First Asian record of Panthera (Leo) fossilis (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia.

Integrative Zoology 2014; 9: 517–530

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

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 1

Geological 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 find of P. fossilis first recorded in Asia considerably extends the current notion of the eastward expansion of the most ancient lions. The Siberian lion is geologically the oldest form and is dimensionally among the largest members of the group of fossil lions on the Eurasian continent. Although known by mandibular remains only, it is readily distinguished from Panthera (Leo) spelaea by a heavy built mandibular corpus with rectangular profile 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 primitive features than the coeval Eurasian cave lion P. (L.) spelaea. Key words: Early Pleistocene, evolution, Panthera fossilis, relationships, Western Siberia

INTRODUCTION Fossil members of the Felidae lion group Panthera (Leo) spp. have a complex history, with numerous species and subspecies described. Lions originated in Africa 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

Correspondence: Marina V. Sotnikova, Geological Institute of Russian Academy of Sciences, Pyzhewsky 7, 119017 Moscow, Russia. Email: [email protected]

widespread in the Holarctic, ranging from Africa and across Europe and Asia to America (Vereshchagin 1971; Hemmer 1974; Kurtén 1985; Turner & Antón 1997; Yamaguchi et al. 2004; Barnett et al. 2009). The Middle Pleistocene distribution of the oldest member of this group of Felidae, Panthera (Leo) fossilis (von Reichenau, 1906), has long been known, but only in Europe (von Reichenau 1906; Freudenberg 1914; Dietrich 1968; Kurtén 1968; Schütt 1969; Hemmer 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; Sabol 2011a).

© 2013 International Society of Zoological Sciences, Institute of Zoology/ Chinese Academy of Sciences and Wiley Publishing Asia Pty Ltd

517

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

M. V. Sotnikova and I. V. Foronova

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

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 pantherine cats (Pei 1934; Harington 1969). Numerous finds of fossil lions in Asian Russia are known from the papers by Riabinin (1919), Gromova (1932), Vangengeim (1961), Vereshchagin (1971), Alexeeva (1980), Foronova (1982, 1999, 2001), Baryshnikov 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 Panthera (Leo) spelaea (Goldfuss, 1810). Middle Pleistocene large lion-like felids in Russia have been reported occasionally but these finds 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 first 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 first 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 oldest and metrically the largest member of the fossil lions in Eurasia. The objective of the present paper is to provide a detailed description and comparison of the Siberian specimen with the similar-sized taxa. This study will also help to expand the existing knowledge of the morphology 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 pantherine felid from Kuznetsk Basin and large Pleistocene lion-like cats of Europe and North America. For com-

518

parison we used collections of P. spelaea from the V.S. Sobolev Institute of Geology and Mineralogy, Siberian Branch of Russian Academy of Sciences, Novosibirsk (IGM SB RAS), the Geological Institute, Russian Academy of Science, Moscow (GIN RAS) and the Zoological Institute, Russian Academy of Science, St. Petersburg (ZIN RAS). Additional morphological and dimensional information was obtained from the literature. In this study, we used a system of standard measurements 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, fig. 6), whereas other paleontologists call this cusp hypoconid (Hankó & Korsós 2007, fig. 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 alveolar measurements. Considering the fact that in P. spelaea 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 finding with the teeth parameters of other fossil lion-like pantherine felids. The studied material is housed at IGM SB RAS, Novosibirsk. All measurements are in millimeters.

RESULTS Remarks on the geological setting of Siberian Panthera Hemmer (2011), referring to Foronova (1998), questions 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 evidence for the occurrence of a large fossil lion in the Siberian Early Pleistocene to be unreliable. The original paper (Foronova 1998, p. 358) includes brief information 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


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

Early Pleistocene Panthera from Siberia

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

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 figure 5 (p. 359) the analyzed mandible IGM-519 from the Bachatsk Quarry is incorrectly numbered 3608. This number belongs to the maxilla from the Mokhovo Quarry, but its image is missing in figure 5. In addition, in figure 3 (p. 357), which shows numerous fossiliferous sections from different quarries, the section of the Bachatsk Quarry that contained mandible IGM-519 is absent. The inaccuracies made in this publication have created the impression that the mandible from the Bachatsk Quarry was found in an obscure geological position. We hope that the following information on the geological and stratigraphic position of the material described and on the associated fauna completely eliminates the misunderstanding. The Kuznetsk Basin is the largest intermontane depression located in the south-eastern part of Western Siberia (Fig. 1). In the Late Cenozoic an almost continuous sequence of Quaternary sediments, over 150 m thick, ranging from the beginning of the Early Pleistocene to the Holocene, was accumulated in this area. These alternating deluvial–proluvial and subaqueous sequences 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 fairly complete paleontological and magnetostratigraphic characteristics. On the basis of these data in the composite stratotype section of the region, the major chronostratigraphic boundaries at 0.8 and 1.8 Ma are recognized and reliably characterized, and the paleomagnetic marker events (i.e. the Brunhes/Matuyama boundary and Jaramillo 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 sediments of 2 extremely diachronous and lithologically-different subaqueous formations. Very compact gray–green sandy clays of Sagarlyk Fm are overlain with a significant 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 authors (IF) in an open part of a large bone-bearing layer 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-

