The Pleistocene lion Panthera spelaea (Goldfuss, 1810) from Poland – A review

Quaternary International xxx (xxxx) xxx

Contents lists available at ScienceDirect

Quaternary International journal homepage: www.elsevier.com/locate/quaint

The Pleistocene lion Panthera spelaea (Goldfuss, 1810) from Poland – A review Adrian Marciszak a, *, Grzegorz Lipecki b, Kamilla Pawłowska c, Gwidon Jakubowski d, ´ ska a, Adam Nadachowski b Urszula Ratajczak-Skrzatek a, Katarzyna Zarzecka-Szubin a

Department of Palaeozoology, Institute of Environmental Biology, Faculty of Biological Sciences, University of Wrocław, Sienkiewicza 21, 50-335, Wrocław, Poland Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Sławkowska 17, 31-016, Krak´ ow, Poland Department of Palaeoenvironmental Research, Institute of Geology, Adam Mickiewicz University, Krygowskiego 12, 61-680, Pozna´ n, Poland d Museum of Earth, Polish Academy of Science, Aleja Na Skarpie 20-27, 00-488, Warszawa, Poland b c

A R T I C L E I N F O

A B S T R A C T

Keywords: Pleistocene Panthera spelaea Dating Lineage Chronosubspecies Competition Extinction

Panthera spelaea was recorded in Poland from 18 open-air and 42 cave sites dated in the range 750–28 ka. Most of these records are located in southern Poland (Silesia) and neighbouring areas. Among them, the find of Panthera spelaea in Kozi Grzbiet mentioned here for the first time is one of the oldest European records of the species. Most of the obtained AMS dates are concentrated on the second half of MIS 3 and showed that the Pleistocene lion disappeared more or less synchronously across Poland. The findings of its remains from open-air sites are mostly accidental discoveries during field works, while those in caves are mostly connected with archaeological research and the exploitation of cave sediments for producing fertilizers. Aside of a few juvenile bones and milk teeth, the remains of adult individuals, mostly males, predominate in the studied sample. Their behavior to hunting cave bear during the times, when the other, more typical food sources were scarce, tendention to hunt very big preys and went into the conflicts with other carnivores than females do, resulted in higher injuries and mortalities. Within the species, three chronosubspecies were recog­ nized, with P. s. fossilis recorded from 7 sites, dated between 750 and 240 ka and P. s. intermedia only partially recognized in Wierzchowska G´ orna Cave. The most numerous are the remains of P. s. spelaea which were found in 48 localities dated between 150 and 28 ka. With the exception of four caves, the Polish finds of P. spelaea are generally less abundant at sites where they occur. Older P. s. fossilis, dated on MIS 19–12, was a large form with robust stature and broad cheek teeth. Younger P. s. fossilis dated on MIS 11–9 was represented by a large and massive specimens, but with more advanced morphological dental features. Polish individuals of P. s. spelaea were slightly smaller and less massive than P. s. fossilis, with narrower cheek teeth. Since MIS 3, a dwarf and gracile specimens appeared, which is correlated with the genetic turnover ca. 48–45 ka. The Pleistocene lion was one of the earliest disappeared large carnivores predated only by a cave hyena. There are no direct evidences of P. spelaea encounters with humans. The extinction of P. spelaea is broadly correlated with the general collapse of the “mammoth steppe” ecosystem and was resulted of multiple reasons like climatic changes, re-building of herbivore guilds, competition and human pressure. Wolf and, to a lesser extent, bears were the main competitors for the Pleistocene lion in Poland.

1. Introduction The Pleistocene lion Panthera spelaea, originally described by Gold­ fuss in 1810, is placed among the most famous Pleistocene mammals. Though generally recognized by most authors as a separate species, the

taxonomical position of this form has for a long time been a subject of controversy. Morphometrical and phylogenetic analysis revealed that P. spelaea was highly distinct and genetically isolated from the modern lion Panthera leo (Linnaeus, 1758) (Argant, 1988, 1991; Marciszak et al., 2014, 2019a, b; Ersmark et al., 2015; Barnett et al., 2016; Argant and

* Corresponding author. E-mail addresses: [email protected] (A. Marciszak), [email protected] (G. Lipecki), [email protected] (K. Pawłowska), gwidon_ [email protected] (G. Jakubowski), [email protected] (U. Ratajczak-Skrzatek), [email protected] (K. ZarzeckaSzubi´ nska), [email protected] (A. Nadachowski). https://doi.org/10.1016/j.quaint.2020.12.018 Received 14 July 2020; Received in revised form 3 December 2020; Accepted 9 December 2020 Available online 17 December 2020 1040-6182/© 2020 Published by Elsevier Ltd.

