<I>Panthera (Leo)

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

Article in Integrative Zoology · September 2014 DOI: 10.1111/1749-4877.12082

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Marina V. Sotnikova Irina V. Foronova Russian Academy of Sciences Russian Academy of Sciences

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1 ORIGINAL ARTICLE 1 2 2 3 3 4 4 5 5 6 6 7 First Asian record of Panthera (Leo) fossilis (Mammalia, 7 8 8 9 Carnivora, Felidae) in the Early Pleistocene of Western Siberia, 9 10 10 11 Russia 11 12 12 13 13 14 Marina V. SOTNIKOVA1 and Irina V. FORONOVA2 14 15 15 1Geological Institute of Russian Academy of Sciences, Moscow, Russia and 2V.S. Sobolev Institute of Geology and Mineralogy, 16 16 17 Siberian Branch of Russian Academy of Sciences, Novosibirsk, Russia 17 18 18 19 19 20 Abstract 20 21 21 22 A lion-like pantherine felid is described as Panthera (Leo) fossilis from the late Early Pleistocene sediments of 22 23 the Kuznetsk Basin (Western Siberia, Russia). The find of P. fossilis first recorded in Asia considerably extends 23 24 the current notion of the eastward expansion of the most ancient lions. The Siberian lion is geologically the old- 24 25 est form and is dimensionally among the largest members of the group of fossil lions on the Eurasian conti- 25 26 nent. Although known by mandibular remains only, it is readily distinguished from Panthera (Leo) spelaea by a 26 27 heavy built mandibular corpus with rectangular profile in the cheek teeth area, a deep, well-outlined and narrow 27 28 anterior section of the masseteric fossa, and a large р4 supported by a big unreduced anterior root. The Siberian 28 29 lion shares these features with the European Middle Pleistocene P. fossilis and the American Late Pleistocene P. 29 30 (Leo) atrox, which suggests their close relationship. P. atrox originated from P. fossilis and was isolated in North 30 31 America south of the Late Pleistocene ice sheets. This explains why the American lion has retained more primi- 31 32 tive features than the coeval Eurasian cave lion P. (L.) spelaea. 32 33 33 34 Key words: Early Pleistocene, evolution, Panthera fossilis, relationships, Western Siberia 34 35 35 36 36 37 37 38 INTRODUCTION widespread in the Holarctic, ranging from Africa and 38 across Europe and Asia to America (Vereshchagin 1971; 39 Fossil members of the Felidae lion group Panthera 39 40 Hemmer 1974; Kurtén 1985; Turner & Antón 1997; Ya- 40 (Leo) spp. have a complex history, with numerous spe- maguchi et al. 2004; Barnett et al. 2009). 41 cies and subspecies described. Lions originated in Afri- 41 The Middle Pleistocene distribution of the oldest 42 ca and dispersed northward into Europe during the early 42 member of this group of Felidae, Panthera (Leo) foss- 43 Middle Pleistocene, 0.75–0.68 Ma (Hemmer 2011). In 43 ilis (von Reichenau, 1906), has long been known, but 44 the Late Pleistocene, this group of pantherine cats was 44 45 only in Europe (von Reichenau 1906; Freudenberg 45 46 1914; Dietrich 1968; Kurtén 1968; Schütt 1969; Hem- 46 47 mer 1974, 2011; Kurtén & Poulianos 1977; Schütt & 47 48 Correspondence: Marina V. Sotnikova, Geological Institute of Hemmer 1978; Sala 1990; Wolsan 1993; Argant et al. 48 49 Russian Academy of Sciences, Pyzhewsky 7, 119017 Moscow, 2007; Cuenca-Bescós & García 2007; Hankó & Korsós 49 50 Russia. 2007; Barycka 2008; Marciszak & Stefaniak 2010; Sab- 50 51 Email: [email protected] ol 2011a). 51

1 Fossil evidence documenting the occurrence of parison we used collections of P. spelaea from the V.S. 1 2 early lions in the Middle Pleistocene of Asia is very Sobolev Institute of Geology and Mineralogy, Siberi- 2 3 poor. Outside Asian Russia, the only large Panthera an Branch of Russian Academy of Sciences, Novosi- 3 4 (Leo) youngi (Pei, 1934) from the Peking Man site of birsk (IGM SB RAS), the Geological Institute, Russian 4 5 Zhoukoudian 1, northeastern China, dated at 0.69–0.42 Academy of Science, Moscow (GIN RAS) and the Zo- 5 6 Ma (Qiu 2006), was referred to the lion group of pan- ological Institute, Russian Academy of Science, St. Pe- 6 7 therine cats (Pei 1934; Harington 1969). tersburg (ZIN RAS). Additional morphological and di- 7 8 Numerous finds of fossil lions in Asian Russia are mensional information was obtained from the literature. 8 9 known from the papers by Riabinin (1919), Gromo- In this study, we used a system of standard measure- 9 10 va (1932), Vangengeim (1961), Vereshchagin (1971), ments of lower jaws and teeth adopted by many authors 10 11 Alexeeva (1980), Foronova (1982, 1999, 2001), Barysh- (Merriam & Stock 1932; Schütt 1969; Vereshchagin 11 12 nikov and Boeskorov (2001), Sotnikova and Nikolskiy 1971; Baryshnikov & Boeskorov 2001). 12 13 (2006), Baryshnikov and Petrova (2008) and Ovodov The analysis shows that the terms used to describe 13 14 and Tarasov (2009). However, the material presented in the premolar cusps of Felidae do not match in various 14 15 these works mainly belongs to the Late Pleistocene Pan- authors. For instance, the posterior additional cusp on 15 16 thera (Leo) spelaea (Goldfuss, 1810). Middle Pleisto- р3–4 is referred to as metaconid in Barycka (2008, p. 16 17 cene large lion-like felids in Russia have been reported 28, fig. 6), whereas other paleontologists call this cusp 17 18 occasionally but these finds have been associated neither hypoconid (Hankó & Korsós 2007, fig. 1; Sabol 2011b). 18 19 with the European P. fossilis nor with Asian P. youngi Therefore, we accept the system of dental elements used 19 20 (Vereshchagin 1971). The assumption of the presence of by US researchers, in particular by Merriam and Stock 20 21 P. fossilis in the Asian part of Russia was first made by (1932), to describe non-carnassial teeth of felids, where 21 22 Baryshnikov and Boeskorov (2001) based on finds of the premolar cusps are designated as anterior, posterior, 22 23 very large lions’ limb bones in the Middle Pleistocene cingular and principal (or main) premolar cusps. 23 24 24 localities along the Volga and Lower Tunguska Rivers. The dental information of the mandible from the 25 25 However, the first clear evidence of a giant lion similar Kuznetsk Basin was obtained only on the basis of alve- 26 26 to the P. fossilis–P. youngi group in Asia is a mandible olar measurements. Considering the fact that in P. spe- 27 27 described by Foronova (1998, 2001, 2005) as Panthera laea the alveolar length of cheek teeth is very close to 28 28 sp. from the Early Pleistocene of the Kuznetsk Basin the dental length (with an error no more than 1.0–1.5 29 29 in Western Siberia. The mandibular size of this felid is mm), we believed it possible to compare the alveolar 30 30 close to that of the largest forms of fossil pantherine cats length of our finding with the teeth parameters of other 31 31 and in some parameters (Lp3-m1) even exceeds them. fossil lion-like pantherine felids. 32 Consequently, the Siberian form is geologically the old- 32 33 The studied material is housed at IGM SB RAS, No- 33 est and metrically the largest member of the fossil lions vosibirsk. All measurements are in millimeters. 34 in Eurasia. 34 35 35 The objective of the present paper is to provide a de- 36 RESULTS 36 tailed description and comparison of the Siberian spec- 37 37 imen with the similar-sized taxa. This study will also 38 Remarks on the geological setting of Siberian 38 help to expand the existing knowledge of the morphol- 39 39 ogy and the stratigraphic and geographic distribution of Panthera 40 40 the oldest members of the lion group in Asia, as well as 41 Hemmer (2011), referring to Foronova (1998), ques- 41 to clarify their relationship with European P. fossilis and 42 tions the correctness of the stratigraphic position of the 42 American P. (Leo) atrox (Leidy, 1853) distributed south 43 mandible of a large pantherine cat derived from the 43 of the Late Pleistocene ice sheets. 44 Sagarlyk Formation in the Bachatsk Quarry of Kuznetsk 44 45 Basin. In this regard, he considers the geological evi- 45 46 MATERIALS AND METHODS dence for the occurrence of a large fossil lion in the Si- 46 berian Early Pleistocene to be unreliable. The original 47 This study is based on a detailed morphological 47 paper (Foronova 1998, p. 358) includes brief informa- 48 analysis of mandibular and dental features of the pan- 48 tion on 2 fragments of Panthera sp. from the Kuznetsk 49 therine felid from Kuznetsk Basin and large Pleistocene 49 Basin, that is, a very large mandible IGM-519 from the 50 lion-like cats of Europe and North America. For com- 50 51 Sagarlyk Formation (Fm) of the Bachatsk Quarry, which 51