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 Figure 1 Location of the Kuznetsk Basin (A) and Kurtak ar- 27 28 cheological area (B). 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 Figure 2 Stratigraphy of the Early–Middle Quaternary deposits 49 of the Kuznetsk Basin and position of Panthera fossilis (IGM- 50 51 519).


519


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

ly Pleistocene of China. Previously, this peculiar ancient elephant was attributed by various paleontologists either to Archidiskodon or Palaeoloxodon, but presently Chinese and Japanese scientists (Wei et al. 2006) believe that it belongs to Mammuthus meridionalis (Nesti, 1825) (=Archidiskodon meridionalis according to taxonomy 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 filled with dense cemented and ferruginized enclosing deposits. According to these fossilization features, it does not differ from other bones in the Sagarlyk Fm. The biostratigraphic characteristics of this part of the formation are supplemented with findings in other sections of Allophaiomys pliocaenicus Kormos, 1932, Prolagurus pannonicus Kormos, 1930, Archidiskodon 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 mandible, teeth of meridionaloid elephants and bones of other mentioned forms are normally magnetized and are correlated 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 paleomagnetic record, permit the correlation of the Sagarlyk fauna with the Tamanian assemblage of European Russia and with faunas of the Early Galerian of Western Europe (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 formation permit its subdivision into heterochronous fossiliferous layers. However, in the studied section the Sagarlyk 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 terminal 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-

520

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, paleontological 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, figure 2; Foronova 2005: 97, figure 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 processes are missing (Fig. 3a–c). Locality: Southeast of Western Siberia, Kuznetsk Basin, 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 supported 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. Within this subgenus, according to the latest morphological 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.


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

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 masseteric fossa reaches forward to the level of the space between 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 midline of the mandibular body in a dorso–ventral direction. The basin between the anterior border of the masseteric 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 broken, and the posterior one is doubled and is situated below 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 greatest and smallest diameters of 7.1/3.5 mm and 11.0/7.7 mm, respectively. Like the ventral margin of the anterior part of the masseteric fossa, the posterior mental foramen is located near the horizontal midline of the mandibular body (Fig. 3a). In lingual view the posterior border of the symphyseal surface is nearly vertically oriented. Its most posteriorly projecting part does not reach the level of the middle part of the diastema between canine and p3 (Fig. 3b). In occlusal view, the space for incisors is moderately wide; the postcanine diastema is long, with length (L) of approximately 28.5 mm. According to the alveolar structure, the lower canine had a large size and a vertically-oriented root. Its estimated length and width are approximately 31.0 and 21.0 mm. Judging by the alveoli, the cheek-tooth series slightly arches buccally. In the tooth row, premolars occupy a clearly more labial position 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 diastema (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 posterior root (Lr = 12.2 mm), whereas the roots of p4 are

nearly equal in size, with Lr/Lr = 13.1 and 13.2 mm, respectively. The m1 crown is supported by a large anterior 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 inner part of the posterior root of m1 beyond the lingual margin of anterior root is also seen on the lower carnassial in the Siberian form.

Comparison and species identification 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 special discussion. The most complete diagnoses of European fossil lions P. fossilis and P. spelaea are reported in Barycka (2008). The data of many authors on cranial 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.