Please cite this article as: Adrian Marciszak, Quaternary International, https://doi.org/10.1016/j.quaint.2020.12.018

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Brugal, 2017; de Manuel et al., 2020). These analyses support the taxonomic recognition of P. spelaea as a separate species that diverged from the lion about 1.9 Ma (Barnett et al., 2009, 2016). In spite of the progress attained during the last decades, it must be noted that several taxonomic problems still exist. One of the basic problems, especially of older publications, is that too often determina­ tion was based exclusively on postcranial remains (osteological basis) so the errors of species identification are frequent, whereas the attribution by means of cranial morphology and teeth is unequivocal. Large, pan­ therinae cats are a highly stereotyped group with a low morphological divergence. This means that morphologically substantial intraspecific differences and, conversely, interspecific similarities may be found among these remains. This paper summarises almost 150 years of the study of P. spelaea remains from Poland, including a particular number of new localities. Previously, Kowalski (1959) listed 12 sites with lion remains, Wolsan (1989a) added a few more, while Barycka (2008) presented 19 localities and described in detail some materials. Our re-examination of old col­ lections in different, mostly Polish but also German museums expanded the list up to 60 localities (Fig. 1). It is also the result of recovering of many specimens, which were previously regarded to be lost. Addition­ ally, a new set of 16 radiocarbon dates is presented, with 13 ones un­ published so far. Accordingly, this paper is intended to present the current state of knowledge concerning the diversity of P. spelaea in Poland based on their geographical and temporal variability within Eurasia.

incomplete and needs a revision. One of such examples is the Pleistocene lion, which is one of the most recognisable member of Eurasian palaeofaunas. The main goal of this paper is to reconstruct the history of the species within Poland territory. The occurrence or absence of particular species is usually regard as its adaptation in response to the environmental changes. However, it will be also analyse third, also important factor, the interspecific competition between them. In this context, the location of Poland as a west-eastern faunal migration corridor also highlighted this problem. 3. Main stages of P. spelaea study in Poland In the context of our studies, the Polish material of P. spelaea is rather ´rna Cave (Ja ´nossy, important, especially that from Wierzchowska Go 1969; Turner, 1984; Kurt´en, 1985; Barycka, 2008; Argant and Brugal, 2017). Althougth the history of the study of Pleistocene lion remains from Poland was already described before (Hoyer, 1937; Kowalski, 1959; Wolsan, 1989; Barycka, 2008), it is strongly incomplete and re­ quires an additional thorough overview. Most of the findings came from cave deposits, and open-air findings are much rarer. Among them is the oldest Polish lion record found in ´w near Szprotawa (Silesia), where an isolated m1 of considerable Witko size was found in the marl mine (Hensel, 1852, 1853). The number of lion records started growing between 1875 and 1939, and it is strongly connected with archaeological excavations and exploitation of cave sediments for fertilization (Wojtal, 2007; Barycka, 2008). However, these finds were mostly mentioned or only briefly described, and the vast majority of these studies based on a single or very few bones, ¨mer, 1883; Gürich, 1885; Ossowski, mandibles or isolated teeth (Ro 1887; Conwentz, 1895; Frech, 1904; Hermann, 1911, 1913; Sonntag, 1919; Hoyer, 1937; Zotz, 1939; Kowalski, 1959). The only more detailed studies of the Polish material of P. spelaea predated the World War II were published by Lubicz-Niezabitowski (1925, 1938a, b) and Ryzie­ wicz (1933). It is even despite that the enormous Ossowski’s collection ´rna Cave was already almost totally excavated. from Wierzchowska Go Wojtusiak (1953) studied most of the cranial and part of the postcranial material in detail after the World War 2, and it is the first comprehensive

2. Regional setting The state of knowledge and understanding of the stages and dy­ namics of the dispersion and changes of the particular faunal elements on areas north of the Carpathians has significantly increased over the last two decades. It was due to a new chronological evidences, a broad revision of historical collections as well as new excavations allowing to re-examine old and describe new materials which particularly increased our knowledge about the formation of fauna in this region. However, data from Central Europe, among them also from Poland still seems to be

