Decreasing size process in the cave (Pleistocene) lion Panthera spelaea (Goldfuss, 1810) evolution – A review

Quaternary International 339-340 (2014) 245e257

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Decreasing size process in the cave (Pleistocene) lion Panthera spelaea (Goldfuss, 1810) evolution e A review Adrian Marciszak a, *, Charles Schouwenburg b, Robert Darga c a

Division of Palaeozoology, Department of Evolutionary Biology and Ecology, Faculty of Biological Sciences, University of Wrocław, Wroclaw, Poland Dorpsstraat 53, 3238BB Zwartewaal, Netherlands c Naturkunde- und Mammut-Museum Siegsdorf, Germany b

a r t i c l e i n f o

a b s t r a c t

Article history: Available online 25 October 2013

This study analyzed size changes in the course of the time among European Middle and Late Pleistocene lions. The general decreasing size as a tendency in cave lion evolution is confirmed. The whole time span of lion presence in Europe was divided into four main periods. The first period covers Middle and late Middle Pleistocene (MIS 19-17 to MIS 7-6) and is characterized by the presence of very large individuals with somewhat primitive morphology. It might be cautiously said that lions in that time were comparatively large throughout the Middle and late Middle Pleistocene. The second period included the last part of the late Middle and the beginning of the Late Pleistocene (MIS 6-5). This period is documented by a mixed presence of large individuals with somewhat primitive features together with smaller, evolutionarily more advanced dentition lions. The third period covers the first part of the last glacial, from MIS 5 to the beginning of MIS 3. For this period, the occurrence of moderate sized specimens with evolutionarily advanced dentition is documented. The last period began with dramatic population decline and genetic variability reduction. It lasted from MIS 3 and 2, and within it size drops sharply. Among moderate sized specimens, very small lions with somewhat lesser posture also appeared. Asian or African affinities as a result of a new migration event for those dwarf lions could not be ruled out. Size decrease showed that Panthera spelaea was a dynamically evolved species. Thus, the size does not seem to be a reliable criterion in determining the age of cave lion findings. The general overview is complicated by the great sexual dimorphism, local evolution and extinction, climatic conditions, and migrations. Size changes in P. spelaea evolution cannot be used as a useful biochronological tool. Ó 2013 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction The majestic posture and great size of the Pleistocene lion have fascinated humans since prehistoric times (Argant et al., 2007). Two forms of the Pleistocene lion were originally distinguished: the Middle Pleistocene Panthera spelaea fossilis (von Reichenau, 1906) and the Late Pleistocene P. spelaea spelaea (Goldfuss, 1810). Since then, the size variation and the possible size decrease within the lion lineage have been subject to debate (Wurm, 1912; Freudenberg, 1914). Some authors maintain that the evolution of the lineage involved a decrease in size (Kurtén, 1960, 1968; Schütt, 1969a; Hemmer and Schütt, 1970; Schütt and Hemmer, 1978; Argant, 1988, 2000, 1991; Guzvica, 1998; Baryshnikov and Boeskorov, * Corresponding author. E-mail addresses: [email protected] (A. Marciszak), c.schouwenburg@ upcmail.nl (C. Schouwenburg), [email protected] (R. Darga). 1040-6182/$ e see front matter Ó 2013 Elsevier Ltd and INQUA. All rights reserved. http://dx.doi.org/10.1016/j.quaint.2013.10.008

2001; Hemmer, 2003, 2004; Bona, 2006; Argant et al., 2007; Hanko, 2007; Barycka, 2008; Baryshnikov and Tsoukala, 2010; Marciszak and Stefaniak, 2010; Argant, 2010a, 2010b; Sotnikova and Foronova, 2012). Others doubt this (Turner, 1997; Nagel et al., 2003; Baryshnikov and Petrova, 2008; Lewis et al., 2010; Hemmer, 2011). The new finds of complete skulls of very large Late Pleistocene lions from eastern and central Europe shed new light on the problem. Records of P. spelaea outside caves are, when compared with cave occurrences, quite rare in central Europe, and the San specimen is the first river find in Poland. From the San River, located in southeeast Poland, some other large, well preserved skulls and long bones have been dredged. They were represented by large herbivores such as wooly mammoth Mammuthus primigenius (Blumenbach, 1799), woolly rhinoceros Coelodonta antiguitatis (Blumenbach, 1807), and auroch Bos primigenius Bojanus, 1827. All paleontological data confirmed the last glacial age of the findings from the San River (Stefaniak pers. comm. 2013).

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2. Methods and materials 2.1. Methods Measurements were taken point to point, with an electronic caliper, to the nearest 0.1 mm. Each value given here is the mean of three measurements, and all measurements are in mm. Osteological and dental terminology and measurements of teeth and bones follow Schmid (1940), Argant (1988, 1991) and Hanko (2007) (Figs. 1e3). As comparative material, the following literature data were used for the morphometric analysis: Dawkins and Sandford 1866, Filhol and Filhol 1871, Hagmann 1899, Boule 1906, Freudenberg 1914, Riabinin 1932, Heller 1953, Del Campana 1954, Pidoplichko 1956, Malez 1963, Rakovec 1964, Terzea 1965, Zapfe 1966, Samson and Kovacs 1967, Dietrich 1968, Schütt 1969a, 1969b, Suire 1970, Vereshchagin 1971, Altuna 1972, Thenius 1972, Hemmer 1974, 1977, Ballesio 1975, Kurtén and Poulianos 1977, Schütt and Hemmer 1978, Ballesio 1980, Clot 1980, Altuna 1981, Kurtén and Poulianos 1981, Bishop 1982, Turner 1982, Toepfer 1983, Nielbock 1987, Argant 1988, Dufour 1989, Alekseeva 1990, Sala 1990, Argant 1991, Gross 1992, Groiss 1992, Fischer 1994, Valensi 1994, Turner 1997, Guzvica 1998, Kleczko 1999, Turner 1999, Baryshnikov and Boeskorov 2001, Garcia 2003, Bona 2006, Sotnikova and Nikolskiy, 2006, Testu 2006, Argant et al., 2007, Hanko 2007, Toskan 2007, Barycka 2008, Diedrich 2008, Ovodov and Zaika 2008, Kempe and Döppes, 2009, Argant 2010a, Baryshnikov and Tsoukala 2010, Diedrich 2011a, 2011b, Sabol 2011a, Argant and Argant 2012 pers. comm., Diedrich 2012, and Diedrich and Rathgeber 2012. 2.2. Material The examined material is an intact skull of an adult, particularly large male lion, lacking the mandible, with both P3 and P4 present (Figs.4e5.). The cranium is kept in the Naturkunde- und MammutMuseum Siegsdorf (Coll. Nr NKM-00664). The specimen was dredged from the San River near Przemysl (SE Poland). Some marks on the sagittal crest and a small injury were detected in the neighbourhood of the right orbit, but none of the injuries seems to have been the reason for the lion’s death. The large size, long and well developed sagittal crest, all sutures and epiphyses fully fused and quite worn teeth indicate that it was fully grown, mature, aged about 7e8 years. Measurements (all in mm): 1-451.0, 2-402.0, 3-375.0, 4-300.0, 5-175.0, 6-230.0, 7-224.0, 8-55.2, 9-138.0, 10-129.2, 11-108.0, 12146.9, 13-93.8, 14-96.5, 15-103.3, 16-322.1, 17-71.1, 18-38.4, 19-90.0, 20-63.0, 21-138.0, 22-92.3, 23-85.6, 24-14.0, 25-44.0, 26-284.9, 2789.0, 28-200.0, 29-62.0, 30-82.4, 31-180.0, 32-46.6, 33-27.1, 3423.5, 35-55.8, 36-36.0, 37-37.4, 38-77.9, 39-176.0; P3: L-30.0/29.8, Lpa-14.6/14.0, Hpa-15.7/16.0, Ba-11.4/11.7, Bp-17.3/17.0; P4: L43.9/44.0, Lpa-15.3/15.1, Hpa-21.5/, Lmt-16.8/16.5, Ba-22.9/22.9, Bp-13.2/13.2. 2.3. Comparative material The examined specimen from the San River was compared in detail with a similar sized skull of the Middle Pleistocene lion P. spelaea fossilis from the Mauer sands. As comparative material, skulls of Middle and Late Pleistocene P. spelaea males were also used. Below, dimensions of those skulls are given, all in mm. Mauer: 1-442.0, 2-393.0, 3-370.0, 4-300.0, 7-225.0, 9-113.0, 12129.0, 13-69.0, 17-70.0, 21-90.0, 22-79.0, 24-15.0, 25-34.0, 29-57.0, 31-150.0, 39-166.6; P3: L-29.3, Lpa-15.7, Hpa-17.2, Ba-13.6, Bp19.2; P4: L-45.1, Lpa-17.5, Hpa-24.4, Lmt-16.8, Ba-21.7, Bp-17.3. Château CHA.1.98.C.3-246: 1-484.7, 9-125.3.