521

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

in these diagnoses. Unfortunately, the mandibular features 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 morphometric analysis of dental characters (Schütt 1969; Hemmer 1974; Schütt & Hemmer 1978). The lack of teeth in the Siberian specimen forced us to conduct a more detailed analysis of the structure of the mandibular body of large pantherine cats and to recognize additional features characteristic of the fossil lions group. One of the major characters that distinguish living lions from tigers and jaguars is the form of mandible (Dawkins & Sanfort 1868; Merriam & Stock 1932; Christiansen & Harris 2009). A mandible with slightly convex ventral margin and anteriorly tapering horizontal body is characteristic of living lions. Conversely, according to Hemmer et al. (2001, 2010), the mandibles of the Pleistocene and living jaguars demonstrate a lesser 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 mandible 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 rectangular outline of the horizontal corpus of the mandible is among the general characteristics of the American fossil 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 characteristic of the most ancient Villafranchian pantherine 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) Length: P3-m1 P4 M1 Depth: anteriorly/at p3 anteriorly/at p4 behind/at m1 Thickness: anteriorly to p3 at p4 behind/at m1

Early–Middle Pleistocene

Late Pleistocene

Asia

North America

Europe

Panthera (Leo) fossilis

P. (L) atrox

1

2

3

4

5

6

7

8

9

10

11

12

90.0 30.0† 31.2†

83.8 32.3 36.0

73.2 25.0 28.4

81.5 — 33.0‡

84.5 31.3 31.3

76.0 28.1 31.0

80.0 28.5 31.0

79.0 31.0 31.0

— 30.3 31.1

69.0–79.5 27.3–32.8 28.3–32.9

89.0 32.3 33.7

86.0 32.1 33.9

62.0 59.7 62.0

62.0 — 59.0

54.6 — /51.5

— — —

58.0 — 60.0

56.0 — —

— — 67.0

/53.0 — 58.0

— 57.4 61.0

41.5–55.0 — —

— /60.7

— /59.7

31.0 32.0 /30.0

— 27.3 /26.1

30.8 — —

— — —

26.9 — 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, Pakefield, Great Britain; 5 Panthera spelaea or ?fossilis, K.14.6, Aze cave, France; 6 Panthera fossilis, KP136, Petralona, Greece; 7–8 Panthera fossilis, no. 1, no 2, Mosbach, Germany; 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.

522


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

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, fig. 2); (c) Panthera (Leo) sp. from Kurtak archeological area, Russia (Ovodov & Tarasov 2009, p. 90, fig. 1a); (d) P. fossilis, KP136, Petralona, Greece (Kurtén & Poulianos 1977, p. 113), Figure reversed; (e) P. youngi, Zhoukoudian 1, China (Pei 1934, table 23, fig. 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, fig. 1); (h) P. spelaea, Indigirka Basin, Russia (Ovodov & Tarasov 2009, p. 90, fig. 1b); (i) P. atrox, LACMMC 29003, RLB, North America (Christiansen & Harris 2009, p. 939, fig. 4), figure reversed.


523

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

Therefore, it would be more logical to consider the combination of characters listed above as a primitive state in the pantherine evolution, and not as features, as put forward 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 plesiomorphic 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 Siberian 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 cited in the literature for the largest fossil lions. The ratio 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 IGM519 (L/W = nearly 31/21) is also similar to the range given by Argant (2010, fig. 29) for the European specimens of P. fossilis. In addition, the Siberian lion closely matches P. fossilis in the great thickness of the mandibular body, in the deep, distinctly outlined and highly-positioned anterior portion of the masseteric fossa (Fig. 4b,d,f), in the ratio of p4/m1 length, and in the large linear size of the p4 anterior root. This set of features distinguishes 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 associated with the African continent where a find of Panthera sp. with features of jaguars, leopards and, to a lesser extent, lions is known from the Pliocene fauna (approximately 3.5 Ma) of the Laetoli site (Hemmer 2011). The specimen from Laetoli was commonly considered 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. Сurrently, the mandible No. 1273 from the Olduvai upper BedII dated at approximately 1.4–1.2 Ma is believed to be the most ancient form of fossil lions. According to Hemmer et al. (2010), only this specimen actually shares

524

apomorphic dental features with the lion group of felids. Other African finds previously assigned to lion-like felids and known from sediments dated at 1.87–1.12 Ma can be classified 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 morphology of this mandible. Petter (1973) notes a straight ventral 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 cingulum 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 similarity in size and dental characters of the Olduvai mandible to that of P. fossilis from the Mosbach type locality in Germany (Hemmer 1974, tab. X1). According to our analysis, the Olduvai specimen (Fig. 4g) has a straight ventral margin of horizontal ramus and equal depth of the latter in front of p3 and behind 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 outline) and apomorphic (reduced p3 and complicated p4), the African form shares with P. fossilis. Thus, the features revealed in P. fossilis and the Olduvai lion indicate their close relationship. This confirms the view of previous researchers that Panthera sp. from the Olduvai upper Bed-II is the ancestor of Eurasian fossil lions. Lion-like pantherine felids dispersed to Europe from Africa in the first 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-Middle 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 important findings of P. fossilis of the first half of the Middle Pleistocene are also known from the localities Westbury-sub-Mendip in Great Britain, Isernia la Pineta 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).