Fig. 1. Distribution of the Pleistocene lion Panthera spelaea in the Middle and Late Pleistocene of Poland: 1 - Mały Baldram; 2 - Bębłowska Dolna Cave; 3 - Biały Kamie´ n Cave; 4 - Bi´snik Cave; 5 - Borsucza Cave; 6 - Brzeziny; 7 - Cave no. 4 on Bir´ ow Hill; 8 - Ciemna Cave; 9 - Deszczowa Cave; 10 - Draby; 11 - Dziadowa Skała Cave; 12 - Dzierzgo´ n; 13 - Głuszyca G´ orna; 14 - Hureczko; 15 - Jasna Strzegowska Cave; 16 - Komarowa Cave; 17 - Kowan´ owek; 18 - Koziarnia Cave; 19 - Kozi Grzbiet; 20 - Krakowska Cave; 21 - Krosinko; 22 - Łokietka Cave; 23 - Magurska Cave; 24 - Maszycka Cave; 25 - Naciekowa Cave; 26 - Na Gołąbcu Cave; 27 - Nied´zwiedzia Cave; 28 - Nied´zwiedzia G´ orna Cave; 29 - Nietoperzowa Cave; 30 - Obłazowa Cave; 31 - Obok Wschodniej Cave; 32 - Oborniki; 33 - Ojc´ ow vicinity; 34 - Pawłowiczki; 35 - Perspektywiczna Cave; 36 - Pod Słupami Cave; 37 - Południowa Cave; 38 - Caves on Połom Hill; 39 - P´ ołnocna Du˙za Cave; 40 - Pyskowice; 41 - Radochowska Cave; , 42 - Raj Cave; 43 - Rakoszyce; 44 - Rog´ oz˙ ka Cave; 45 - San River; 46 - Skarszyn; 47 - Sob´ otka; 48 - Sosnowiec-Milowice; 49 - Stajnia Cave; 50 - Szkaradowo; 51 - Towarna Cave; 52 - Tunel Wielki Cave; 53 - Waplewo Wielkie; 54 - Węgry; 55 - Wierzchowska G´ orna Cave; 56 - Witk´ ow; 57 - Wschodnia Cave; 58 - Zamkowa Dolna Cave; 59 - Zawalona Cave; 60 - Zb´ ojecka Cave; 61 - Zegar Cave. 2

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

research of the Pleistocene lion from Poland. In overall, the interest in study and re-examination of the old lion material was growing between 1950 and 2020 (e.g. Wiszniowska, 1978; Barycka, 2008), and de­ scriptions of new materials appeared at that time (Barycka, 2008; Marciszak and Stefaniak, 2010; Marciszak et al., 2011, 2014, 2019a, b; Woroncowa-Marcinowska et al., 2013). In meantime, alongside with the ´rna most numerous Polish collection of P. spelaea from Wierzchowska Go Cave, two other sites, Bi´snik and Nied´zwiedzia Caves, also provided abundant lion material (Wiszniowska, 1978, 1989; Barycka, 2008; Marciszak and Stefaniak, 2010; Marciszak et al., 2011, 2014, 2019a, b). Lion materials from other sites were usually mentioned and never described in detail (Kowalski, 1972; Cyrek et al., 2000; Lipecki et al., 2001; Lipecki and Wolsan, 2003; Nadachowski et al., 2009; Pawłowska, 2015b).

Table 1 Records of Panthera spelaea from Poland (MNI – Minimum Number of In­ dividuals, NISP – Number of Identified Specimens, ♂ – male, ♀ – female). Site Panthera spelaea fossilis Kozi Grzbiet Tunel Wielki Cave Południowa Cave Wschodnia Cave Draby Bi´snik Cave, layers 19ad–18 Deszczowa Cave Wierzchowska G´ orna Cave Panthera spelaea intermedia Wierzchowska G´ orna Cave Panthera spelaea spelaea Biały Kamie´ n Cave Bi´snik Cave, layers 12–4 Borsucza Cave Brzeziny Cave no. 4 on Bir´ ow Hill Ciemna Cave, layers 18–1 Dziadowa Skała Cave, layer 7 Hureczko Jasna Strzegowska Cave Komarowa Cave, layers G–D Kowan´ owek Koziarnia Cave Krakowska Cave Krosinko Łokietka Cave Magurska Cave Maszycka Cave Naciekowa Cave na Gołąbcu Cave Nied´zwiedzia Cave Nied´zwiedzia G´ orna Cave Nietoperzowa Cave Obłazowa Cave Oborniki Pawłowiczki pod Słupami Cave Caves of Połom Mt Perspektywiczna Cave P´ ołnocna Du˙za Cave Pyskowice Radochowska Cave Raj Cave, layers 10–2 Rakoszyce Rog´ oz˙ ka Cave San river Skarszyn Sob´ otka Sosnowiec–Milowice Stajnia Cave, layers D–C Towarna Cave Waplewo Wielkie Węgry Wierzchowska G´ orna Cave Wschodnia Cave Zamkowa Dolna Cave Zawalona Cave Zb´ ojecka Cave Zegar Cave Panthera spelaea ssp. Bi´snik Cave, layers 15–14 Bębłowska Dolna Cave Ojc´ ow (vicinity) Dzierzgo´ n Mały Baldram Głuszyca G´ orna Szkaradowo Wierzchowska G´ orna Cave Witk´ ow