Fig. 1. Scheme of measurements of the lion skull, modified from Argant (1988, 1991, 2010a): 1 e total length (prosthion-acrocranion) (TOT), 2 e condylobasal length (CBL), 3 e basal length (prosthion-basion) (BAS), 4 e bizygomatic breadth (ZYG), 5 e rostrum length, 6 e facial length (prothion-point F), 7 e neurocranium length (point Facrocranion), 8 e nasal breadth, 9 e least breadth at the canine alveoli, 10 e least breadth between infraorbital foramina, 11 e least breadth between orbits (entorbitale entorbital) (IOB), 12 e frontal least breadth (ectorbitaleectorbital) (EOB), 13 e postorbital least breadth (postorbital bar) (POB), 14 e maximal neurocranium breadth, 15 e least breadth at parieto-temporale suturae, 16 e neurocranium length (nasion-basion), 17 e maximal height of zygomatic arch, 18 e minimal height of zygomatic arch, 19 e maximal length of orbit, 20 e maximal height of orbit, 21 e cranial height (acrocranion-basion), 22 e upper tooth row length on alveoles (P2-M1), 23 e upper premolars row length on alveoles (P2eP2), 24 e diastema length (posterior margin of alveole of C1-anterior margin of alveole of P2), 25 e alveole length of C1, 26 e basifacial length (prosthion-synsphenion), 27 e basicranium length (synsphenion-basion), 28 e palatal length, 29 e maximal incisor row breadth at alveolus (I3eI3 B), 30 e minimal breadth between canines at internal margins of canine alveolus (C1eC1 B), 31 e maximal palatal breadth at P4 external alveoles, 32 e minimal breadth between pterygoides, 33 e maximal breadth at cavita gleonidae, 34 e height of foramen magnum, 35 e length of bulla tympanica, 36 e breadth of bulla tympanica, 37 e foramen magnum breadth, 38 e maximal breadth of occipital condyli, 39 e mastoid breadth (MAS).

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Fig. 2. Scheme of measurements of P3, modified from Schmid (1940) and Hanko (2007); A e lateral view, B e occlusal view, Ps e parastyle, Pa e paracone, Ms e metastyle, C e cingulum, mes e mesial side, dist e distal side, ling e lingual side, bucc e buccal side, L e total length of crown, Lpa e paracone length, Hpa e paracone height, Ba e anterior breadth of crown, Bp e posterior breadth of crown.

Fig. 3. Scheme of measurements of P4, modified from Schmid (1940) and Hanko (2007); A e lateral view, B e occlusal view, Pp e preparastyle, Ps e parastyle, Pr e protocone, Pa e paracone, Ms e metastyle, mes e mesial side, dist e distal side, ling e lingual side, bucc e buccal side, L e total length of crown, Lpa e paracone length, Hpa e paracone height, Lms e metacone þ metastyle length, Ba e anterior breadth of crown, Bp e posterior breadth of crown.

Château CHA.1.06.C.3.431: 1-465.0, 9-114.1. Mokhnevskaya Cave: 1-ca 475.0, 2-422.0, 3-398.0, 7-270.0, 9138.0, 10-127.7, 11-110.0, 14-98.7, 19-73.3, 21-140.0, 23-90.0, 28216.0, 31-138.0; P3: L-32.6, Bp-19.1, P4: L-41.8, Ba-22.9. Crayford LBL 1.4: 1-450.0, 4-300.0. Petralona Cave PEC 90: 1-416.0, 2-372.0, 3-350.0, 5-169.9, 6233.0, 7-207.5, 9-115.2, 10-108.1, 11-89.4, 14-98.7, 19-73.3, 21-132.5, 23-81.5, 28-178.5, 31-145.0; P3: L-27.9, Bp-13.6; P4: L-39.6, Ba23.6. Petralona Cave PEC 64: 4-þ262.0, 7-218.0, 12-125.0, 13-77.0, 1770.0, 39-166.0. Azé Cave K 13.29: 1-417.4, 2-368.6, 3-347.7, 4-283.4, 5-162.5, 6233.8, 7-207.9, 8-62.2, 9-112.3, 10-121.2, 11-93.0, 12-133.9, 13-95.3, 14-111.3, 15-99.6, 16-243.3, 17-72.7, 18-37.0, 19-80.5, 20-50.4, 21126.5, 22-89.3, 23-84.9, 24-7.0, 26-247.5, 27-106.4, 28-181.0, 2949.0, 30-58.8, 31-143.3, 32-53.7, 33-39.2, 34-27.0, 35-54.0, 36-34.0, 37-32.6, 38-74.0, 39-162.5; P3: L-29.9/30.6, Lpa-15.2/15.0, Hpa-

Fig. 4. Late Pleistocene cave lion Panthera spelaea spelaea from San River (dorsal view).

17.0/15.0, Ba-13.1/13.6, Bp-18.8/19.0; P4: L-41.3/41.3, Lpa-15.5/15.5, Hpa-22.6/22.6, Lmt-15.7/15.7, Ba-23.0/22.8, Bp-17.6/17.7. Duvannyi Yar IPBPS 1: 1-359.4, 3-307.0, 5-140.0, 9-105.0, 1176.5, 13-67.1, 29-51.5, 30-57.0, 39-152.0, LP3-24.5, Bp-14.4. Smolensk GIN 1123: 1-360.0, 2-328.0, 3-309.0, 4-238.2, 5-140.0, 9-107.0, 11-75.5, 13-65.5, 29-46.5, 30-62.5, 36-25.8, 38-66.2, 39146.0, LP3-25.0. Chernigov: 1-375.0, 2-355.0, 3-322.0, 4-245.0, 8-63.0, 9-111.0, 10-103.0, 11-78.0, 12-115.0, 13-71.0, 15-77.0, 16-185.0, 17-50.0, 21115.0, 22-80.0, 24-6.0, 28-161.0, 29-55.0, 30-61.0, 31-132.0, 32-54.0, 34-29.0, 35-48.0, 36-35.0, 37-31.0, 38-71.0, 39-1520; P3: L-28.0, Bp-14.5; P4: L-39.0, Ba-18.5. Kondakovka K 1: 1-345.1, 2-326.2, 3-305.0, 4-249.0, 5-140.0, 9103.0, 11-72.0, 13-62.0, 29-47.0, 30-57.5, 36-26.5, 38-69.5, 39-153.0; P3: L-28.0, Bp-16.3; P4: L-40.0, Ba-20.8.