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

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 Italy, and was dated at about 0.61 Ma (Sala 1990; Hemmer 2011). However, to date, the oldest known finding of a lion-like cat in Europe is, apparently, a specimen from Pakefield in Suffolk, Great Britain, dated within MIS 17 around 0.68 Ma, or even MIS 19 around 0.75 Ma (Lewis et al. 2010). A poorly presented mandibular fragment from Pakefield 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 possible assignment of specimen M-6165 from Pakefield 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 Pleistocene faunas, but by the end of the Middle Pleistocene they occupied a prominent place in European faunal assemblages (for a summary see Argant et al. 2007; Barycka 2008; Marciszak & Stefaniak 2010; Sabol 2011a). At the same time, it should be noted that among the European Pleistocene specimens there are forms of lions with the so-called ‘transitional’ or ‘intermediate’ features (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. fossilis and P. spelaea from the Austrian site Repolusthöhle and abundant lion remains from the Biśnik Cave (Schütt & Hemmer 1978; Marciszak & Stefaniak 2010; Hemmer 2011). The relationships between P. spelaea and P. fossilis have not been fully elucidated. Most scientists consider them as chronosubspecies of P. spelaea (Barycka 2008; Argant 2010; Marciszak & Stefaniak 2010) or P. leo (Schütt & Hemmer 1978). Meanwhile, Sotnikova and Nikolsky (2006), based on the cranial characters, believe that P. spelaea and P. fossilis are different species. The latter record agrees well with the results of cladistic 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 lineage. These conclusions agree to some extent with the

scenario by Sabol (2011a), according to which in the Middle Pleistocene (approximately in MIS 6) cave lions were separated from hypothetical ancient local populations of lion-like felids that penetrated into the Alpine regions. Initially P. spelaea became widespread in the mountain areas of Europe and only in the Late Pleistocene 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 another. However, to confirm this hypothesis a detailed morphological revision of the well-stratified latest Middle– earliest Late Pleistocene finds 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; however, this material was not described in detail and its attribution to the Panthera genus is not reliably justified. Further development of the genus in this region can be traced by finds of felids close in size to jaguar-like members of the Panthera genus. These finds were reported 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 Quaternary 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 Panthera (Mazák 2010). Analysis of P. palaeosinensis features is beyond the scope of the present work, but it should be noted that its mandibular morphology is closer to that of the Early–Middle Pleistocene jaguar-like felids than to the lion group of pantherine cats. Consequently, 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 factual data.


525

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

Information on fossil lions in China is confined to the finding of a lion-sized mandible in Locality 1 of Zhoukoudian (ZKD) (Pei 1934). Views on the age of the ZKD-1 fauna are quite variable but modern researchers 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 affinities 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 Eurasian cave lion and P. youngi are conspecific. Examination of the pictures figured by Pei (1934, fig. 40b, pl. XXIII-1a–c) showed that the mandible from ZKD1, similarly to most specimens of P. fossilis, have massive mandidular body with a straight ventral margin. Its fourth premolar also resembles P. fossilis in bearing a strong anterior cusp supported by a large anterior root, and a well-developed posterior cingular area lacking a distinct cingular cusp. At the same time, a relatively short p4 in P. youngi indicates its more derived position 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 Hystrix 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 locality, on the left bank of the Yenisei River, Kurtak archeological area, near Krasnoyarsk (Fig. 1). This find can be presumably associated with the Middle Pleistocene sediments 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 morphologically agrees with that of P. fossilis in having a strong and massive horizontal ramus with straight lower border, narrow and well-outlined anterior part of the

526

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 significantly shorter than m1, as also is seen in P. youngi from ZKD-1. The analysis of other findings presumably associated with the Middle Pleistocene sediments in the territory eastward from the Ural Mountains does not confirm their similarity with P. fossilis. For instance, Sabol (2011a), referring to Vereshchagin (1971) and Sotnikova (2006), reports finds of fossil lions in the Middle Pleistocene of Siberia, whereas Sotnikova (2006) only notes the occurrence of jaguar-like members of pantherine cats in Transbaikalia. As for the findings listed in Vereshchagin (1971), a well-stratified material is derived only from Late Paleolithic sites, whereas the ages of older specimens require a detailed examination. Several 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). However, 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 confirmed by the presence of P. youngi in ZKD-1 and the findings of very large forms in the Kurtak archeological area and in the Middle Ural Mountains (Mokhnevskaya Cave). Compared to the European finds, the Asian specimens are very few. Nevertheless, keeping in mind the earliest finding 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 information 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 lion-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-