1 we describe in this article, and a maxilla (No. 3608) of Equus aff. suessenbornensis Wüst, 1901 and Equus 1 2 found in deposits of the younger Sergeevo Fm in the sanmeniensis Teilhard de Chardin and Piveteau, 1930 2 3 Mokhovo Quarry. In the caption to figure 5 (p. 359) the were also found in the same layer, as well as a molar of 3 4 analyzed mandible IGM-519 from the Bachatsk Quar- Archidiskodon sp. similar to Archidiskodon tokunagai 4 5 ry is incorrectly numbered 3608. This number belongs (Matsumoto) Teilhard and Trassaert, 1937 from the Ear- 5 6 to the maxilla from the Mokhovo Quarry, but its image 6 7 is missing in figure 5. In addition, in figure 3 (p. 357), 7 8 which shows numerous fossiliferous sections from dif- 8 9 ferent quarries, the section of the Bachatsk Quarry that 9 10 contained mandible IGM-519 is absent. The inaccura- 10 11 cies made in this publication have created the impres- 11 12 sion that the mandible from the Bachatsk Quarry was 12 13 found in an obscure geological position. We hope that 13 14 the following information on the geological and strati- 14 15 graphic position of the material described and on the 15 16 associated fauna completely eliminates the misunder- 16 17 standing. 17 18 The Kuznetsk Basin is the largest intermontane de- 18 19 pression located in the south-eastern part of Western Si- 19 20 beria (Fig. 1). In the Late Cenozoic an almost contin- 20 21 uous sequence of Quaternary sediments, over 150 m 21 22 thick, ranging from the beginning of the Early Pleis- 22 23 tocene to the Holocene, was accumulated in this area. 23 24 These alternating deluvial–proluvial and subaqueous se- 24 25 quences are recovered in large open coal pits, which 25 26 provide comprehensive biostratigraphic characteristics 26 27 of the Quaternary sediments of the region. The Early Figure 1 Location of the Kuznetsk Basin (A) and Kurtak ar- 27 28 Pleistocene sediments are represented upward from the cheological area (B). 28 29 base by deposits of the Mokhovo, Sagarlyk, Sergeevo 29 30 and the lowermost part of the Kedrovka Fm with fair- 30 31 ly complete paleontological and magnetostratigraphic 31 32 characteristics. On the basis of these data in the compos- 32 33 ite stratotype section of the region, the major chronos- 33 34 tratigraphic boundaries at 0.8 and 1.8 Ma are recognized 34 35 and reliably characterized, and the paleomagnetic mark- 35 36 er events (i.e. the Brunhes/Matuyama boundary and Ja- 36 37 ramillo Subchron) are recorded (Foronova 1998, 1999, 37 38 2001, 2005) (Fig. 2). 38 39 39 The Panthera mandible IGM-519 was found in the 40 40 Bachatsk Quarry, in the section represented by sedi- 41 41 ments of 2 extremely diachronous and lithologically-dif- 42 42 ferent subaqueous formations. Very compact gray–green 43 43 sandy clays of Sagarlyk Fm are overlain with a signifi- 44 44 cant hiatus by the lacustrine–alluvial gray–blue plastic 45 45 clays of the Kedrovka Fm. The subaerial deposits of the 46 46 Sergeevo Fm separating these subaqueous sequences in 47 47 the complete sections are missing in this outcrop. 48 48 The fossil lion find was revealed by one of the au- 49 Figure 2 Stratigraphy of the Early–Middle Quaternary deposits 49 thors (IF) in an open part of a large bone-bearing lay- 50 of the Kuznetsk Basin and position of Panthera fossilis (IGM- 50 51 er in the upper part of the Sagarlyk Fm. Bones and teeth 519). 51