Fig. 5. Late Pleistocene cave lion Panthera spelaea spelaea from San River (ventral view).

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Siegsdorf: 1-384.0, 2-352.5, 3-329.5, 4-250.5, 5-151.0, 6-226.0, 7-189.0, 8-58.0, 9-106.0, 10-103.0, 11-76.8, 12-107.5, 13-64.5, 16223.0, 17-63.0, 18-35.0, 19-70.3, 21-111.0, 23-77.0, 26-238.0, 2797.0, 29-53.0, 30-58.5, 31-147.5, 34-33.0, 35-46.0, 36-27.0, 37-32.5, 38-71.3, 39-152.0; P3: L-28.0, Bp-15.5; P4: L-40.3, Ba-20.0. Arrikrutz: 1-407.5, 3-339.0, 4-284.0, 5-162.0, 6-253.0, 7-201.5, 9-112.0, 10-112.0, 11-102.0, 16-233.0, 21-101.0, 23-84.5, 24-10.0, 26246.0, 27-101.0, 31-151.0, 35-46.0, 36-24.0, 39-160.5; P3: L-28.5, Bp-15.5; P4: L-43.0. Moggaster Cave Mog. 85/7: 1-ca 430.0, 4-ca. 280.0, 9-120.0, 1195.0, 12-134.0, 14-115.0, 19-66.0, 21-124.0, 22-90.0, 23-83.0, 31163.0, 34-20.0, 35-50.0, 36-30.0, 37-31.0, 38-72.0, 39-160.0; P3: L31.8, Lpa-16.4, Hpa-16.4, Ba-13.1, Bp-16.2; P4: L-43.1, Lpa-16.7, Hpa-19.8, Lmt-16.6, Ba-21.9, Bp-16.1. Vence: 1-354.0, 2-320.0, 4-241.0, 7-185.0, 12-111.0, 13-78.0, 1768.0, 39-142.0; P3: L-26.0; P4: L-39.0. Cajarc: 1-393.0, 2-350.0, 4-265.0, 7-205.0, 12-126.0, 13-67.0, 1768.0, 39-155.0; P3: L-26.0; P4: L-37.0. L’Herm Cave: 1-410.0, 2-388.0, 3-350.0, 4-245.0, 5-153.0, 7187.0, 9-110.0, 11-90.0, 12-113.0, 13-62.0, 17-73.0, 29-55.0, 39-145.0. Zoolithen Cave BT 5421: 1-395.0, 2-353.0, 3-335.0, 4-261.0, 5153.9, 6-221.0, 7-196.6, 9-107.4, 10-107.4, 11-88.5, 19-62.1, 21-121.1, 23-77.3, 28-171.2, 31-135.9; P3: L-26.0; P4: L-38.0. Zoolithen Cave Ma. 50947: 1-355.0. Zoolithen Cave MB. 48115: 1-417.0, 3-340.0, 4-264.0, 9-116.0, 23-74.0, 30-147.5, P4: L-42.0. Zoolithen Cave Ma. 50948: 1-402.0, 3-325.0, 4-253.0, 23-79.0; P3: L-28.0; P4: L-40.0. Huttenheim 6816.5.6.73.1: 1-375.0, 9-113.0, 17-71.0; P4: L-32.0. Edingen 6617.1.9.79.2: 1-380.0, 17-72.0; P3: L-26.0. Bottrop MBOT 6349/19a-b: 1-440.0, 17-66.0; P3: L-26.0. Slouper Cave R4406: 1-390.0, 2-350.0, 3-335.0, 4-265.0, 9115.0, 10-111.0, 11-93.0, 28-180.0, 31-150.0, 35-55.0, 36-35.0, 39160.0; P3: L-28.0; P4: L-41.0. Slouper Cave OK 130570: 1-378.0, 17-72.0. Lezetxiki: 1-421.5, 2-382.0, 3-366.0, 4-274.5, 9-107.0, 11-74.5, 12-122.5, 13-88.0, 21-115.0, 28-191.5, 31-169.5, 37-31.2, 38-71.5, 39-165.5; P3: L-31.0, Ba-12.1, Bp-16.1; P4: L-40.8, Bp-22.7. Romain-la-Roche RO.86-R.20-54: 1-400.9, 6-233.0, 7-207.8, 989.4, 10-127.4, 11-76.3, 12-112.6, 13-96.6, 27-105.5, 29-51.6, 32-44.6, 33-33.5, 34-13.5, 35-46.7, 36-27.4, 37-28.7, 38-64.3, 39-155.0. Urs¸ilor Cave SIER-UrsCave 1: 1-350.0. Duvanny Yar YIG 6397: 2-294.0, 3-275.2, 4-212.0, 5-125.2.0, 6143.0, 7-150.0, 9-91.4, 10-91.4, 11-59.1, 13-66.0, 19-66.0, 23-69.3, 28-157.0, 31-123.3, 35-47.0, 36-29.8, 38-64.0, 39-122.0; P3: L-22.9, Bp-10.3; P4: L-37.0, Lmt-13.3, Ba-17.2. Duvanny Yar YIG 3190/1: 1-309.0, 2-295.1, 3-275.4, 4-213.8, 5126.0, 6-148.0, 7-162.0, 9-88.0, 10-88.0, 11-61.0, 13-61.4, 19-61.4, 2195.3, 23-64.3, 28-155.0, 31-120.8, 35-55.8, 36-47.1, 38-60.8, 39125.6; P3: L-25.3, Bp-13.5; P4: L-38.1, Lmt-14.4, Ba-17.3. Bolshoy Lyakhovski Island ZIN 15572: 13-55.6, 21-95.4, 3547.3, 36-34.0, 38-67.9, 39-150.1. Veternica: 18-45.1, 22-88.0; P3: L-29.0; P4: L-41.3. Uzhur: 1-350.0, 2-323.0, 3-300.0, 4-234.0, 6-206.0, 7-165.0, 865.5, 9-109.5, 10-104.0, 11-83.0, 12-116.0, 14-106.0, 17-54.0, 21117.0, 28-170.0, 30-47.5, 31-140.0, 35-45.0, 36-27.0, 38-71.5, 39142.0; P3: L-26.5; P4: L-39.1. Ural: 1-355.0, 2-330.0, 4-244.0; P3: L-25.5; P4: L-37.0. Altai: 1-334.0, 2-310.4, 3-289.4, 4-222.0, 7-171.0, 8-62.0, 9-95.0, 10-96.0, 11-69.0, 12-100.0, 13-61.5, 14-129.0, 21-101.0, 28-158.2, 31130.0, 35-45.0, 36-28.0, 39-137.0; P3: L-24.0, Bp-13.6; P4: L-36.5, Ba-18.2. Kuzntetskij basin: 1-320.0, 2-317.4, 3-298.0, 4-220.0, 8-68.3, 994.0, 10-97.0, 11-73.0, 12-102.0, 13-65.0, 14-121.0, 31-132.5, 35-46.0, 36-30.0, 39-140.0; P3: L-25.0; P4: L-35.0.