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

ons in P. spelaea and the southern North American form in P. atrox. This opinion was later confirmed by morphological and molecular records (Baryshnikov & Boeskorov 2001; Sotnikova & Nikolsky 2006; Barnett et al. 2009). To date, only finds 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 similarity of P. atrox to jaguars and even to tigers (summarized 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; Baryshnikov & Boeskorov 2001; Sotnikova & Nikolsky 2006). Nevertheless, Christiansen and Harris (2009) attempted 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 studies of mitochondrial DNA of a living and Late Pleistocene lions, P. spelaea and P. atrox, undertaken by Barnett 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 Pleistocene forms strictly group with living lions. Although the inclusion of P. atrox as a separate species in the lion group of felids is currently beyond question, 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 (Yamaguchi et al. 2004) or Beringian (Barnett et al. 2009; Hemmer 2011) populations of cave lions; others consider it as being derived from the Chinese P. youngi (Harington 1969) or from a hypothetical member of the Asian lion-like group of felids Panthera ‘leo’ spp. (Sabol 2011a). However, results of the present study suggest a different 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 average, these lions are larger than P. spelaea, and their size is comparable to that of P. fossilis. Despite the apparent similarity in size of P. atrox and P. fossilis, morphological comparison of these taxa has never been done and their relationship has not been discussed. Exclusion of P. fossilis from comparative analysis has likely resulted from the lack of its findings 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 fill, to some extent, a geographic gap in the Eurasian history of the most ancient group of lions. Examination of the Siberian finding revealed a close affinity 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 resembles P. fossilis in the nearly straight ventral mandibular outline, the narrow and very deep anterior part of the masseteric fossa, the rectangular profile of the horizontal ramus, the large p4 with well-developed additional cusps and basal cingulum, as well as in the subequal size of the p4 roots. Among these features, we consider 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 mandible than in other lion-like felids and a slightly more reduced 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

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 Figure 5 Late Pleistocene occurrence of Panthera spelaea and 49 Panthera atrox in North America (modified after Barnett et al. 50 51 2009, fig. 1).


527


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

are also observed in mandibles of P. atrox. The derived conditions are represented by the development of a long and ventrally-deflected angular process and by a posterior 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 advanced 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 appeared 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 pantherine 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 identification of the Illinoian specimen (P. fossilis–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 chronology of the fossiliferous deposits concerned deserve close study.” In general, American finds, namely, northern P. spelaea and southern P. atrox, are mainly late Late Pleistocene in age (Harington 1969; Stuart & Lister 2011). Despite the fact that both species occurred in America almost simultaneously, P. atrox has retained far more primitive features than the Late Pleistocene Eurasian and American P. spelaea. Such a combination of characters could have been formed in P. atrox owing to a rapid 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 lions to America is unknown but molecular data indicate 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 providing additional data on the material from Kurtak archaeological area. We thank Alexey Tesakov for scientific discussion that critically improved the paper. We are also indebted to Katya Firsova and Igor Foronov, who kindly prepared the design of figures for the purpose of this study. M. Sotnikova acknowledges the fi-

528

nancial support from the Russian Foundation for Basic 1 2 Research, project 12-05-00-904a.

REFERENCES Alexeeva EV (1980). Pleistocene Mammals of Southeast of Western Siberia. Nauka, Moscow. (In Russian.) Argant A (1988). Etude de l’exemplaire de Panthera spelaea (Goldfuss, 1810) (Mammalia, Carnivora. Felidae) du gisement Pleistocene moyen recent de la grotte d’Aze (Saône-et-Loire). Revue de Paléobiologie 7, 449–66. Argant A (2010). Carnivores (Canidae, Felidae et Ursidae) de Romain-la-Roche (Doubs, France) Genève. Revue de Paléobiologie 29, 495–601. Argant A, Jeannet M, Argant J, Erbajeva M (2007). The big cats of the fossil site Château Breccia Northern Section (Saône-et-Loire, Burgundy, France): stratigraphy, palaeoenvironment, ethology and biochronological dating. Courier Forschunginstitut Senckenberg 259, 121–40. Barnett R, Shapiro B, Barnes I et al. (2009). Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a Late Pleistocene reduction in genetic diversity. Molecular Ecology 18, 1668–77. Barycka E (2008). Middle and Late Pleistocene Felidae and Hyaenidae of Poland. Fauna Poloniae–Fauna Polski. Museum and Istitute of Zoology Polish Academy of Science, Warszawa. Baryshnikov G (2001). The Pleistocene black bear (Ursus thibetanus) from the Urals (Russia). Lynx (Praha), n. s., 2002 32, 33–43. Baryshnikov GF (2003). Pleistocene small porcupine from the Ural Mountains, Russia, with note on taxonomy of Hystrix vinogradovi (Rodentia, Hystricidae). Russian Journal of Theriology 2, 43–7. Baryshnikov G, Boeskorov G (2001). The Pleistocene cave lion Panthera spelaea (Carnivora, Felidae) from Yakutia, Russia. Cranium 18, 7–24. Baryshnikov GF, Petrova EA (2008). Cave lion (Panthera spelaea) from the Pleistocene of Chuvashiya, European Russia. Russian Journal of Theriology 7, 33–40. Burger J, Rosendahl W, Loreille O et al. (2004). Molecular phylogeny of the extinct cave lion Panthera leo spelaea. Molecular Phylogenetics and Evolution 30, 841–9.