4. Material and methods 4.1. Material Presence of the Pleistocene lion was documented from 60 sites from Poland, 42 cave and 18 open-air localities. In this paper materials from 55 sites were re-evaluated, while bones from 5 other localities are missing or their status is unknown. The materials of P. spelaea are currently divided among different, mostly Polish institutions. Most of the palaeontological remains are housed at the Department of Palae­ ozoology, University of Wrocław (27 sites) and Institute of Systematics ´w (17 and Evolution of Animals, Polish Academy of Sciences in Krako localities). Smaller samples are stored in the Department of Evolutionary Palaeobiology, Polish Academy of Sciences and Institute of Geological Sciences, Polish Academy of Sciences, both in Warszawa and Archaeo­ logical Museum Wrocław. Few bones are also located in Podlaskie Museum in Białymstok, Museum of Fossils and Minerals (Robert Szybiak collection) in Dubiecko, Museum of the Earth, Polish Academy of Sci­ ences in Warszawa, National Museum in Kielce, Geological Museum of the Polish Geological Institute-National Research Institute in Warszawa and Regional Museum name Wojciech Dutkiewicz in Rogo´zno. Single findings are also housed in two German museums: Naturkunde-und Mammut-Museum in Siegsdorf and Museum für Naturkunde, Hum­ ¨t in Berlin. boldt Universita 4.2. Morphometric analysis The identification of P. spelaea remains was performed using the basic morphometric analysis. Measurements were taken point to point, with an electronic calliper, up to the nearest 0.1 mm. Each value given here is the mean of three measurements (all in mm). The mean is rounded to one decimal place. Measuring schemes and morphological terminology were taken and modified from Schmid (1940) and Argant (2010) and showed in Appendix 2. Measurements of lion material are given in Tables S1–S9 in Appendix 2. Throughout the text, upper teeth are referred as capital letters (e.g. P4), while the lower ones as lowercase letters (e.g. p4). The MIS (Marine Isotopic Stages) bouduaries after Lisiecki and Raymo (2005). 4.3. Radiocarbon dating The new radiocarbon dates (AMS) presented in this paper (Table 1) ´ Radiocarbon Laboratory (Poland) following were obtained in Poznan the pre-treatment protocol for the collagen extraction (Longin, 1971; Piotrowska and Goslar, 2002). Before extraction, the degree of collagen degradation was checked by measuring the content of nitrogen and carbon in bone samples using the analyser Flash EA 1112 Series (Ther­ mo-Scientific). The samples were regarded suitable for collagen dating if the nitrogen content was not lower than 0.6%, and C/N ratio was not higher than 5. Suitable bones were crushed mechanically to granulation <0.3 mm, and the obtained bone powder was then treated with 2M HCl 3

MIS

NISP/MNI

19 12 15–12 15–12? 11 10–9/8 9–8 10–9

2/1 (♂) 36/2 (1 ♂, 1 ♀) 4/1 (♂) 1/1 (♂) 2/2 (♂, ♀) 210/9 (7 ♂♂, 2 ♀♀) 9/1 (♂) 7/3 (3 ♂♂)

8–7/6

28/5 (4 ♂♂, 1 ♀)

4–3 6–3 5–3 5–3 3 6–3 3 5–3 5–3 5–3 3 5–3 5–3 3 5–3 5–3 3 5–3 5–3 5–3 4–3 5–3 3 3 4–3 4–3 5–3 4–3 4–3 3 3 4–3 3 3 6–5? 3 3 3 3 4–3 LP LP 6/5–3 5–3 4–3 4–3 LP 4–3

1/1 91/15 (11 ♂♂, 4 ♀♀) 4/1 (♂) 1/1 (♀) 1/1 23/7 (5 ♂♂, 2 ♀♀) 2/1 2/1 (♀) 1/1 21/2 (♂, ♀) 1/1 (♂) 1/1 (♂) 1/1 (♂) 2/2 (2 ♀♀) 7/2 (♂, ♀) 4/1 (♀) 1/1 (♂) 136/6 (5 ♂♂, 1 ♀) 1/1 (♀) 336/8 (7 ♂♂, 1 ♀) 20/3 (2 ♂♂, 1 ♀) 33/4 (3 ♂♂, 1 ♀) 3/1 (♀) 1/1 (♀) 1/1 2/1 (♂) 22/3 (2 ♂♂, 1 ♀)

8/7–6 ? ? ? ? ? ? 10–3 ?