3. Results 3.1. Systematical analysis The examined San specimen is characterized by very large size, with dimensions exceeding most of the other European cave lions (Fig.6.). The rostrum is short and wide, with well marked widening at the canines level and at the P2 level. Generally speaking, the facial part is strongly widened (Figs.6e8.). The incisors toothrow is broad. Massive zygomatic arches are widely spaced. The temporal region is compressed, proportionally short and wide. The postorbital bar is broad and the postorbital process is almost exactly in the middle of the temporal region. Frontal profile (in the lateral view) is strongly convex, with well-marked concavity in the medium of frontal part (Fig.9.). Orbits are large and rounded. Mastoid processes are broad and widely spaced. Bullae tympanic are large, more inflated, and strongly convex with large tympanic chamber. P3 is narrow, with well developed anterior and posterior cingulum ridges. The transition between the anterior and posterior part of the crown is well marked. The posterior part of the crown is long and formed well developed flat surface of triangular shape. Parastyle is poorly developed and oriented anterio-lingualy. Paracone is proportionally long, low, and oval. Metastyle is large and high, medially positioned, sharply bordering from posterior cingulum. In occusal view P4 is slend and elongated. Anterior part of the crown is broad, with reduced, lower and more rounded protocone pushed more posterio-lingualy. Preparastyle is well developed. Considerable marked is anterior margin constriction between protocone and parastyle. Parastyle is large, high and more oval, with strongly developed anterio-lateral bulging of the anterior wall and well separated from the paracone by deep and sharply pointed valley. Paracone is low and shorter than metastyle and the notch between paracone and metastyle considerable. Posterior part of the crown is narrow, strongly convex buccally and slightly thickened on the buccal side. Metastyle is longer than the paracone, sharply ended with the blade oriented almost horizontally (Fig.10.).

Fig. 6. Total skull length (TOT) and C1eC1 least breadth (C1eC1 B) at the canines alveoli of the Middle and Late Pleistocene lion Panthera spelaea. Note large size and proportionally narrow stout of Panthera spelaea fossilis from Château and Mauer (for citation see Comparative material).

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Fig. 7. Scheme of the Pleistocene lion skulls (upper view): left e Middle Pleistocene lion Panthera spelaea fossilis from Mauer (according to Wurm, 1912 and Charles Schouwenburg photos, modified), right e Late Pleistocene lion Panthera spelaea spelaea from San River. Note the broader snout and shorter and wider temporal region of the San individual.

In the Mauer skull, the snout is proportionally longer but narrower (Fig.6.). Its shape is almost triangular, without widening of the facial part at the canines level. Incisors are small and incisors toothrow is narrow (Figs.6e8.). Slightly narrower zygomatic arches are less spaced than in the San specimen. The longer and narrower temporal region is less compressed than in the San individual. Sharply marked narrow postorbital bar is pushed fairly forward of the postorbital process (Fig.7.). Frontal profile (in the lateral view) is almost straight, without any crossing between maxillae and frontal region (Fig.9.). Orbits are quite small and rounded. Mastoid region is narrow, with close compressed mastoid processes. Bullae

tympanic are quite small, poorly convex and with small tympanic chambers. Broad P3 has less developed anterior and posterior cingulum ridges. Parastyle is small, but well marked, shifted slightly anteriolingualy, while paracone is proportionally short, rounded and high. Metastyle is large and low, placed slightly more posterio-buccaly and poorly distinctness from posterior cingulum. Posterior part of the crown is very broad, strongly convex of buccal margin in the posterior part, less developed flat surface of triangular shape. P4 in general occlusal view is broad and compact. Well developed, high and triangular protocone is oriented almost exactly vertically to the

Fig. 8. Scheme of the Pleistocene lion skulls (bottom view): left e Middle Pleistocene lion Panthera spelaea fossilis from Mauer (according to Wurm, 1912 and Charles Schouwenburg photos, modified), right e Late Pleistocene lion Panthera spelaea spelaea from San River. Note the broader snout and more inflated bulla tympanic of the San individual.

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The San skull, compared to the Mauer lion, is characterized by a more progressive morphology, typical for the P. s. spelaea form. It differs from the Mauer cranium in a proportionally shorter and broader snout, larger orbits, better developed temporal and mastoid areas as well as less inflated bulla tympanica, broader incisors toothrow and narrower upper teeth with better cutting projection (P3 and P4). All these features indicate a more advanced position of the San specimen (when compared with the Mauer skull) and well fit with Late Pleistocene lion morphology. Although clearly different in morphology, the two specimens (San and Mauer) share features which distinguish them from the modern lion. First is their greater size. The overall cranial size of both individuals considerably exceeds even the record African lion: TOT ¼ 419.0 mm for the South African lion (Hemmer, 1974) and 402.0 mm for the East African lion (Mazak, 1979). Although the monograph of Panthera leo (Haas et al., 2005) lists an enormous skull of the total length of w460.0 mm, Yamaguchi (2005a, 2005b) doubts this information. The V-shaped, deep and round median indentation at the posterior end of the palate at the opening of the choanae, in both cases indicates P. spelaea (instead of W-shaped for the African lion) (Vereshchagin, 1971; Argant, 1988; Argant and Argant, 2011). Fig. 9. Scheme of the Pleistocene lion skulls (lateral view): top e Middle Pleistocene Panthera spelaea fossilis from Mauer (according to Wurm, 1912 and Charles Schouwenburg photos, modified), bottom e Late Pleistocene lion Panthera spelaea spelaea from San River. Note the less developed temporal region, proportionally larger orbits and convex frontal profile of the San individual.

crown. Anterior margin constriction between protocone and parastyle is quite poorly marked. Preparastyle is absent. Parastyle is small, rounded, flat, with well marked anterio-lateral bulging of the anterior wall. Parastyle is strongly associated with the paracone (constriction between parastyle and paracone is negligible). Paracone is high and longer than metastyle and the notch between paracone and metastyle is shallower and not well pronounced. Angle between paracone and metastyle more compressed. Metastyle is shorter than the paracone, with blade oriented somewhat upward. Generally, the posterior part of the crown is broad, strongly thickened buccally and lingually, with a straight buccal margin (Fig.10.).

3.2. Size fluctuation We compiled available dimensions of selected bones (skulls, long bones, calcaneus, metapodials and teeth) and plotted them together. Only the total length of examined skeleton elements has been used and is shown as single dimensions, due to the small number of samples from each locality and the great variety of both Middle and Late Pleistocene lions. We compared dimensions of the Pleistocene lions with a large, sexually and geographically diverse sample of modern P. leo given by Gross (1992). It is clear that in almost all bones of smaller P. spelaea, individuals (females) are comparable in size with recent African male lions, as was found by Lewis et al. (2010). There is well- pronounced sexual dimorphism in both species, where males far exceed dimensions of females. For example, skulls of the male P. spelaea are on average 10 cm longer than female skulls (Gross, 1992; Diedrich, 2007). Considering P. spelaea skull male dimensions, scenario proposed by Argant et al. (2007) of the size decrease with time is well

Fig. 10. Scheme of the Pleistocene lion P4 (lateral view): left e Middle Pleistocene lion Panthera spelaea fossilis from Mauer, right e Late Pleistocene lion Panthera spelaea spelaea from San River. Note presence of the preparastyle, larger, oval paracone, well-marked border between parastyle and paracone, shorter and lower paracone (even if slightly worn), longer metastyle and generally higher as well as less massive crown of the San individual.