1 ly Pleistocene of China. Previously, this peculiar an- blage is correlated with Tiraspolian and Viatkian faunas 1 2 cient elephant was attributed by various paleontologists of Russia, as well as with the Cromerian (excluding the 2 3 either to Archidiskodon or Palaeoloxodon, but present- earliest one) faunas of Western Europe. 3 4 ly Chinese and Japanese scientists (Wei et al. 2006) be- Thus, considering all available geological, paleonto- 4 5 lieve that it belongs to Mammuthus meridionalis (Nesti, logical and paleomagnetic data, it can be concluded that 5 6 1825) (=Archidiskodon meridionalis according to taxon- the age of the mandible described corresponds to the 6 7 omy accepted in Russia). time of accumulation of the upper part of the Sagarlyk 7 8 All fossil bones from Sagarlyk Fm are very heavy Fm; namely, to the Jaramillo Subchron of the uppermost 8 9 and strongly mineralized. The described mandible of Matuyama Chron (i.e. to the late Early Pleistocene). 9 10 dark brown color is massive and heavy; its hollows and 10 Systematic paleontology 11 cavities were also filled with dense cemented and ferru- 11 12 ginized enclosing deposits. According to these fossiliza- Order Carnivora Bowdich, 1821 12 13 13 tion features, it does not differ from other bones in the Family Felidae Fischer, 1817 14 Sagarlyk Fm. The biostratigraphic characteristics of this 14 Subfamily Pantherinae Pocock, 1917 15 part of the formation are supplemented with findings in 15 16 other sections of Allophaiomys pliocaenicus Kormos, Genus Panthera Oken, 1816 16 17 1932, Prolagurus pannonicus Kormos, 1930, Archidis- Subgenus Leo Brehm, 1829 17 18 kodon meridionalis tamanensis Dubrovo, 1964, E. aff. Panthera (Leo) fossilis (von Reichenau, 1906) 18 19 suessenbornensis, E. sanmeniensis, very large Alces aff. Panthera sp.: Foronova 1998: 358–359 part, figure 19 20 latifrons (Johnson, 1874) and Bison sp. (ex gr. priscus 5(1) – No. 519; Foronova 1999: 73,77 part; Foronova 20 21 Bojanus, 1827). 2001: 69–70 part, 196-197: table IV, figure 2; Foronova 21 22 22 The sediments of the upper part of the Sagarlyk Fm 2005: 97, figure 3. Panthera leo fossilis: Hemmer 2011, 23 23 bearing fossils including the described Panthera mandi- p. 203. 24 24 ble, teeth of meridionaloid elephants and bones of other Material: Horizontal ramus of the left mandible 25 25 mentioned forms are normally magnetized and are cor- (IGM-519) with the canine and p3 alveoli; with 2 roots 26 26 related with the Jaramillo Subchron of the Matuyama exposed inside the alveolus of p4 and a fractured crown 27 27 Chron (Fig. 2). Consequently, the evolutionary stage of m1. The anterior part of the symphyseal region is 28 28 of major fossils (i.e. rodents, meridionaloid elephants, partly destroyed; coronoid, condylar and angular pro- 29 29 horses and broad-fronted moose), along with the paleo- cesses are missing (Fig. 3a–c). 30 30 magnetic record, permit the correlation of the Sagarlyk 31 Locality: Southeast of Western Siberia, Kuznetsk Ba- 31 fauna with the Tamanian assemblage of European Rus- 32 sin, Bachatsk Quarry (geographic position: 54.28°N, 32 sia and with faunas of the Early Galerian of Western Eu- 33 86.16°E), upper part of the Sagarlyk Fm. 33 rope (Foronova 1998, 1999, 2001, 2005). 34 Age: Late Early Pleistocene. 34 The overlying deposits of the Kedrovka Fm in its 35 Taxonomic remarks 35 36 complete volume have a rather wide age range, and in 36 37 the most part are correlated with the Brunhes Chron. The lion affinities of large pantherine felids from 37 38 The rich paleontological characteristics of this forma- the Pleistocene of Eurasia and North America had been 38 39 tion permit its subdivision into heterochronous fossilif- clarified by numerous morphological studies and sup- 39 40 erous layers. However, in the studied section the Sagar- ported by the molecular data of Burger et al. (2004) and 40 41 lyk Fm is overlain by only basal layers of the Kedrovka Barnett et al. (2009). 41 42 Fm, which are reversely magnetized, and, consequently, Consequently, following the concept of Vereshchagin 42 43 the accumulation of the formation started in the termi- (1971) we grade the lion group of pantherine cats up to 43 44 nal Matuyama Chron (Fig. 2). This part of the formation the rank of their own subgenus Leo Brehm, 1829. With- 44 45 contains significantly younger fauna compared to the in this subgenus, according to the latest morphologi- 45 46 Sagarlyk Fm; namely, Mammuthus trogontherii, Equus cal and molecular record (Sotnikova & Nikolskiy 2006; 46 47 mosbachensis, Rangifer sp., Bos sp. and large ‘priscoid’ Barnett et al. 2009), we include 4 taxa; that is, the living 47 48 Bison. Panthera leo (Linnaeus, 1758) (Africa and South Asia) 48 49 As for fossilization, the bones of Kedrovka Fm differ and Pleistocene P. (L.) spelaea (Eurasia and Beringia), 49 50 substantially from Sagarlyk bones, having lower degree P. (L.) fossilis (Eurasia), and P. (L.) atrox distributed in 50 51 of mineralization and lighter color. This faunal assem- North America south of the Late Pleistocene ice sheet. 51

1 Description nearly equal in size, with Lr/Lr = 13.1 and 13.2 mm, re- 1 2 spectively. The m1 crown is supported by a large anteri- 2 3 The fragment of mandible IGM-519 has a strong and or root with Lr/root width directly below a tooth crown 3 4 unusually massive horizontal body, with the greatest (Wr) = 20.0/13.5 mm and a smaller posterior root with 4 5 thickness below p4 (thickness [T] = 32.0 mm) and with Lr/Wr = 10.0/9.0 mm. The lingual protrusion of the in- 5 6 an equal-in-size depth in front of p3 and behind m1. An ner part of the posterior root of m1 beyond the lingual 6 7 almost straight mandibular ventral margin below p3-m1 margin of anterior root is also seen on the lower carnas- 7 8 is also characteristic of this specimen. sial in the Siberian form. 8 9 In buccal view a deep and distinctly outlined masse- 9 10 teric fossa reaches forward to the level of the space be- Comparison and species identification of the 10 11 tween 2 roots of m1. The anterior part of the masseteric Siberian form 11 12 fossa is dorso–ventrally narrower than in P. spelaea and 12 The resemblance of the Siberian specimen to the 13 P. leo. Its ventral margin only slightly crosses the mid- 13 group of the largest fossil pantherine cats is beyond 14 line of the mandibular body in a dorso–ventral direction. 14 question but its species identification requires spe- 15 The basin between the anterior border of the masseter- 15 cial discussion. The most complete diagnoses of Euro- 16 ic fossa and its deepest part, which is usually shallow in 16 pean fossil lions P. fossilis and P. spelaea are reported 17 mandibles of P. spelaea and P. leo, is very deep in the 17 in Barycka (2008). The data of many authors on crani- 18 Siberian specimen. The anterior mental foramen is bro- 18 al and dental characteristics of the taxa are summarized 19 ken, and the posterior one is doubled and is situated be- 19 20 low the anterior root of p3 and the middle of the c-p3 20 21 diastema. The greatest diameter of the doubled foramen 21 22 is 15.5 mm, whereas each of the foramens has the great- 22 23 est and smallest diameters of 7.1/3.5 mm and 11.0/7.7 23 24 mm, respectively. Like the ventral margin of the anteri- 24 25 or part of the masseteric fossa, the posterior mental fo- 25 26 ramen is located near the horizontal midline of the man- 26 27 dibular body (Fig. 3a). 27 28 In lingual view the posterior border of the symphyse- 28 29 al surface is nearly vertically oriented. Its most posteri- 29 30 orly projecting part does not reach the level of the mid- 30 31 dle part of the diastema between canine and p3 (Fig. 31 32 3b). 32 33 In occlusal view, the space for incisors is moderate- 33 34 ly wide; the postcanine diastema is long, with length 34 35 (L) of approximately 28.5 mm. According to the alveo- 35 36 lar structure, the lower canine had a large size and a ver- 36 37 tically-oriented root. Its estimated length and width are 37 38 approximately 31.0 and 21.0 mm. Judging by the alveo- 38 39 li, the cheek-tooth series slightly arches buccally. In the 39 40 tooth row, premolars occupy a clearly more labial posi- 40 41 tion in relation to m1, so the lingual margin of p4 roots 41 42 is displaced at 6.7 mm buccally relative to the lingual 42 43 border of the m1 root. There is also a relatively long di- 43 44 astema (L = 3.7 mm) between p4 and m1 roots. There 44 45 is no any overlap or crowding of the cheek teeth in the 45 46 mandible of Siberian specimen. The alveolar length of 46 47 p3-m1 is very large (Fig. 3c, Table 1). 47 48 The anterior root of p3 (root length directly below 48 49 a tooth crown [Lr] = 10.0 mm) is smaller than the pos- Figure 3 The mandible of Panthera fossilis (IGM-519). (a) 49 50 terior root (Lr = 12.2 mm), whereas the roots of p4 are lateral view; (b) occlusal view; (c) lingual view. Scale bar = 50 51 50 mm. 51