23/3 (♂, 2 ♀♀) ? 1/1 1/1 (♂) ? 1/1 (♀) 1/1 (♂) 461/15 (12 ♂♂, 3 ♀♀) 1/1 (♂)

2/1 (♂) 5/1 (♂) 1/1 16/3 (2 ♂♂, 1 ♀) 2/1 (♂) 2/1 (♂) 1/1 (♂) 2/1 (♂) 1/1 (♀) 1/1 (♀) 3/1 (♂) 18/2 (♂, ♀) 1/1 (♂) 1/1 (♂) 43/6 (5 ♂♂, 1 ♀) 5/1 (♂) 4/1 (♂) 2/1 3/1 10/2 (♂, ♀)

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

(20 min), and 0.1M NaOH (60 min). After each step of the treatment, the sample was centrifuged and the residuum was collected. Extraction of the collagen was processed in HCl (pH = 3, 80 ◦ C, 10 h), and the residuum was removed after centrifugation. The extracted collagen was then ultrafiltered on pre-cleaned Vivaspin 15 MWCO 30 kD filters. Quality of the collagen was ultimately assessed based on C/N ratio (interval of acceptance 2.7–3.5) and collagen extraction yield (accep­ tance threshold 0.5%). On demand, the carbon and nitrogen stable isotopic composition in the collagen can be determined (Bronk Ramsey et al., 2004).

Boeskorov and Lazarev (1997); Nagel (1997); Turner (1997, 1999); Guˇzvica (1998); Kleczko (1999); Baryshnikov and Boeskorov (2001); ´ ´ (2005); Anto ´n et al. (2005); Foronova (2001); Garci´ a (2003); Abelov a ˜ Castanos (2005); Bona (2006); Pacher (2006, 2018); Sotnikova and Nikolsky (2006); Testu (2006); Argant et al. (2007); Diedrich (2007a, b, ´ (2007); 2008, 2009a, b, 2010, 2011a-f, 2012, 2014a, 2017); Hanko Toˇskan (2007); Barycka (2008); Baryshnikov and Petrova (2008); ¨ppes (2009); Ovodov and Ovodov and Zaika (2008); Kempe and Do Tarasov (2009); Baryshnikov and Tsoukala (2010); Cr´egut-Bonnoure et al. (2010); Lewis et al. (2010); Baryshnikov (2011, 2016); Cr´ egut-­ Bonnoure (2011); Hemmer and Keller (2011); Pautret-Homerville et al. (2011); Sabol (2011a, 2014); Bona and Sardella (2012, 2014); Diedrich ´n ˇ a and Roblíˇckova ´ (2013); Marciszak et al. and Rathgeber (2012); Ka (2014, 2019a, b); Sotnikova and Foronova (2014); Fourvel et al. (2015); Argant and Brugal (2017); Sabol et al. (2018).

4.4. Abbreviations used in the text The list of abbreviations used in the paper: B Ba Bp CBL C1/c1 d L LP M MIS mm MNI mtcp mtts N NISP pa ph I-1(2/3/4/5)-a ph I-2(3/4/5)-p ph II-1(2/3/4/5)-a ph II-2(3/4/5)-a ph III pr SD V

breadth anterior breadth posterior breadth condylobasal length upper canine/lower canine distal length Late Pleistocene mean Marine Isotopic Stage millimeter Minimum Number of Individuals metacarpal metatarsal number of specimens in sample Number of Identified Specimens paracone(id) first (second/third/fourth/fifth) proximal phalanx of manus second (third/fourth/fifth) proximal phalanx of pes first (second/third/fourth/fifth) medial phalanx of manus second (third/fourth/fifth) medial phalanx of pes distal phalanx protocone(id) standard deviation variation