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documented in the fossil record (Fig.11.). The total length of P. s. fossilis male skulls from Château (L ¼ 484.7 mm and L ¼ 465.0 mm) and Mauer (L ¼ 442.0 mm) fairly exceeds the length of the later ones. However, the ratio is limited to only three specimens, and the cranial material of early Middle Pleistocene lionesses is even more poorly represented. Only two maxilla pieces (maybe even from one individual) from Bilzingsleben are known. The lioness from Bilzingsleben is characterized by rather medium dimensions and falls in the size range of Late Pleistocene P. spelaea females (Toepfer, 1983; Turner, 1997). The same was found for the female’s mandible from Forest Bed (Baryshnikov and Petrova, 2008). Five late Middle Pleistocene lion crania from Crayford (L ¼ 450.0 mm), Petralona Cave (L ¼ 416.0 mm), Azé Cave (L ¼ 417.4 mm), Moggaster Cave (L ca 430.0 mm) and L’Herm Cave (L ¼ 410.0 mm), also very large, are less impressive. Dated to the transitional period MIS 6-5, lions from Lezetxiki (L ¼ 421.5 mm), Mokhnevskaya Cave (L ca 475.0 mm), Nied zwiedzia Cave (L ¼ 416.4 mm), Arrikrutz (L ¼ 407.5 mm) and Romain-la-Roche (L ¼ 400.9 mm) are still characterized by great size. Two other particularly large individuals from Edingen (L ca 440.0 mm) and San River (L ¼ 451.0 mm), although lacking good stratigraphical position, based on morphology are not older than MIS 6-5. For both specimens, in view on their advanced morphology and great size, the most probable geological age is the first half of the last glacial or less probably, the latest part of MIS 6. From the last glacial, none of 17 male skulls exceed 400.0 mm in total length. The only exception, the largest skull (holotype) from Zoolithen Cave, measured 417.0 mm in total length according to Kempe and Döppes (2009) and only 402.0 mm according to Diedrich (2008, 2012). The Zoolithen Cave is widely regarded as a classical, Late Pleistocene cave bear and cave hyena den, and the detailed stratigraphical position of specimen MB 48115 is unknown. It came from an old, historical collection, for which the lack of stratigraphy is a normal situation. Apart this individual, all lion skulls with more or less good stratigraphical position with Late Pleistocene age have total lengths less than 400.0 mm. Dated at 47,180 BP (Stuart and Lister, 2010), the male from Siegsdorf with TOT ¼ 384.0 mm is a typical, moderate sized specimen from the European last glacial (Gross, 1992). Other skeleton elements including long bones and metapodiales showed on average visible smaller dimensions than Middle Pleistocene lions. Baryshnikov and Tsoukala (2010) and Baryshnikov and

Boeskorov (2001) noted, “In P. l. spelaea from the Late Pleistocene, the basal length is noticeably smaller, not exceeding 340 mm”. In a population, beside normal specimens of average dimensions, record size specimens are noted. The two particularly large, Late Pleistocene individuals from San River and Edingen might be just simply exceptionally large specimens, and their great size should not be regarded as any rule. Complete long bones of the steppe lion are even less numerous than skulls. The few known specimens lie in the upper range of size variability, although single specimens (ulna from Mosbach 2) are above the size of other specimens (Fig.11.). Similarly to skulls, the limited number of Middle Pleistocene lion findings excluded any detail analysis. Much more numerous and better preserved are metapodiales and calcaneus bones. Generally, dimensions of the Middle and the late Middle Pleistocene lions follow the size variability of Late Pleistocene lions, but usually in the uppermost part (Fig.12.). Single specimens of the Middle Pleistocene lion, e. g. giant metatarsal III from Château with an incredible length of w192.0 mm, fairly exceeds the size of other individuals (Argant, 1991). Quite small Middle Pleistocene lionesses, e. g. metatarsal V, represent mostly specimens from localities dated to MIS 11-9, including Aldene and Lunel-Viel (Bonifay 1971). In teeth, this regularity is not so obvious, especially in m1 (Fig.13.). The high number of large m1 in the Late Pleistocene

Fig. 11. Measurements of condylobasal skull length (CBL) and total length of long bones comparisons of the Middle and Late Pleistocene lion Panthera spelaea plotted with recent, sexually divided African lion Panthera leo (recent lion according to Gross, 1992; for citation of the Pleistocene lion see Comparative material). White rhombus e Middle Pleistocene lions, white circles e late Middle Pleistocene lions, black dots e Late Pleistocene lions, all measurements in mm.

Fig. 12. Measurements of metapodiales and calcaneus bone comparisons of the Middle and Late Pleistocene lion Panthera spelaea plotted with recent, sexually divided African lion Panthera leo (recent lion according to Gross, 1992; for citation of the Pleistocene lion see Comparative material). White rhombus e Middle Pleistocene lions, white circles e late Middle Pleistocene lions, black dots e Late Pleistocene lions, all measurements in mm.

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Fig. 13. Measurements of teeth comparisons of the Middle and Late Pleistocene lion Panthera spelaea plotted with recent, sexually divided African lion Panthera leo (recent lion according to Gross, 1992; for citation of the Pleistocene lion see Comparative material). White rhombus e Middle Pleistocene lions, white circles e late Middle Pleistocene lions, black dots e Late Pleistocene lions, all measurements in mm.

individuals can be easily explained by an elongation process in the Pleistocene lion lineage evolution. The blade and cutting projection surface of lower carnassials was in later and more evolutionary advanced forms longer, which provided more efficient meat slicing (Barycka, 2008). Based on those figures, which illustrate the summarized size variability of P. spelaea from different time periods, P. spelaea fossilis was on average slightly larger in linear size than P. spelaea spelaea. Generally speaking, Middle Pleistocene lions were slightly larger in body size (skull and postcranial bones) than Late Pleistocene specimens, but teeth were roughly the same size. The modern African lion also is characterized by great size variability. On average, the South African lion P. leo krugeri (Roberts 1929) is recently the largest lion subspecies, while the West African lion P. leo senegalensis (Mayer 1826) is regarded as (on average) one of the smallest. The most known East African lion P. leo nubica (Blainville 1843) is rather moderate in size, but there are many very big specimens, far larger than South African lions. Specimen variability also should be taken into consideration in fossil material (Hemmer, 1974; Mazak, 1975; Lupták, 2009). 4. Discussion 4.1. Systematical analysis The lion morphology has been subject to numerous detailed studies. Most of the features distinguishing the Middle (fossilis sensu stricto) and Late Pleistocene (spelaea sensu stricto) lion are dentition characters (Wurm, 1912; Heller, 1953; Kabitzsch, 1960; Kurtén, 1960; Schütt, 1969a; Schütt and Hemmer, 1978; Argant, 1988, 1991; Janossy, 1990; Hemmer, 2004; Bona, 2006; Hanko, 2007; Barycka, 2008; Marciszak and Stefaniak, 2010; Argant, 2010a; Sabol, 2011a). Complete skulls of the Late Pleistocene lion are rare, and remains of the Middle Pleistocene lion are extremely rare. Until now, the only complete skull is the famous Mauer specimen. The main Mauer fauna has been dated between MIS 15 and 14 (Wagner et al., 2011) and the same age is postulated for the Mauer skull (Hemmer, 2011). Another two, quite well preserved, skulls from Château Breccia are still under extensive research (Argant, 1991, 2000; Argant et al., 2007; Argant and Argant, 2012 pers. comm.). Three among major morphological characters of the skull distinguish fossilis from spelaea: narrow and long rostrum, long and narrow temporal region and almost flat frontal region (in the lateral view) are well marked in all three. Moreover, the C1-C1