3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

Christiansen P, Harris JM (2009). Craniomandibular morphology and phylogenetic affinities of Panthera atrox: implications for the evolution and paleobiology of the lion lineage. Journal of Vertebrate Paleontology 29, 934–45. Cuenca-Bescós G, García N (2007). Biostratigraphic succession of the Early and Middle Pleistocene mammal faunas of the Atapuerca cave sites (Burgos, Spain). Courier Forschunginstitut Senckenberg 259, 99–110. Dawkins WB, Sandfort WA (1868). The British Pleistocene Mammalia. I. The British Pleistocene Felidae. Palaeontographical Society, London. Dietrich WO (1968). Fossile Löwen im europäischen und afrikanischen Pleistozän. Paläeontologische Abhandlungen. Abteilung A–Palaozoologie 3, 93– 366. Foronova IV (1982). New finds of mammals in the Pleistocene deposits of the Kuznetsk Basin. Trudy Zoologicheskogo Instituta AN SSSR 111, 50–7. (In Russian.) Foronova IV (1998). Early Quaternary Mammals of Kuznetsk Basin (Southwestern Siberia). Mededelingen Nederlands Instituut voor Toegepaste Geowetenschappen TNO 60, 353–74. Foronova IV (1999). Quaternary Mammals and Stratigraphy of Kuznetsk Basin (Southwestern Siberia). Antropozoikum ‘Quaternary of Siberia’ 23, 71–98. Foronova IV (2001). Quaternary Mammals of the South-East of Western Siberia (Kuznetsk Basin): Phylogeny, Biostratigraphy, and Palaeoecology. Publishing House of SB RAS GEO, Novosibirsk. (In Russian.) Foronova IV (2005). Large mammal faunas from southwestern Siberia of the Plio–Pleistocene boundary and Lower/Middle Pleistocene transition. Quaternary International 131, 95–9. Freudenberg W (1914). Die Säugetiere des älteren Quartärs von Mitteleuropa. Geologische und Palaeontologische. Abhandlungen 12, 1–219. Gromova V (1932). New materials on the Quaternary fauna from the Volga River basin and on the mammalian history of Eastern Europe and Northern Asia in general. Trudy Komissii po Izucheniyu Сhetvertichnogo Perioda 2, 69–180. (In Russian.) Hankó E, Korsós Z (2007). A cladistic analysis of the teeth and mandible morphological characters of Pleistocene lions from Hungary. Állattani Közlemének 92, 39–51.