5. Results 5.1. General remarks The data presented here are a result of an extensive revision and the first comprehensive account of all so far known records of the Pleisto­ cene lion P. spelaea from Poland. The work includes both quantitative and qualitative updates, however it is neither exhaustive nor complete. It is very likely that there are still many finds that have not been reported to scientific institutions or museums, or else remain in private collec­ tions. They were often found by local searchers and may have been sold on account of their great value and rarity. The material from some lo­ calities, e.g. caves of Połom, Pyskowice and Sosnowiec-Milowice, is scattered over many museums and private collections, and it still con­ tains hundreds of bones of different species to be further studied. The occurrence of the Pleistocene lion P. spelaea has been recorded from 60 Polish sites dated from the early Middle until the Late Pleisto­ cene (Fig. 1), and covered the timespan between MIS 19 and MIS 2 (Tables 1 and 2). The obtained AMS dates demonstrated that P. spelaea survived in Poland until the transition between MIS 3 and MIS 2 (Table 1). Most of the dates are concentrated on the second half of MIS 3 and showed that the species disappeared more or less synchronously in the entire Poland (Table 1). However, given the limitations of the data, it is possible that later survivals may be discovered yet in some areas. These findings should be taken with caution because of the small sample sizes, but their potential significance is valid. Anyway, it means that P. spelaea is one of the species which disappeared much earlier as compared to other carnviores, and it was predated only by Crocuta crocuta spelaea Goldfuss, 1810. The latter has vanished from the Polish territory even earlier. However, our knowledge of P. spelaea in Poland is still incomplete and scanty, and needs further detailed research. Probably many undis­ covered lion remains are held in private hands, and many sand mines are still active. Among these 60 records, 18 are open-air sites alongside the entire territory of Poland (Fig. 1). Most of these findings are loose, single or few bones, with lose context, and came mostly from sand and marl mines or gravel pits. The increasing in the number of these findings in the second half of the nineteenth century was connected with a period of intense indus­ ´ trialisation of western Poland (Pomorze, Wielkopolska and Sląsk) and thus a significant intensification of earthworks related to the railway and road infrastructure, development of towns and villages, as well as a broad-based action of drainage and regulation of watercourses. The accidental character of such findings means that the exact location, stratigraphical context and other information are usually scarce, if present at all. The lion material from the Polish open-air sites was probably accumulated through natural processes and there is no evi­ dence of human activity. The accumulation of remains may be an outcome of natural or catastrophic death, slope or fluvial processes, activity of carnivores and some additional factors. Remains were also

4.5. References analysed for this study Morphometrical analysis is based on our own measurements and data set taken from the literature: Dawkins and Sandford (1866); Filhol ¨mer (1883); Woldˇrich (1893, and Filhol (1871); Bourguignat (1879); Ro 1897); Hagmann (1899); Kafka (1903); Boule (1906a, b); von Reichenau ˇ (1906); Wurm (1912); Freudenberg (1914); Zelizko (1918); Fabiani ¨ (1919); Hilzheimer (1922, 1927); Klahn (1922); Schmidtgen (1922); Lubicz-Niezabitowski (1925, 1938a); Jaeckel (1927); Gromova (1928); Edinger (1931); Dubois and Stehlin (1932); Riabinin (1932); Ryziewicz (1933); Gromov (1935); Leonardi (1935); Schmid (1940); Koby (1941); ´ and Casajuana (1950); Pasa (1947); Mottl (1949); Crusafont-Pairo Rakovec (1951, 1958, 1965); Heller (1953); Wojtusiak (1953); Anelli (1954); Del Campana (1954); Lehmann (1954); Petrbok (1954); ´zvorka (1954); Pidoplichko (1956); Arambourg (1960); Musil (1960, Za 1969, 1996); Malez (1963); Terzea (1965); Zapfe (1966); Samson and Kovacs (1967); Dietrich (1968); Martin (1968); J´ anossy (1969a, b, 1990); Mostecký (1969); Schütt (1969a, b); Hemmer and Schütt (1970); Suire (1970); Vereshchagin (1971); Altuna (1972, 1981); Thenius (1972); Hemmer (1974, 1977, 2001); Adam (1975); Ballesio (1975, ¨ck (1975); Kurt´ 1980); Daxner-Ho en and Poulianos (1977, 1981); Caloi and Palombo (1978); Schütt and Hemmer (1978); Wiszniowska (1978); Cr´egut (1979); Piccoli et al. (1979); Clot (1980); Clot et al. (1980); Bishop (1982); Riedel (1982); Erdbrink (1981, 1983a, b, 1986, 1988); Siegfried (1983); Toepfer (1983); Nielbock (1987); Ruprecht (1987); Argant (1988, 1991, 2010); Lumley et al. (1988); Duforur (1989); Alekseeva (1990); Sala (1990); Cervera (1992); Groiβ (1992, 2002); Gross (1992); Astruc et al. (1994); Fischer (1994); Valensi (1994); 4

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Table 2 The list of radiocarbon (C14) and Uranium–Thorium (UTh) dates obtained for Panthera spelaea from Poland. The calibration for dates was made with the OxCal software (version OxCal v. 4.3, Bronk Ramsey et al., 2004). Atmospheric data were obtained from Reimer et al. (2013). Presented calibrated date with 95.4% probability. An amount of a nitrogen (N), carbon (C) and collagen (coll.) is presented in percentage. Site

Bone

Coll. no.