B/TOT ratio distinguishes them from those of younger lions (Argant and Argant, 2012 pers. comm.) (Fig.6.). A few other skulls of Pleistocene lions have been described as P. spelaea fossilis or P. leo fossilis, but in fact none of them represents this subspecies. One is a partially damaged specimen from Moggaster Cave, whose stratigraphic position is unclear (Groiss, 1992). It was found together with a scanty material of the cave bear Ursus spelaeus Rosenmüller, 1794, a relatively small wolverine Gulo gulo (Linnaeus, 1758) and a red deer Linnaeus, 1758 (Groiss, 1992; Döppes, 2001). Groiss (1992) and Argant et al. (2007) assigned the lion remains from Moggaster Cave to fossilis; recently they were re-described as intermediate between fossilis and spelaea form (Marciszak and Stefaniak, 2010). The skull size is similar to that of the Mauer specimen, and it has also a narrow snout and postorbital bar. However, the teeth morphology shows more progressive features: massive P3 with short and low paracone, wide P4 with paracone and metastyle of similar length, narrow p4 with long and low protoconide, low and narrow m1 (B/L ratio ca. 50.5) and p4 and m1 almost equal in length. The morphology of the Moggaster Cave individuals is similar to the lion from Bisnik Cave, dated as MIS 8-7. In both cases, dental morphology confirms a similar stage of evolutionary development and indicates an intermediate form, dated generally to the late Middle Pleistocene (Marciszak and Stefaniak, 2010). The skull from Petralona Cave was assigned to fossilis (Hemmer, 1974; Kurtén and Poulianos, 1977, 1981; Schütt and Hemmer, 1978) or spelaea (Tsoukala, 1989, 1991; Baryshnikov and Tsoukala, 2010; Sabol, 2011b). The morphology shows a mixture of features of both fossilis and spelaea forms. The typical characters of fossilis are proportionally narrow (POB/TOT ¼ 17.0) and long postorbital bar (postorbital processes displaced far anteriorly) and less inflated and relatively small bullae tympanica (Hemmer, 1974; Kurtén and Poulianos, 1977; Schütt and Hemmer, 1978; Baryshnikov and Tsoukala, 2010). The characters of spelaea are visible in the shortening and widening of the rostrum (C1-C1 B/TOT ¼ 27.7), the large, rounded orbits and the convex frontal skull profile (although the curvature of the profile is less marked than in the Late Pleistocene individuals) (Hemmer, 1974; Baryshnikov and Tsoukala, 2010). The lower dentition is also characterized by intermediate features: the very wide p4 and m1 (as in fossilis) stand in clear opposition to the narrow premolars and p4 is distinctly shorter than M1 (as in spelaea) (Hemmer, 1974; Schütt and Hemmer, 1978; Baryshnikov and Tsoukala, 2010). The upper dentition (P3 and P4) of the Petralona skull is so worn that it excludes any detailed morphometric analysis (Baryshnikov and Tsoukala, 2010; Baryshnikov pers. comm. 2011). The stratigraphic position of the Petralona skull is unclear. The layers and sediments with the lion remains from Petralona are generally dated as ca. 400e350 ka (MIS 11-10) (Kurtén and Poulianos, 1977; Schütt and Hemmer, 1978; Kurtén, 1983; Baryshnikov and Tsoukala, 2010; Kahlke et al., 2011; Sabol, 2011b). The carnivore fauna from Petralona was recently divided into three faunal assemblages. The second assemblage, dated as the late Middle Pleistocene, includes Ursus deningeri, Crocuta crocuta spelaea, P. spelaea and Felis silvestris (Baryshnikov and Tsoukala, 2010). Pleistocene lions with progressive morphology closer to P. s. spelaea first appeared in western Asia, eastern and southeeast Europe (Sabol, 2011b), and the lions from Petralona Cave with their intermediate features, fit well with this theory. The last skull recognized by Sotnikova and Nikolskiy (2006) as the fossilis was an intact cranium from Azé Cave, assigned by most authors to primitive spelaea (Argant, 1988, 1991, 2000, 2010a, 2010b). More recently, the individual was classified as the intermediate form, with mixed features characteristic of both Middle and Late Pleistocene lion (Argant, 2010b). Sotnikova and Nikolskiy (2006) justified this assignation with the existence of features

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which were distinctly different from the typical spelaea, and which generally indicated fossilis: large size, less inflated bulla tympanica, very small incisors, narrow rostrum at the level of canines and upper carnassials, relatively broader nuchal surface and proportionally small orbits. It is true that the Azé skull shows many primitive features, but it also displays many progressive characters (compared to the Mauer specimen), closer to spelaea. The mastoid area is relatively broad (MAS/TOT ¼ 38.9), frontal skull profile is convex and the temporal region is proportionally short with very broad postorbital bar (POB/TOT ¼ 22.8) (Fig.6.). The dentition also confirms the intermediate position of the Azé lion. P3 is broad with short and low paracone and very wide (anteriorly and posteriorly) P4 with minute preparastyle and metastyle comparable in length with paracone. Broad p3 and p4 have long and low protoconides, and very wide and high m1 has a poorly reduced talonide and is comparable in length with p4 (Argant, 1988, 1991, 2010a, 2010b). Based on morphology, we question Sotnikova and Nikolsky’s (2006) assignation of the Azé cranium to fossilis and agree with Argant (1991), who regarded the specimen as a large, primitive cave lion or, more recently, intermediate, late Middle Pleistocene form (Argant, 2010b). Moreover, according to Argant (in Baryshnikov and Tsoukala, 2010) the Middle Pleistocene lion did not survive to MIS 9. The detailed morphological and statistical analysis of the dental material of the Middle, late Middle and Late Pleistocene lions confirmed that generally the so called “typical fossilis” occurred in MIS 17 (19) (for details see Lewis et al., 2010) to MIS 10 (Argant in Baryshnikov and Tsoukala, 2010). The lion remains dated at MIS 11-9 from central and western Europe showed a clearly more progressive morphology (Janossy, 1990; Hanko, 2007; Marciszak and Stefaniak, 2010). The Azé sediments, from which the lion remains come, are dated as MIS 6 (Argant, 1988, 1991). Lion remains from a Polish locality Deszczowa Cave, also dated as MIS 6, are very similar to the Azé specimen in their morphology and state of preservation (Barycka, 2008). The size and morphology of the San individual are very similar to the Edingen specimen, whose stratigraphic position is also not quite clear (Diedrich and Rathgeber, 2012). The remains of intermediate lions from Poland (Bisnik Cave, Deszczowa Cave and Wierzchowska Górna Cave), dated to late Middle Pleistocene (MIS 8-6), although large sized, display many primitive features and distinctly differ in their morphology from the San specimen (Barycka, 2008; Marciszak and Stefaniak, 2010; Marciszak, 2012). The most probable age of the San and Edingen specimens is younger than MIS 6, although a slightly older age can not be entirely excluded. However, considering the advanced morphology of the San specimen, it is rather unexpected. Until now, all dredged large mammals (mainly ungulates) from San River were radiocarbon dated to Late Pleistocene (Stefaniak pers. comm. 2013.). Both individuals might be possibly quoted as the exceptions among Late Pleistocene lions mentioned by Argant et al. (2007) and Argant (2010a). 4.2. Discussion e size fluctuation For a long time it has been commonly accepted that large size is more typical for the early lions. According to Lewis et al., (2010), “There has long been a general perception that the earliest European lions were particularly large”. It was widely accepted that the Middle Pleistocene lions were larger, the Late Pleistocene ones were smaller, and the size decreased in the late Middle Pleistocene (Kurtén, 1968; Schütt and Hemmer, 1978; Bishop, 1982; Groiss, 1992; Baryshnikov and Boeskorov, 2001; Hemmer, 2004; Bona, 2006; Sotnikova and Nikolskiy, 2006; Barycka, 2008; Lupták, 2009; Baryshnikov, 2011). In the past, the Mauer skull was used as an example of the great size of the Middle Pleistocene lion. To