1 in these diagnoses. Unfortunately, the mandibular fea- shows, tigers and jaguars more often retain their man- 1 2 tures are almost missing in them and the species and dible shape, with a rectangular outline of the horizontal 2 3 subspecies differentiation of large lion-like felids from body between p3 and m1 (Christiansen & Harris 2009). 3 4 the Pleistocene of Europe is mainly based on morpho- The similar characters to a greater or lesser degree are 4 5 metric analysis of dental characters (Schütt 1969; Hem- demonstrated by fossil Pleistocene lions of Europe and 5 6 mer 1974; Schütt & Hemmer 1978). The lack of teeth in North America (Fig. 4b,d,f,i). For instance, according 6 7 the Siberian specimen forced us to conduct a more de- to Christiansen and Harris (2009, p. 939), a rectangu- 7 8 tailed analysis of the structure of the mandibular body lar outline of the horizontal corpus of the mandible is 8 9 of large pantherine cats and to recognize additional fea- among the general characteristics of the American fos- 9 10 tures characteristic of the fossil lions group. sil lion Panthera atrox. A rather straight ventral margin 10 11 One of the major characters that distinguish liv- of the mandibular corpus below the cheek teeth row is 11 12 ing lions from tigers and jaguars is the form of mandi- also common for the group of the fossil European lions 12 13 ble (Dawkins & Sanfort 1868; Merriam & Stock 1932; (Barycka 2008). 13 14 Christiansen & Harris 2009). A mandible with slightly 14 15 This similar structure of mandible as a whole is char- 15 convex ventral margin and anteriorly tapering horizon- acteristic of the most ancient Villafranchian panther- 16 tal body is characteristic of living lions. Conversely, ac- 16 ine cats, such as Panthera toscana (Schaub, 1949), and 17 cording to Hemmer et al. (2001, 2010), the mandibles 17 Panthera palaeosinensis (Zdansky, 1924), as well as of 18 of the Pleistocene and living jaguars demonstrate a less- 18 the oldest known member of the Leo subgenus, Panthera 19 er depth of the horizontal corpus behind the molar than 19 sp. from the Olduvai upper Bed-II in Africa (Fig. 4g). 20 in the diastema in front of p3. However, as the analysis 20 21 21 22 22 23 23 24 24 25 25 Table 1 Comparative measurements of mandibles and lower dents of Panthera (Leo) fossilis, related Eurasian forms, and the largest 26 26 specimens of Panthera (Leo) atrox 27 27 28 Measurements Early–Middle Pleistocene Late Pleistocene 28 29 (mm) Asia Europe North America 29

30 Panthera (Leo) fossilis P. (L) atrox 30 31 1 2 3 4 5 6 7 8 9 10 11 12 31 32 Length: 32 33 33 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 34 34 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 35 35 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 36 36 37 Depth: 37 38 anteriorly/at p3 62.0 62.0 54.6 — 58.0 56.0 — /53.0 — 41.5–55.0 — — 38 39 anteriorly/at p4 59.7 — — — — — — — 57.4 — /60.7 /59.7 39 40 behind/at m1 62.0 59.0 /51.5 — 60.0 — 67.0 58.0 61.0 — 40 41 Thickness: 41 42 anteriorly to p3 31.0 — 30.8 — 26.9 — — — — — — — 42 43 at p4 32.0 27.3 — — — — — — — — — — 43 44 behind/at m1 /30.0 /26.1 — — 31.6 — — — — — 29.2 36.9 44 45 1 Panthera fossilis: IGM-519, Kuznetsk Basin, Russia (this paper); 2 Panthera sp., Kurtak archeological area, Russia; 3 Panthera 45 46 youngi, Zhoukoudian 1, China; 4 Panthera leo or ?fossilis, NHM M6165, Pakefield, Great Britain; 5 Panthera spelaea or ?fossi- 46 47 lis, K.14.6, Aze cave, France; 6 Panthera fossilis, KP136, Petralona, Greece; 7–8 Panthera fossilis, no. 1, no 2, Mosbach, Germa- 47 48 ny; 9 Panthera fossilis, ISEZ MF/320/6803, Wierzchowska Górna, Poland; 10 Panthera fossilis, Cromer–Mindel, Europe; 11–12 48 49 Panthera atrox, 2901-3, UC14001, RLB, North America. Measurements from: Ovodov and Tarasov (2009); Pei (1934); Lewis et 49 50 al. (2010); Baryshnikov and Petrova (2008); Argant (1988); Kurtén and Poulianos (1977); von Reichenau (1906); Barycka (2008); 50 51 Merriam and Stock (1932). †Alveolar length. ‡Measurements are taken on the image. Dashes indicate missing data. 51