Lab. no.

Uncal. date BP

Cal. date b2k

N

C

source

Raj Cave, layer 10 Raj Cave, layer 10 Obłazowa Cave Stajnia Cave, layer D Zamkowa Dolna Cave Towarna Cave Naciekowa Cave Nied´zwiedzia Cave Nietoperzowa Cave, layer 5 Wierzchowska G´ orna Cave Wierzchowska G´ orna Cave Zawalona Cave, layer E Sob´ otka Oborniki Krosinko Nied´zwiedzia Cave

cuboid atlas mandible ph III talus p3 mandible calcaneus humerus mandible tibia P4 mandible mtcp V mandible cranium

MF/1254/1306 MF/1254/639 MF/6412 JS/Pss/1 ZD/6/1 MF/6587 JNW/Pss/111 JNK/Pss/133 MF/6254 MF/320/6889 MF/320/6857 MF/2117 MB/P/68 MRR-687/G; Ob/2013/8.F3 KR1 111.F71 JNK/Pss/1

Poz-58067 OxA-11096 Poz-58065 Poz-43095 Poz-111419 Poz-58066 Poz-25168 Poz-25167 Poz-23627 Poz-59970 OxA-10087 OxA-11156 Poz-28256 Poz-96211 OxA-26790 W-2845

23250 ± 25190 ± 25900 ± 31900 ± 35070 ± 37200 ± 37200 ± 37300 ± 37900 ± 38200 ± 38650 ± 38800 ± 39800 ± 40000 ± >48300 60000 ±

27772-27347 30326-28611 30698-29682 39676-33850 41161-38507 42512-40902 43141-40118 43460-39996 43297-41221 43834-41312 43714-41911 45057-41433 45757-42149 45658-42390

1.7

3.7

0.6 1.2 2.6 2.7 1.4 1.8 5.9 3.7

2.4 5.5 6.4 6.4 7.1 6.4 19.0 8.6

4.6 3.0

13.4 9.5

this paper Stuart and Lister, (2011) this paper this paper this paper this paper this paper this paper this paper this paper Barnett et al., (2009) Stuart and Lister, (2011) this paper this paper this paper this paper

found in 42 cave localities concentrated mostly in Sudety Mts and ´w-Częstochowa Upland (Silesia) (Fig. 1). The increasing of Polish Krako P. spelaea findings from cave deposits since the second half of the nineteenth century is strongly connected with the growing interest to archaeological research of caves and due to the exploitation of their sediments for producing fertilizers. A number of lion bones found in particular caves is diversified, and varied from single specimens up to more than 500 bones. Generally, the number of bones is low in most localities, varied between 10 and 15 bones or, in few caves, more than 20 bones were found. Only from 4 caves the number of P. spelaea re­ mains is considerably higher, encounting 480 bones in Wierzchowska ´rna Cave, 324 in Bi´snik Cave, 336 in Nied´zwiedzia Cave and 136 in Go Naciekowa Cave. Beside a single open-air record from Pawłowiczki and a few caves sites, all other Polish finds of P. spelaea are represented by adult in­ dividuals, mostly adult, but not senile (Table 2). Very old specimens are rare, and the vast preponderance of large adult males was noticed. Since the bones with clear signs of carnivore or human activity are very rare in Polish sites, this state of affairs and the predominance of can be partially explained by their specialisation in hunting on cave bears during the times when the other more typical food sources like ungulates were scarce. Spelaeoid bears U. spelaeus sensu lato were atypical prey for the Pleistocene lion, but when food was scarce, lions entered deeply into caves and hunted them (Argant, 1988, 1991, 2000; Diedrich, 2012). It was risky business facing this powerful animal in total darkness, and many lions were killed during such incidents, but not eaten by herbiv­ orous bears. Another explanation of the predominance of males is that larger and stronger male lions more often tend to hunt very large preys and went into the conflicts with other carnivores than females which ´n, 1997) resulted in higher injuries and mortality (Turner and Anto (Table 2). Among the Polish material of P. spelaea, almost all anatomical parts are represented, with the predominance of maxillar and mandibular fragments, isolated teeth, long bones, metapodials, carpals, tarsals and phalanges. Only four cranii are known, two of which are more complete (those from San River and Nied´zwiedzia Cave), and two neurocrania ´rna caves). Complete long (from Nied´zwiedzia and Wierzchowska Go bones are uncommon, and general state of preservation of most bones is relatively good, even those from open-air sites. The entire Polish ma­ terial of P. spelaea represents three different chronosubspecies, mor­ phometrical description and taxonomical classification of which are discussed in detail below.