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describe the Mauer individual, adjectives such as “largest”, “exceptionally”, “unusual” “extremely”, and “giant” were used (Kurtén, 1960; Dietrich, 1968; Hemmer, 1974; Kurtén and Poulianos, 1977; Hemmer, 2003). However, the dimensions of this individual were in the last thirty years exceeded by new findings of similar or even larger specimens (all identified as adult males) from Château, Mokhnevskaya Cave, Edingen and San River (Baryshnikov and Tsoukala, 2010; Argant and Argant, 2012 person. com.; Diedrich and Rathgeber, 2012). According to Argant et al. (2007:127) the “size drops steadily in the course of time (with some exceptions, however) until it reaches that of the living lion”. In 2010, Argant added (2010b:60): “In that case the size can give a fable chronological data but the gender of the individuals must also be determined, because the sexual dimorphism of lions is great and must necessarily be taken into account. Between these two terms the evolution becomes more confusing. During Late Middle Pleistocene (stage MIS 6), smaller forms alternated with big sized forms. During the stage MIS 5, at the beginning of the Late Pleistocene, a period where there still existed some big sized forms, we can observe some smaller individuals. It becomes more difficult to determine a biochronological position. Dating precisely the sites of this period remains always difficult and the cave lions are not well known there, in spite of recent progress in that field”. Comparison of all available dimensions of various cranial and postacranial elements with more or less good stratigraphical position showed that dimensions of the Pleistocene lion from different time periods overall strongly overlap. Simultaneously, the Middle Pleistocene individuals are placed in the upper limit of the range variation of the skull and postcranial dimensions. The main problem with P. s. fossilis is the very limited number of satisfactory preserved specimens allowing more detailed analysis. Only materials from Mosbach 2 and Château Breccia are more numerous, although still scanty. However, the same is also true for the Late Pleistocene lion. The samples from each, even the largest lion collections of Zoolithen Cave, Nied zwiedzia Cave, or Wierzchowska Górna Cave, are not numerous enough to make statistically reliable analyses (minimum number of 30 individuals) (Argant, 2010a). Based on this, it is hard to demonstrate that early lions were “significantly larger” than later forms. We can only give a general overview on dimensions, and based on them, try to give a satisfactory explanation. Lions in Europe in the early Middle Pleistocene really had impressive posture. Their great size is documented by findings from Pakefield (Lewis et al., 2010), Château Breccia (Argant, 1991, 2000; Argant et al., 2007; Argant, 2010a; Argant and Argant pers. comm. 2012), Mosbach 2 (von Reichenau, 1906; Schütt, 1969a,b; Hemmer, 1974; Schütt and Hemmer, 1978) or Mauer (von Reichenau, 1906; Dietrich, 1968; Hemmer, 2004). It might be cautiously said that Pleistocene lions throughout the Middle and late Middle Pleistocene (MIS 19/17 to MIS 6) were comparatively large. Lions from Wierzchowska Górna (MIS 8-5) and Bisnik Cave (MIS 9-6) strongly overlap in size with specimens from Mosbach 2, Château, and La Belle-Roche (von Reichenau, 1906; Schütt, 1969a,b; Hemmer, 1974; Argant, 1991; Kleczko, 1999). Because great sexual dimorphism exists, when comparing different populations from different time periods, the best way is to compare males and females separately. Since the last interglacial and the beginning of the last glacial, the situation seems to be more complicated. Size slowly decreased, and since MIS 5 specimens moderate in size are typical. However, very large individuals, such as specimens from Santenay (Argant, 1991), Mokhnevskaya Cave (Baryshnikov and Tsoukala, 2010) or Repolust Cave (Schütt and Hemmer, 1978), still existed. Large individuals, still retaining numerous primitive features, came also from the Spanish localities Lezetxiki and Arrikrutz (dated at MIS 6-

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5) (Altuna, 1972, 1981; Sotnikova and Foronova, 2012). The Iberian Peninsula, the westernmost area in the European continent, can be regarded as one of the last “primitive cave lions refuges”, where huge individuals with a mixture of ancestral and advanced characters still survived. Somewhere between 48 and 45 ka BP, Pleistocene lion populations rapidly declined and their genetic variability was dramatically reduced, which caused a genetic bottleneck (Barnett et al., 2009). This period, due to a massive stadial within MIS 3, affected haplotype loss, strongly pronounced size decline, and decreasing massiveness. This last period covers MIS 3 and 2 and is characterized by presence of many small or even dwarf specimens, comparable in size with medium-sized modern lions. It is not quite clear if those small, gracile lions represented an eastern (close to spelaea-vereshchagini) or southern (closer to leo) migration event, which replaced larger lions. Another explanation is the ecological response of the local, surviving populations to harsh environmental conditions and declination in prey abundance. This scenario is recently well documented in West Africa, where the short maned, quite gracile and smaller Senegal lion inhabited harsh habitats with limited number of large ungulates (Lupták, 2009). The last European lions were really small. Specimens from Bois de Cantet (Lm1 ¼ 23.2 mm) (Clot et al., 1980), Azoleta Cave (Castanos, 2005 and references therein) and the Romanian locality of Ghidfalaucariera (skull basal length of w249.0 mm) (Samson and Kovacs, 1967) fall into the size variability of average size modern lions. Decline in massiveness was also noted in cave lion evolution. Many authors noted not only considerable size, but also very robust body build (Hemmer, 1974; Argant, 1991, 2000; Hemmer, 2003, 2004; Argant et al., 2007; Argant, 2010a; Hemmer, 2011; Sabol, 2011b). Hemmer (2011) noted that body mass of P. spelaea is 160e180% of the body mass of the similar sized (in linear dimensions) modern African lion. Anyonge (1996) noted that the long bones of the American lion Panthera (leo) atrox (Leidy, 1853) were much thicker than bones of similar-sized modern P. leo. Being very similar, robustly build and big as the American lion, European Pleistocene lions also had extremely powerful forelimbs. It might had given a great advantage, when hunting cave bear in narrow, dark cave corridors. U. spelaeus was atypical prey for the Pleistocene lion, but when food was scare, lions entered deeply into caves and hunted bears (Argant, 1988, 1991, 2000; Diedrich, 2012). It was risky business facing this powerful animal in total darkness, but those strong forelimbs might had helped subdue prey, allowing delivery of a lethal bite. Additionally, the large size also gave them an advantage, as larger and stronger male lions hunt buffalos Syncerus caffer (Sparrman, 1779) and other very big ungulates (Turner and Antón, 1997) more often than do females. 5. Conclusions Analysis of large series of different skeleton elements showed that it might be cautiously suggested that, as proposed by Argant et al. (2007) size declination in the course of the time in the evolution of Panthera spelaea lineage occurred. The long history of this species in Europe can be divided into four main periods, based on size fluctuations. First, the longest period covers the time span since the earliest known lion remains from Pakefield (Lewis et al., 2010), dated at MIS 17 (19), to the latter part of the late Middle Pleistocene (MIS 7-6). It is characterized by presence of very large and massive lions, which can be regarded as among the largest cats that ever existed (Kurtén, 1968; Argant et al., 2007). Lion fossil material from Mosbach 2 (von Reichenau, 1906; Schütt, 1969a,b; Hemmer, 1974), Sima de los Huesos (Cervera, 1992; Garcia, 2003) and especially Château Breccia well documented this great size of those early lions. However, lions from the late Middle Pleistocene