Harington CR (1969). Pleistocene remains of the lion-like cat (Panthera atrox) from the Yukon Territory and northern Alaska. Canadian Journal of Earth Sciences 6, 1277–88. Hemmer H (1967). Wohin gehört ‘Felis’ palaeosinensis Zdansky, 1924 in systematischer Hinsicht? Neues Jahrbuch für Geologie und Paleontologie Abhandlungen 129, 83–96. Hemmer H (1974). Untersuchungen zur Stammesgeschichte der Pantherkatzen (Pantherinae). Teil III. Zur Artgeschichte des Löwen, Panthera (Panthera) leo (Linnaeus 1758). Ver öffentlichungen der Zoologischen Staatssammlung München 17, 167–280. Hemmer H (2011). The story of the cave lion Panthera leo spelaea (Goldfuss, 1810)–a review. Quaternaire, Hors-série 4, 201–8. Hemmer H, Kahlke R-D, Vekua AK (2001). The jaguar–Panthera onca gombaszoegensis (Kretzoi, 1938) (Carnivora: Felidae) in the late Lower Pleistocene of Akhalkalaki (south Georgia, Transcaucasia) and its evolutionary signiﬁcance. Geobios 34, 475–86. Hemmer H, Kahlke R-D, Vekua AK (2010). Panthera onca georgica ssp. nov. from the Early Pleistocene of Dmanisi (Republic of Georgia) and the phylogeography of jaguars (Mammalia, Carnivora, Felidae). Neues Jahrbuch für Geologie und Paläontologie 257, 115–27. Krukover AA, Chekha VP (1999). Small fossil mammals from the Kurtak archaeological region (Kranoyarsk District). Antropozoikum ‘Quaternary of Siberia’ 23, 71–98. Kurtén B (1968). Pleistocene Mammals of Europe. Weidenfeld and Nicolson, London. Kurtén B (1985). The Pleistocene lion of Beringia. Annales Zoologici Fennici 22, 117–21. Kurtén B, Anderson E (1980). Pleistocene Mammals of North America. Columbia University Press, NY. Kurtén B, Poulianos N (1977). New stratigraphic and faunal material from Petralona Cave with special reference to the Carnivora. Anthropos 4, 47–130. Lewis M, Pacher M, Turner A (2010). The larger Carnivora of the West Runton Freshwater Bed. Quaternary International 228, 116–35. Marciszak A, Stefaniak K (2010). Two forms of cave lion: Middle Pleistocene Panthera spelaea fossilis Reichenau, 1906 and Upper Pleistocene Panthera spelaea spelaea Goldfuss, 1810 from the Biśnik Cave, Poland. Neues Jahrbuch für Geologie und Paläontologie 258, 339–51.


529

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

Mazák JH (2010). What is Panthera palaeosinensis? Mammal Review 40, 90–102. Merriam JC, Stock C (1932). The Felidae of Rancho La Brea. Carnegie Institution of Washington Publications 442, 1–231. Ovodov ND, Tarasov AY (2009). Big cat (Panthera sp.) and a small cave bear in Siberia. In: Eniseiskaya provinciya. Paleontologiya i speleologiya 4, 86–92. (In Russian.) Pei WC (1934). On the Carnivora from Locality 1 of Choukoutien. Palaeontologia Sinica. (Ser. C) 8, 1–216. Petter G (1973). Carnivores Pleistocènes du ravin d’Olduvai (Tanzanie). In: Leakey LSB, Savage RJG, Coryndon SC, eds. Fossil vertebrates of Africa 3, Academic Press, London, pp. 43–100. Qiu Z-X (2006). Quaternary environmental changes and evolution of large mammals in north China. Vertebrata PalAsiatica 4, 109–32. Riabinin A (1919). The fossil lions from the Ural Mountains and Volga River Basin. Memoires du Comité géologie. NS 168, 1–24. (In Russian.) Sabol M (2011a). Masters of the lost word: a hypothetical look at the temporal and spatial distribution of lion-like felids. Quaternaire Hors-serie 4, 229–36. Sabol M (2011b). A record of Pleistocene lion-like felids in the territory of Slivakia. Quaternaire Hors-serie 4, 215–28. Sala B (1990). Panthera leo fossilis (v. Reichenau, 1906) (Felidae) de Isernia la Pineta (Pléistocène moyen inférieur d’Italie). Geobios 23, 189–94. Schütt G (1969). Untersuchungen am Gebiss von Panthera leo fossilis (v. Reichenau, 1906) und Panthera leo spelaea (Goldfuss, 1810). Neues Jahrbuch für Geologie und Paläontologie 134, 192–220. Schütt G, Hemmer H (1978). Zur Evolution des Löwen (Panthera leo L.) im europäischen Pleistozän. Neues Jahrbuch für Geologie und Paläontologie 4, 228–55. Sotnikova MV (2006). Pliocene–Early Pleistocene carnivore assemblages of Transbaikalian Area, Russia. In: Alexeeva NA, Erbajeva MA, Mironov GA, eds. Stratigraphy, Paleontology and Paleoenvironment of Pliocene–Pleistocene of Transbaikalia and Interregional Correlations. Publishing house of SB RAS, Ulan-Ude, pp. 84–5.