1 1 2 2 3 3 4 4 5 5 6 6 7 7 8 8 9 9 10 10 11 11 12 12 13 13 14 14 15 15 16 16 17 17 18 18 19 19 20 20 21 21 22 22 23 23 24 24 25 25 26 26 27 27 28 28 29 29 30 30 31 31 32 32 33 33 34 34 35 35 36 36 37 37 38 38 39 39 40 40 41 41 42 42 43 43 44 44 45 45 Figure 4 Mandibles of selected members of Pleistocene fossil lions group from Eurasia and North America. Scale bar = 50 mm. (a) 46 Panthera fossilis Kuznetsk Basin, (IGM-519); (b) P. fossilis from Solymár, Hungary (Hankó & Korsós 2007, p. 42, fig. 2); (c) Pan- 46 47 thera (Leo) sp. from Kurtak archeological area, Russia (Ovodov & Tarasov 2009, p. 90, fig. 1a); (d) P. fossilis, KP136, Petralona, 47 48 Greece (Kurtén & Poulianos 1977, p. 113), Figure reversed; (e) P. youngi, Zhoukoudian 1, China (Pei 1934, table 23, fig. 1c); (f) 48 49 P. fossilis, Mosbach type locality, Germany (Hemmer 1974, table 12); (g) Panthera (Leo) sp., Olduvai upper Bed II, Africa (Petter 49 50 1973, table 8, fig. 1); (h) P. spelaea, Indigirka Basin, Russia (Ovodov & Tarasov 2009, p. 90, fig. 1b); (i) P. atrox, LACMMC 2900- 50 51 3, RLB, North America (Christiansen & Harris 2009, p. 939, fig. 4), figure reversed. 51

1 Therefore, it would be more logical to consider the com- apomorphic dental features with the lion group of felids. 1 2 bination of characters listed above as a primitive state in Other African finds previously assigned to lion-like fe- 2 3 the pantherine evolution, and not as features, as put for- lids and known from sediments dated at 1.87–1.12 Ma 3 4 ward by Christiansen and Harris (2009), that confirm can be classified only as the ancestral forms belonging 4 5 the relationship between Panthera atrox and jaguars. to the lion stem group (Hemmer 2011). 5 6 Consequently, an almost square shape of the central part Despite the fact that most researchers consider the 6 7 of the mandible of Siberian form demonstrates its plesi- Olduvai specimen to be a possible ancestor of Panthera 7 8 omorphic nature and, together with other features listed (Leo), little information is available on the morpholo- 8 9 below, permits its attribution to the group of fossil lions. gy of this mandible. Petter (1973) notes a straight ven- 9 10 The alveolar length of p3, p4 and m1 of the Siberi- tral margin and a massive mandibular body, as well as 10 11 an lion fall well within the range observed for Eurasian incisors, with narrow and long cross-section, differing 11 12 specimens of P. spelaea and P. fossilis, but the length in shape from those of P. leo. Vereshchagin (1971) point 12 13 of the cheek teeth row exceeds all measurements cit- out the strong development of the posterior basal cin- 13 14 ed in the literature for the largest fossil lions. The ra- gulum on р4 and a lion type of wide coronoid process 14 15 tio Lm1/Lp4 = 101.3% of the studied form is within the in specimen no. 1273. Hemmer (2011) mentions a lion 15 16 range of those values (97.5%–105.5%) given by Schütt condition of P3 in this specimen and showed the simi- 16 17 and Hemmer (1978) for P. fossilis from Mosbach and larity in size and dental characters of the Olduvai man- 17 18 Mauer. The estimated size of the lower canin of IGM- dible to that of P. fossilis from the Mosbach type locali- 18 19 519 (L/W = nearly 31/21) is also similar to the range ty in Germany (Hemmer 1974, tab. X1). 19 20 20 given by Argant (2010, fig. 29) for the European speci- According to our analysis, the Olduvai specimen 21 21 mens of P. fossilis. In addition, the Siberian lion closely (Fig. 4g) has a straight ventral margin of horizontal ra- 22 22 matches P. fossilis in the great thickness of the mandib- mus and equal depth of the latter in front of p3 and be- 23 23 ular body, in the deep, distinctly outlined and highly-po- hind m1, as well as a long p4 with relatively short main 24 24 sitioned anterior portion of the masseteric fossa (Fig. cusp, strong anterior and posterior cusps and developed 25 25 4b,d,f), in the ratio of p4/m1 length, and in the large lin- posterior basal cingulum. The large anterior part of p4 26 26 ear size of the p4 anterior root. This set of features dis- supported by a strong root is also among these features. 27 27 tinguishes the Siberian specimen from P. spelaea but These characters, both plesiomorphic (mandubular out- 28 28 shows its similarity to P. atrox (Fig. 4h,i). However, the line) and apomorphic (reduced p3 and complicated p4), 29 29 American form, as shown below, is a more advanced the African form shares with P. fossilis. Thus, the fea- 30 30 member of the P. fossilis–P. atrox lineage. tures revealed in P. fossilis and the Olduvai lion indicate 31 31 their close relationship. This confirms the view of previ- 32 32 ous researchers that Panthera sp. from the Olduvai up- 33 DISCUSSION 33 per Bed-II is the ancestor of Eurasian fossil lions. 34 Origin and European occurrence of Panthera 34 35 Lion-like pantherine felids dispersed to Europe from 35 36 fossilis Africa in the first half of the Middle Pleistocene. Finds 36 37 of the oldest European lions dated in the time span 37 The early history of lion-sized pantherine cats is as- from the Cromer Complex (early Middle Pleistocene, 38 sociated with the African continent where a find of Pan- 38 39 about 0.78 Ma) to the Holstein interglacial (mid-Mid- 39 thera sp. with features of jaguars, leopards and, to a dle Pleistocene, about 0.45 Ma) are rare. Nowadays all 40 lesser extent, lions is known from the Pliocene fau- 40 41 of them are referred to P. fossilis that was described by 41 na (approximately 3.5 Ma) of the Laetoli site (Hemmer von Reichenau (1906) based on well-preserved fossils 42 2011). The specimen from Laetoli was commonly con- 42 43 from Mosbach and Mauer sands. Other most import- 43 sidered an ancestor of the lion group of pantherine felids ant findings of P. fossilis of the first half of the Middle 44 (Turner & Antón 1997; Werdelin & Lewis 2005), but 44 45 Pleistocene are also known from the localities West- 45 Hemmer et al. (2001) consider it to be a stem species bury-sub-Mendip in Great Britain, Isernia la Pine- 46 for the whole group of felids mentioned above. Сurrent- 46 47 ta in Italy, Château in France, Vértesszólós in Hungary, 47 ly, the mandible No. 1273 from the Olduvai upper Bed- and from Cromerian beds of Petralona Cave in Greece 48 II dated at approximately 1.4–1.2 Ma is believed to be 48 49 (Freudenberg 1914; Dietrich 1968; Schütt 1969; Kurtén 49 the most ancient form of fossil lions. According to Hem- & Poulianos 1977; Sala 1990; Argant et al. 2007; Hankó 50 mer et al. (2010), only this specimen actually shares 50 51 & Korsós 2007; Hemmer 2011). 51