also reached particularly large dimensions. Specimens from Wierzchowska Górna Cave, Bisnik Cave, and Moggaster Cave (Groiss, 1992) have in most cases roughly the same size and stout posture as stratigraphically older lions. The second period is much shorter, but much more dynamic. It includes the last part of the late Middle and the first part of the Late Pleistocene (MIS 6-5). This period is documented by the mixed presence of surviving large individuals with somewhat still primitive features, e. g. from Lezetxiki (Altuna, 1972), L’Herm Cave (Clot, 1980) and Deszczowa Cave (Barycka, 2008). Especially well fitting into this scenario is the giant lion from Mokhnevskaya Cave, which is among the three largest lion skulls ever recorded (Baryshnikov and Tsoukala, 2010). Together with them, smaller, with evolutionary more advanced dentition lions also appeared (Argant, 1991; Hemmer, 2011; Sabol, 2011b). The great morphology and size variability of P. spelaea fossil material from this period is probably connected with the irregular replacement of P. s. fossilis by P. s. spelaea, which entered Europe from the east (Marciszak and Stefaniak, 2010). At roughly the same time, the eastern late Middle Pleistocene lions showed a more advanced morphology, compared to those from Western Europe (Schütt and Hemmer, 1978; Barycka, 2008; Hemmer, 2011; Sabol, 2011b). The third period covers the first part of the last glacial, from MIS 5 to the beginning of MIS 3. For this period, occurrence of moderate sized specimens with evolutionary advanced dentition is documented. As “typical” from this time span individuals from Siegsdorf (Gross, 1992) or Zoolithen Cave (Diedrich, 2012) can be give. Apart from one exception, the holotype from Zoolithen Cave with unclear stratigraphical position (Diedrich, 2008), none of male skulls from this period exceed 400.0 mm in total length. The last period began with dramatic population declination and genetic variability reduction (Barnett et al., 2009). It lasted from MIS 3 to 2, and within it size drops sharply. Among moderate sized specimens, comparable in size with those from first part of the last glacial, very small (sometimes even regarded as dwarfs) lions also appeared. Apart from smaller dimensions, they are also characterized by much less stout posture, and in bone shapes and dimensions they are generally closer to the African lion than the Late Pleistocene lion. It is not quite clear that they represented autochthonous, European populations, whose smaller size might be regarded as ecological response for the hard environmental conditions, as recently in Western Africa (Lupták, 2009). However, migration events from adjacent areas and African or Asian affinities of those small lions are not excluded (Hemmer, 2011; Sabol, 2011b). Even considering that most of those extremely small lions were lionesses, the dimensions are comparable at the most with medium sized specimens of modern African lions. Described in this paper for the first time, the well preserved and particularly large skull of a male cave lion from San River somewhat complicates this scenario. Although the stratigraphical position of this finding is unclear and both dating methods (Radiocarbon and UraniumeThorium) failed, its geological age can be estimated with high probability based on cranial and teeth morphology. Comparison with the similar in size, but with very different morphology P. spelaea fossilis skull from Mauer confirmed a high evolution stage of development. The San lion is characterized by advanced morphology, typical for the Late Pleistocene lion P. spelaea spelaea. It can be seen in the short and strongly widening at the canines level stout, compressing and shortening of the temporal region as well as strongly convex frontal skull profile (in the lateral view). Also, upper dentition features are typical for P. s. spelaea morphology: narrow P3 with proportionally long and low paracone and narrow P4 with reduced and low protocone, well developed preparastyle, paracone shorter than metastyle and strongly convex buccally posterior part of the crown. This represented a much more advanced and higher evolutionary level than the other Eemian

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Polish lion skull from Nied zwiedzia Cave (Sudety Mts). Generally, if the Late Pleistocene age of this individual is accepted, the great size might be resolved in two different ways. Although it could be regarded as one of those large specimens surviving until the Late Pleistocene, the advanced cranial and dental morphology argues against this. It seems that the second explanation, that it is simply a record size specimen among the Late Pleistocene lions with impressive dimensions and advanced morphology typical for the last glacial, might be more reliable. A similar situation can be observed recently, where among medium size African lions some particularly large individuals exceed the dimensions of others (Hemmer, 1974; Yamaguchi, 2005a, 2005b). In conclusion, the general tendency to decreasing size in the evolution of the P. spelaea lineage has been documented. However, it is not useful as a biochronological tool, in view of the great variability between populations and strongly pronounced sexual dimorphism. It seems that lion in western Eurasia was a dynamically evolving population with size fluctuations driven by climate, but influenced by migration between regions and immigration from outside. The overview is also limited by great sexual dimorphism, individual variability, prey abundance and limited number of findings, especially in Middle Pleistocene lions. All those factors make detail analysis of the size fluctuation extremely difficult, and in view of scarcity of P. s. fossilis postcranial material not statistically significant. Perhaps in the future, more numerous finds of especially Middle Pleistocene lions can shed a new light on this problem. Acknowledgements We are very grateful to Alain and Jacqueline Argant for their unpublished data on the two Château lion skulls and to Prof. Dr hab. Beata M. Pokryszko for linguistic improvements. We would like to express also our thanks to Martin Sabol (Comenius University, Bratislava), Helmut Hemmer (Johannes-Gutenberg-Universität Mainz), Alain and Jacqueline Argant (LAMPEA, Lyon), Philippe Fosse (Université de Toulouse le Mirail) and Pawe1 Socha (University of Wroc1aw) for the inspiring discussion and valuable comments on the topic. Last but not least, we express thanks to the referees for their critical reading and helpful comments on the manuscript. This research was supported by the Ministry of Science and Higher Education, Poland (project 1018/S/KBEE/2012). References Aleekseva, L.I., 1990. Late Pleistocene Mammal Fauna of Eastern Europe. Nauka, Moscov (in Russian). Altuna, J., 1972. Fauna de Mamíferos de los yacimientos prehistóricos de Guipúzcoa. In: Con catálogo de los mamíferos cuaternarios del Cantábrico y del Pirineo Occidental. Munibe 24, pp. 1e464. Altuna, J., 1981. Fund eines Skeletts des Höhlenlöwen (Panthera spelaea GOLDFUSS) in Arrikrutz, Baskenland. Bonner Zoologische Beiträge 32, 31e46. Anyonge, W., 1996. Locomotor behavior in Plio-Pleistocene sabre-tooth cats: a biomechanical analysis. Journal of Zoology 238, 395e413. Argant, A., 1988. Etude de L’exemplaire de Panthera spelaea (GOLDFUSS, 1810) (Mammalia, Carnivora, Felidae) du gisement Pleistocene moyen recent de la grotte d’Azé (Soane et Loire). Révue de Paléobiologie 7, 449e466. Argant, A., 1991. Carnivores quaternaires de Bourgogne. In: Documents des Laboratoires de Géologie de la Faculté des Sciences de Lyon 115, pp. 1e301. Argant, A., 2000. Les sites paléontologiques du Pleistocène moyen en Mâconnais. Bulletin de la Société Préhistorique Française 97, 609e623. Argant, A., 2010a. Carnivores (Canidae, Felidae et Ursidae) de Romain-la-Roche (Doubs, France). Revue de Paléobiologie 29, 495e601. Argant, A., 2010b. The cave lion of the Áze cave (Burgundy, France). Biochronology of the cave-lion: an attempt to date the Panthera (Leo) spelaea. In: Argant, A. (Ed.), Program and Guide book of excursion, 16th International Cave Bear and Lion Symposium, Azé (Saõne-et-Loire, France), September 22nde26th, 2010, p. 60. Argant, A., Argant, J., 2011. The Panthera gombaszogensis story: the contribution of the Château Breccia (Saône-et-Loire, Burgundy, France). Quaternaire, Horsesérie 4, 247e269.

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