530

Sotnikova M, Nikolskiy P (2006). Systematic position of the cave lion Panthera spelaea (Goldfuss) based on cranial and dental characters. Quaternary International 142–3, 218–28. Stuart AJ, Lister AM (2011). Extinction chronology of the cave lion Panthera spelaea. Quaternary Science Reviews 30, 2329–40. Turner A, Antón M (1997). The Big Cats and their Fossil Relatives, Columbia University Press, New York. Vangengeim EA (1961). Paleontological Foundation of the Anthropogene Stratigraphy of Northeastern Siberia (on Mammals). Academy of Sciences of the USSR, Moscow. (In Russian.) Vereshchagin NK (1971). The cave lion and its history in the Holarctic and on the territory of the USSR. Trudy Zoologicheskogo Instituta AN USSR 49, 123– 99. (In Russian.) von Reichenau W (1906). Beiträge zur näheren Kenntnis der Carnivoren aus den Sanden von Mauer und Mosbach. Abhandlungen der Großherzoglichen Hessischen Geologischen Landesanstalt zu Darmstadt 4, 125. Whitmore FC, Foster HL (1967). Panthera atrox (Mammalia, Felidae) from Central Alaska. Journal of Paleontology 41, 247–51. Wei GB, Taruno H, Kawamura Y, Jin CZ (2006). Pliocene and Early Pleistocene primitive mammoths of Northern China: their revised taxonomy, biostratigraphy and evolution. Journal of Geosciences, Osaka City University 49, 59–101. Werdelin L, Lewis ME (2005). Plio–Pleistocene Carnivora of eastern Africa: species richness and turnover patterns. Zoological Journal of the Linnean Society 144, 121–44. Wolsan M (1993). Évolution des Carnivores Quaternaires en Europe Centrale dans leur Contexte Stratigraphique et Paléoclimatique. L’Anthropologie 97, 203–22. Yamaguchi N, Cooper A, Werdelin L, Macdonald DW (2004). Evolution of the mane and group-living in the lion (Panthera leo): a review. Journal of Zoology 263, 329–42. Zdansky O (1924). Jungtertiäre Carnivoren Chinas. Palaeontologia Sinica 2, 1–49.


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51

First record of Mylagaulid rodents (Rodentia, Mammalia) from the Miocene of Eastern Siberia (Olkhon Island, Baikal Lake, Irkutsk Region, Russia).

Mitochondrial genome of the African lion Panthera leo leo.

Late Pleistocene carnivores (Carnivora: Mammalia) from a cave sedimentary deposit in northern Brazil.

Fibroblastic osteosarcoma in a lion (Panthera leo).

Outbreak of pox disease among carnivora (felidae) and edentata.

Early impact of mammographic screening in Western Siberia.

Haemangiosarcoma in a captive Asiatic lion (Panthera leo persica).

Caesarean of Lion (Panthera leo) at Dulahajra Safari Park, Bangladesh.

Dual phylogenetic origins of Nigerian lions (Panthera leo).

A metapopulation approach to African lion (Panthera leo) conservation.

The first record of Deinotherium (Proboscidea, Mammalia) in the Miocene of Adygea.

The complete mitochondrial genome of the Servaline Genet, Genetta servalina, the first representative from the family Viverridae (Mammalia, Carnivora).

Prevalence of arterial hypertension in the Krasnoyarsk Krai (Siberia, Russia).

The complete mitochondrial genome of the Spotted Linsang, Prionodon pardicolor, the first representative from the family Prionodontidae (Mammalia, Carnivora).

Selected clinical, biochemical, and electrolyte alterations in anesthetized captive tigers (Panthera tigris) and lions (Panthera leo).

Developmental changes in pituitary-gonadal function in free-ranging lions (Panthera leo leo) of the Serengeti Plains and Ngorongoro Crater.

Carnivora: the primary structure of the major hemoglobin component from adult European lynx (Lynx lynx, Felidae).

Detection and genetic characterization of a wide range of infectious agents in Ixodes pavlovskyi ticks in Western Siberia, Russia.

Cope's rule and the evolution of body size in Pinnipedimorpha (Mammalia: Carnivora).

Neogene amphibians and reptiles (Caudata, Anura, Gekkota, Lacertilia, and Testudines) from the south of Western Siberia, Russia, and Northeastern Kazakhstan.

Mucoperiosteal exostoses in the tympanic bulla of African lions (Panthera leo).

Presence of the 54-chromosome common vole (Mammalia) on Olkhon Island (Lake Baikal, East Siberia, Russia), and the occurrence of an unusual X-chromosome variant.

The inability of the lion, Panthera leo, L. to desaturate linoleic acid.

Molecular evidence for a recent demographic expansion in the puma (Puma concolor) (Mammalia, Felidae).