1 The earliest occurrence of P. fossilis in the European scenario by Sabol (2011a), according to which in the 1 2 mainland was established on the basis of a single upper Middle Pleistocene (approximately in MIS 6) cave li- 2 3 carnassial derived from the Isernia la Pineta site in Ita- ons were separated from hypothetical ancient local pop- 3 4 ly, and was dated at about 0.61 Ma (Sala 1990; Hemmer ulations of lion-like felids that penetrated into the Al- 4 5 2011). However, to date, the oldest known finding of a pine regions. Initially P. spelaea became widespread in 5 6 lion-like cat in Europe is, apparently, a specimen from the mountain areas of Europe and only in the Late Pleis- 6 7 Pakefield in Suffolk, Great Britain, dated within MIS 17 tocene dispersed throughout Eurasia. According to this 7 8 around 0.68 Ma, or even MIS 19 around 0.75 Ma (Lew- scenario, relict populations of P. fossilis could have been 8 9 is et al. 2010). A poorly presented mandibular frag- retained on the European plains by the last glacial time; 9 10 ment from Pakefield was described as P. leo by Lewis et that is, until the time of global distribution of cave lions 10 11 al. (2010), but a distinctly labial position of premolars in Eurasia (Sabol 2011a). 11 12 in relation to the molar, spaced p4 and m1, as well as a Such an interpretation of relationships between P. 12 13 massive mandibular body beneath the diastema c1-p3 fossilis and P. spelaea suggests that ‘intermediate’ forms 13 14 and large dental size as a whole do not eliminate a pos- could be advanced members of the P. fossilis lineage 14 15 sible assignment of specimen M-6165 from Pakefield to rather than being transitional from one species to anoth- 15 16 P. fossilis. er. However, to confirm this hypothesis a detailed mor- 16 17 Recent research indicated that major findings of P. phological revision of the well-stratified latest Middle– 17 18 fossilis and similar forms were obtained from the Middle earliest Late Pleistocene finds of fossil lions is required. 18 19 Pleistocene sediments of Europe. Large lion-like felids 19 Fossil Pleistocene pantherine felids of Asian 20 appeared not earlier than MIS 19–17 (0.75–0.68 Ma) 20 21 and were relatively rare forms in the early Middle Pleis- mainland 21 22 tocene faunas, but by the end of the Middle Pleistocene 22 According to Qiu (2006), the oldest Asian remains of 23 they occupied a prominent place in European faunal as- 23 Panthera sp. dated at 3.6–2.6 Ma were found in China 24 semblages (for a summary see Argant et al. 2007; Bary- 24 in the Late Pliocene deposits of the Yushe Basin; how- 25 cka 2008; Marciszak & Stefaniak 2010; Sabol 2011a). 25 ever, this material was not described in detail and its at- 26 At the same time, it should be noted that among the Eu- 26 tribution to the Panthera genus is not reliably justified. 27 ropean Pleistocene specimens there are forms of lions 27 Further development of the genus in this region can 28 with the so-called ‘transitional’ or ‘intermediate’ fea- 28 29 tures (Hemmer 1974; Schütt & Hemmer 1978; Argant be traced by finds of felids close in size to jaguar-like 29 30 et al. 2007). These forms occurred in the latest Middle– members of the Panthera genus. These finds were re- 30 31 earliest Late Pleistocene of Europe. For instance, among ported within faunal assemblages of Henan (Lok. 39), 31 32 them are atypical forms with mixed characters of P. fos- Yangguo and Longdan as P. palaeosinensis (Zdansky 32 33 silis and P. spelaea from the Austrian site Repolusthöhle 1924; Qiu 2006). Faunas of these sites are correlated 33 34 and abundant lion remains from the Biśnik Cave (Schütt with mammalian unit NCMQ1 of the Chinese Quaterna- 34 35 & Hemmer 1978; Marciszak & Stefaniak 2010; Hem- ry continental scale and are dated within 2.6–1.3 Ma (Qiu 35 36 mer 2011). 2006). 36 37 The relationships between P. spelaea and P. fossi- Panthera palaeosinensis was initially regarded as the 37 38 lis have not been fully elucidated. Most scientists con- ancestral tiger (Hemmer 1967). Subsequently, it was 38 39 sider them as chronosubspecies of P. spelaea (Baryc- considered as the most primitive member of the lion or 39 40 ka 2008; Argant 2010; Marciszak & Stefaniak 2010) or leopard clade and even as an ancestor of the genus Pan- 40 41 P. leo (Schütt & Hemmer 1978). Meanwhile, Sotniko- thera (Mazák 2010). Analysis of P. palaeosinensis fea- 41 42 va and Nikolsky (2006), based on the cranial characters, tures is beyond the scope of the present work, but it 42 43 believe that P. spelaea and P. fossilis are different spe- should be noted that its mandibular morphology is clos- 43 44 cies. The latter record agrees well with the results of cla- er to that of the Early–Middle Pleistocene jaguar-like 44 45 distic analysis of pantherine felids from the Pleistocene felids than to the lion group of pantherine cats. Conse- 45 46 of Europe conducted by Hankó and Korsós (2007). The quently, the analysis of fossil data leaves no doubt that 46 47 result of the study by Hankó and Korsós gave reason to the origin of the lion group of felids is associated with 47 48 believe that P. spelaea was not a direct descendant of the African continent, and the assumption of Mazák 48 49 P. fossilis, but represents a separate more advanced lin- (2010) about their Asian roots is not supported by factu- 49 50 eage. These conclusions agree to some extent with the al data. 50 51 51

