al cusps and basal cingulum, as well as in the subequal
size of the p4 roots. Among these features, we consid-
er the morphology of the mandibular body as primitive.
Of the derived features uniting these 2 taxa, we note, on
average, a larger general size and more massive mandi-
ble than in other lion-like felids and a slightly more re-
duced area of the anterior cusp on p4 compared to p4
of the ancestral Panthera sp. from the Olduvai upper
Bed-II. In addition, Sotnikova and Nikolskiy (2006) list
some cranial features of P. atrox that characterize the
American lion as a very advanced form. Such characters
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are also observed in mandibles of P. atrox. The derived
conditions are represented by the development of a long
and ventrally-deected angular process and by a poste-
rior expansion of the coronoid process (Fig. 4i).
Thus, we can conclude that P. atrox evolved on the
morphological basis of the most ancient members of the
Eurasian lion group but it has developed its own ad-
vanced features. Its ancestor was probably P. fossilis,
which dispersed to America from Siberia (via the Bering
Land Bridge) earlier than P. spelaea did. The latter ap-
peared in American Beringia at the beginning of the last
glacial (Hemmer 1974; Schütt & Hemmer 1978; Barnett
et al. 2009), whereas the earliest migration wave nearly
did not leave ancient evidence in the history of panther-
ine cats of North America.
It is commonly believed that the oldest fossil lions
dispersed to America at the beginning of the Illinoian
glaciation. However, according to Harington (1969), the
species identication of the Illinoian specimen (P. fos-
silis–atrox group or P. spelaea) was not carried out and
its age estimate is also in doubt. Harington (1969, p.
1285) wrote: “Although the specimen may represent P.
atrox the evidence is uncertain. The fauna and chronolo-
gy of the fossiliferous deposits concerned deserve close
study.”
In general, American nds, namely, northern P. spe-
laea and southern P. atrox, are mainly late Late Pleis-
tocene in age (Harington 1969; Stuart & Lister 2011).
Despite the fact that both species occurred in Ameri-
ca almost simultaneously, P. atrox has retained far more
primitive features than the Late Pleistocene Eurasian
and American P. spelaea. Such a combination of charac-
ters could have been formed in P. atrox owing to a rap-
id penetration of P. fossilis southward and its subsequent
isolation in North America south of the Late Pleistocene
ice sheets. The age of penetration of the most ancient li-
ons to America is unknown but molecular data indi-
cate that the genetic isolation of P. atrox occurred in the
Middle Pleistocene approximately 340 ka (Barnett et al.
2009).
ACKNOWLEDGMENTS
We gratefully acknowledge Nikolai Ovodov for pro-
viding additional data on the material from Kurtak ar-
chaeological area. We thank Alexey Tesakov for scien-
tific discussion that critically improved the paper. We
are also indebted to Katya Firsova and Igor Foronov,
who kindly prepared the design of gures for the pur-
pose of this study. M. Sotnikova acknowledges the fi-
nancial support from the Russian Foundation for Basic
Research, project 12-05-00-904a.
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