1 Information on fossil lions in China is confined to the masseteric fossa, and p4 with posterior cingular basin 1 2 finding of a lion-sized mandible in Locality 1 of Zhouk- bearing a ridge-like cingular cusp. However, unlike the 2 3 oudian (ZKD) (Pei 1934). Views on the age of the conditions observed in P. fossilis, the Kurtak specimen 3 4 ZKD-1 fauna are quite variable but modern research- has p4 significantly shorter than m1, as also is seen in P. 4 5 ers support an age estimate of the main fossiliferous youngi from ZKD-1. 5 6 layers of ZKD 1 within the time span of 0.69–0.42 Ma The analysis of other findings presumably associat- 6 7 (Qiu 2006). Pei (1934) created a new species P. youngi ed with the Middle Pleistocene sediments in the terri- 7 8 for the ZKD-1 pantherine felid and noted its affinities to tory eastward from the Ural Mountains does not con- 8 9 both the cave lion and the American P. atrox. Hemmer firm their similarity with P. fossilis. For instance, Sabol 9 10 (1974) has also supported the attribution of this form (2011a), referring to Vereshchagin (1971) and Sotniko- 10 11 to the lion group of fossil felids. Moreover, Harrington va (2006), reports finds of fossil lions in the Middle 11 12 (1969) believed that the American fossil lion, the Eur- Pleistocene of Siberia, whereas Sotnikova (2006) only 12 13 asian cave lion and P. youngi are conspecific. Exam- notes the occurrence of jaguar-like members of pan- 13 14 ination of the pictures figured by Pei (1934, fig. 40b, therine cats in Transbaikalia. As for the findings listed 14 15 pl. XXIII-1a–c) showed that the mandible from ZKD- in Vereshchagin (1971), a well-stratified material is de- 15 16 1, similarly to most specimens of P. fossilis, have mas- rived only from Late Paleolithic sites, whereas the ages 16 17 sive mandidular body with a straight ventral margin. of older specimens require a detailed examination. Sev- 17 18 Its fourth premolar also resembles P. fossilis in bear- eral mandibles of Panthera spelaea were described from 18 19 ing a strong anterior cusp supported by a large anterior the Adycha (MM6880) and Kolyma (ZIN 29405, GIN 19 20 root, and a well-developed posterior cingular area lack- 833-104) basins from supposedly Middle Pleistocene 20 21 ing a distinct cingular cusp. At the same time, a relative- sediments (Baryshnikov & Boeskorov 2001). Howev- 21 22 ly short p4 in P. youngi indicates its more derived posi- er, according to the authors (pp. 8–9), this material was 22 23 tion compared to most European specimens of P. fossilis found on the beach and its morphology is typical for P. 23 24 (Table 1, Fig. 4e). spelaea. 24 25 25 As noted above, there are few reports indicating the Thus, apart from the mandible from Kuznetsk Basin, 26 26 Early or Middle Pleistocene occurrence of fossil lions distribution of lions with features resembling P. fossilis 27 27 in Asian Russia. A skull of a giant pantherine cat (with in the Pleistocene of Asia is confirmed by the presence 28 28 condylobasal length [CBL] = 422 mm and zygomatic of P. youngi in ZKD-1 and the findings of very large 29 29 breadth [ZB] = 312 mm) was found together with Hys- forms in the Kurtak archeological area and in the Mid- 30 30 trix vinogradovi Argyropolo, 1941 and Ursus thibetanus dle Ural Mountains (Mokhnevskaya Cave). Compared 31 31 permjak Baryshnikov, 2001 in the Mokhnevskaya Cave, to the European finds, the Asian specimens are very few. 32 32 Middle Ural Mountains, in the deposits correlated with Nevertheless, keeping in mind the earliest finding of a 33 33 a warm stage of the late Middle Pleistocene (MIS 7) or fossil lion in Kuznetsk Basin, we can assume that the 34 34 the last interglacial (MIS 5e) (Baryshnikov 2001, 2003). expansion of the oldest lions from Africa to Eurasia via 35 35 Unfortunately, the skull was not available for detailed the Asian continent could also take place. New infor- 36 36 study, as it is part of a private collection. mation on the Early Pleistocene occurrence of P. fossilis 37 37 A mandible of a very large pantherine felid with in central Siberia located quite close to the territory of 38 38 mandibular depth of 62.0 mm and Lm1 = 36.0 mm was Beringia also supports the idea that this group of felids 39 39 found on the beach in the area of the Berezhekovo local- could invade America much earlier than the cave lion P. 40 40 ity, on the left bank of the Yenisei River, Kurtak archeo- spelaea. 41 41 logical area, near Krasnoyarsk (Fig. 1). This find can be 42 American Pleistocene lion Panthera atrox and its 42 presumably associated with the Middle Pleistocene sed- 43 43 iments exposed in the Berezhekovo section (Krukover relationship with Eurasian pantherine felids 44 44 & Chekha 1999). The Kurtak specimen was described 45 It has long been thought that all the Pleistocene li- 45 as Panthera sp. (Ovodov & Tarasov 2009). It is similar 46 on-like pantherine felids of North America belong to 1 46 in size to the largest members of the lion group of the 47 species, P. atrox (Whitmore & Forster 1967; Harington 47 Pleistocene felids (Table 1, Fig. 4c). This mandible mor- 48 1969). Subsequently, Vereshchagin (1971) assumed that 48 phologically agrees with that of P. fossilis in having a 49 Beringian lions from north-eastern Siberia, Alaska and 49 strong and massive horizontal ramus with straight low- 50 the Yukon territory are P. spelaea rather than P. atrox. 50 er border, narrow and well-outlined anterior part of the 51 Kurtén (1985) placed the Beringian (Alaska/Yukon) li- 51

1 ons in P. spelaea and the southern North American form Middle Pleistocene in Europe and the first occurrence 1 2 in P. atrox. This opinion was later confirmed by mor- of fossil lions in America (Fairbanks, Alaska) during the 2 3 phological and molecular records (Baryshnikov & Boe- Illinoian glaciation. Our data from Siberia fill, to some 3 4 skorov 2001; Sotnikova & Nikolsky 2006; Barnett et al. extent, a geographic gap in the Eurasian history of the 4 5 2009). To date, only finds of pantherine cats south of the most ancient group of lions. 5 6 Late Pleistocene ice sheets are regarded as P. atrox (Fig. Examination of the Siberian finding revealed a close 6 7 5). affinity between Eurasian samples of P. fossilis and the 7 8 Despite the fact that some authors point to the sim- population of P. atrox from the collection of Rancho 8 9 ilarity of P. atrox to jaguars and even to tigers (sum- La Brea, USA, described by Merriam and Stock (1932) 9 10 marized in Christiansen & Harris 2009), most research (RLB). Judging from the descriptions by Merriam and 10 11 based on morphological data infers their attribution to Stock (1932) and Christiansen and Harris (2009), we 11 12 the group of fossil lions (Harington 1969; Vereshchagin can summarize that the American lion closely resem- 12 13 1971; Hemmer 1974; Kurtén & Anderson 1980; Barysh- bles P. fossilis in the nearly straight ventral mandibu- 13 14 nikov & Boeskorov 2001; Sotnikova & Nikolsky 2006). lar outline, the narrow and very deep anterior part of the 14 15 Nevertheless, Christiansen and Harris (2009) attempt- masseteric fossa, the rectangular profile of the horizon- 15 16 ed to prove that P. atrox was, indeed, a species separate tal ramus, the large p4 with well-developed addition- 16 17 from the other great cats and descended from an early al cusps and basal cingulum, as well as in the subequal 17 18 jaguar. The situation has become clear after recent stud- size of the p4 roots. Among these features, we consid- 18 19 ies of mitochondrial DNA of a living and Late Pleisto- er the morphology of the mandibular body as primitive. 19 20 cene lions, P. spelaea and P. atrox, undertaken by Bar- Of the derived features uniting these 2 taxa, we note, on 20 21 nett et al. (2009). The authors completely disprove the average, a larger general size and more massive mandi- 21 22 postulated link between P. atrox and jaguar and show ble than in other lion-like felids and a slightly more re- 22 23 that the ancient DNA sequences of the studied Pleisto- duced area of the anterior cusp on p4 compared to p4 23 24 cene forms strictly group with living lions. of the ancestral Panthera sp. from the Olduvai upper 24 25 Bed-II. In addition, Sotnikova and Nikolskiy (2006) list 25 Although the inclusion of P. atrox as a separate spe- 26 some cranial features of P. atrox that characterize the 26 cies in the lion group of felids is currently beyond ques- 27 American lion as a very advanced form. Such characters 27 tion, its origin and relationships to other fossil lions are 28 28 still the subject of debate. Most authors believe that the 29 29 American lion descended from the ancient (Yamagu- 30 30 chi et al. 2004) or Beringian (Barnett et al. 2009; Hem- 31 31 mer 2011) populations of cave lions; others consider 32 32 it as being derived from the Chinese P. youngi (Har- 33 33 ington 1969) or from a hypothetical member of the 34 34 Asian lion-like group of felids Panthera ‘leo’ spp. (Sab- 35 35 ol 2011a). 36 36 37 However, results of the present study suggest a dif- 37 38 ferent scenario for the evolution of the American lion. 38 39 As is known, the largest forms of P. atrox were derived 39 40 from the Late Pleistocene asphalt beds of the Rancho 40 41 La Brea in the southern part of North America. On aver- 41 42 age, these lions are larger than P. spelaea, and their size 42 43 is comparable to that of P. fossilis. Despite the appar- 43 44 ent similarity in size of P. atrox and P. fossilis, morpho- 44 45 logical comparison of these taxa has never been done 45 46 and their relationship has not been discussed. Exclusion 46 47 of P. fossilis from comparative analysis has likely re- 47 48 sulted from the lack of its findings in the territories of 48 Central Asia and Siberia intermediate between Europe 49 Figure 5 Late Pleistocene occurrence of Panthera spelaea and 49 and America, and also from the fact that there is a long 50 Panthera atrox in North America (modified after Barnett et al. 50 51 break between the existence of P. fossilis in the early 2009, fig. 1). 51

1 are also observed in mandibles of P. atrox. The derived nancial support from the Russian Foundation for Basic 1 2 conditions are represented by the development of a long Research, project 12-05-00-904a. 2 3 and ventrally-deflected angular process and by a poste- 3 4 rior expansion of the coronoid process (Fig. 4i). REFERENCES 4 5 Thus, we can conclude that P. atrox evolved on the 5 6 morphological basis of the most ancient members of the Alexeeva EV (1980). Pleistocene Mammals of South- 6 7 Eurasian lion group but it has developed its own ad- east of Western Siberia. Nauka, Moscow. (In Rus- 7 8 vanced features. Its ancestor was probably P. fossilis, sian.) 8 9 which dispersed to America from Siberia (via the Bering Argant A (1988). Etude de l’exemplaire de Panthera 9 10 Land Bridge) earlier than P. spelaea did. The latter ap- spelaea (Goldfuss, 1810) (Mammalia, Carnivora. Fe- 10 11 peared in American Beringia at the beginning of the last lidae) du gisement Pleistocene moyen recent de la 11 12 glacial (Hemmer 1974; Schütt & Hemmer 1978; Barnett grotte d’Aze (Saône-et-Loire). Revue de Paléobiolo- 12 13 et al. 2009), whereas the earliest migration wave nearly gie 7, 449–66. 13 14 14 did not leave ancient evidence in the history of panther- Argant A (2010). Carnivores (Canidae, Felidae et Ursi- 15 15 ine cats of North America. dae) de Romain-la-Roche (Doubs, France) Genève. 16 16 It is commonly believed that the oldest fossil lions Revue de Paléobiologie 29, 495–601. 17 17 dispersed to America at the beginning of the Illinoian 18 Argant A, Jeannet M, Argant J, Erbajeva M (2007). The 18 glaciation. However, according to Harington (1969), the 19 big cats of the fossil site Château Breccia Northern 19 species identification of the Illinoian specimen P.( fos- 20 Section (Saône-et-Loire, Burgundy, France): stratig- 20 silis–atrox group or P. spelaea) was not carried out and 21 raphy, palaeoenvironment, ethology and biochrono- 21 its age estimate is also in doubt. Harington (1969, p. 22 logical dating. Courier Forschunginstitut Sencken- 22 1285) wrote: “Although the specimen may represent P. 23 berg 259, 121–40. 23 atrox the evidence is uncertain. The fauna and chronolo- 24 24 gy of the fossiliferous deposits concerned deserve close Barnett R, Shapiro B, Barnes I et al. (2009). Phylogeog- 25 25 study.” raphy of lions (Panthera leo ssp.) reveals three dis- 26 tinct taxa and a Late Pleistocene reduction in genetic 26 In general, American finds, namely, northern P. spe- 27 diversity. Molecular Ecology 18, 1668–77. 27 28 laea and southern P. atrox, are mainly late Late Pleis- 28 tocene in age (Harington 1969; Stuart & Lister 2011). Barycka E (2008). Middle and Late Pleistocene Felidae 29 and Hyaenidae of Poland. Fauna Poloniae–Fauna 29 30 Despite the fact that both species occurred in Ameri- 30 ca almost simultaneously, P. atrox has retained far more Polski. Museum and Istitute of Zoology Polish Acad- 31 emy of Science, Warszawa. 31 32 primitive features than the Late Pleistocene Eurasian 32 Baryshnikov G (2001). The Pleistocene black bear (Ur- 33 and American P. spelaea. Such a combination of charac- 33 34 ters could have been formed in P. atrox owing to a rap- sus thibetanus) from the Urals (Russia). Lynx (Praha), 34 35 id penetration of P. fossilis southward and its subsequent n. s., 2002 32, 33–43. 35 36 isolation in North America south of the Late Pleistocene Baryshnikov GF (2003). Pleistocene small porcupine 36 37 ice sheets. The age of penetration of the most ancient li- from the Ural Mountains, Russia, with note on taxon- 37 38 ons to America is unknown but molecular data indi- omy of Hystrix vinogradovi (Rodentia, Hystricidae). 38 39 cate that the genetic isolation of P. atrox occurred in the Russian Journal of Theriology 2, 43–7. 39 Middle Pleistocene approximately 340 ka (Barnett et al. 40 Baryshnikov G, Boeskorov G (2001). The Pleistocene 40 2009). 41 cave lion Panthera spelaea (Carnivora, Felidae) from 41 42 Yakutia, Russia. Cranium 18, 7–24. 42 43 ACKNOWLEDGMENTS Baryshnikov GF, Petrova EA (2008). Cave lion (Pan- 43 44 44 We gratefully acknowledge Nikolai Ovodov for pro- thera spelaea) from the Pleistocene of Chuvashiya, 45 45 viding additional data on the material from Kurtak ar- European Russia. Russian Journal of Theriology 7, 46 46 chaeological area. We thank Alexey Tesakov for scien- 33–40. 47 47 tific discussion that critically improved the paper. We 48 Burger J, Rosendahl W, Loreille O et al. (2004). Molec- 48 are also indebted to Katya Firsova and Igor Foronov, 49 ular phylogeny of the extinct cave lion Panthera leo 49 who kindly prepared the design of figures for the pur- 50 spelaea. Molecular Phylogenetics and Evolution 30, 50 pose of this study. M. Sotnikova acknowledges the fi- 51 841–9. 51

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