120 350 160 1300 630 500 900 1000 700 800 600 1100 1100 1000 2000

5.2. Taxonomical analysis of P. spelaea from Poland 5.2.1. Panthera spelaea fossilis (von Reichenau, 1906) The most ancient remains of P. s. fossilis have been recorded from seven Polish sites dated back between 750 and 300 ka. Among the Polish findings of P. s. fossilis, the noteworthy is the find from Kozi Grzbiet which is the oldest Polish lion record. The material is not very numerous but well-preserved and informative. It includes right, complete upper canine and third phalanx, which metrically and morphologically are indistinguishable from those of P. spelaea. Both specimens belonged to very large lion, and certainly cannot be assigned to any another Euro­ pean big cat (Figs. 2 and 3). It is very big and robust canine of conical toothed cat, totally outstanding in shape from the canines of sabre tooth cats. It belonges to rather young individual, judging from the size to the male (Figs. 2 and 3). A wide nutrient foramen at the base of the massive root is visible. Elongated and sharply pointed crown holds well definied mesial and distal crests with clear and sharp relief. Alongside the most crown length running two parallel grooves, one on buccal and one on the lingual side. The only cat that could be mistaken for this canine from Kozi Grzbiet is Panthera gombaszoegensis. However, while similar morphologically, canines of this cat are far smaller than described specimen from Kozi Grzbiet. Even the extremely large jaguar’s canine from Vallonnet Cave, assigned previously to the lion (Lumley et al., 1988), was latter re-classified to be a P. gombaszoegensis (Hemmer, 2001). To tell the truth, in view of uniform shape of canines of pan­ therinae cats is is a hazardous matter try to classified based on single canine of intermediate size. This canine from Vallonnet Cave can belonges to the great male of P. gombaszoegensis or small female of P. s. fossilis. Anyway, in case of tooth from Kozi Grzbiet assignation is rather clear (Fig. 3). ´nossy (1969a, b) found that canines of P. s. fossilis are slender and Ja more elongated than canines of P. s. spelaea. Other previous studies showed that P. s. fossilis possesses less flattened and comparatively narrower canines as compared to P. s. spelaea (Sotnikova and Nikolsky, 2006; Barycka, 2008; Sabol, 2014). However, more recent studies (Argant and Brugal, 2017; Marciszak et al., 2019b) demonstrated opposite situation, where canines of P. s. fossilis are larger and more massive than those of P. s. spelaea. A broad revision based on European material dated on the last 750 ka showed, that from MIS 8 canines of P. spelaea shortened and narrowed, and it crown narrowes in relation to its length (Marciszak et al., 2019b). Great, upper canine from Kozi Grzbiet well corroborates with this trend (Fig. 3). Alongside with the British records from Pakefield and West Runton (Lewis et al., 2010), it may currently be the oldest known appearance of this chronospecies in Europe as well, not only in mainland (Parfitt et al., 5

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Fig. 2. Upper, right canine (KG/1/1) of Panthera spelaea fossilis from Kozi Grzbiet (layer 2a-c, MIS 19). Together with phalanx 3 this tooth represented the oldest Polish record of the Pleistocene lion and one of the oldest from Europe. Scale bar 30 mm.

Fig. 3. Graph showing the relationship between the crown length of C1 and the breadth of C1 crown of Panthera spelaea and Panthera gombaszoegensis. For citation see the chapter Comparative material.

2005; Lewis et al., 2010). Because the Polish material of this form was already studied and described in detail (Barycka, 2008; Marciszak et al., 2019b), and its morphological characterisation is limited up to the short description. P. s. fossilis from Polish Middle Pleistocene sites is characterised by very large size and stout build, and size of some individuals (e.g. lions from Południowa Cave or Draby) placed them among the largest ever known lions. Their incisors are relatively smaller and narrower, whereas

canines are exceptionally large and robust, and less flattened laterally. Large and broad P3 has short and high paracone with relatively weakly reduced additional cusplets (protocone and hypocone), and with strongly developed posterior cingulum which makes strongly marked ridge; there is a pronounced border between hypocone and cingulum ridge. Posterior part of the crown is strongly elongated on the lingual side towards the posterior root and creats a long, almost flat traingular surface. The P4 are metrically heterogeneous but morphologically 6

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

homogeneous. All the specimens have a triangular, well-developed, broad and relatively high protocone. The centrally positioned apex of this cusp is well developed. The cusp is oriented almost vertically to the entire tooth, and separated from the parastyle by a wide U-shaped val­ ley. The rounded and low parastyle is proportionally small, and closely appressed to the paracone. A shallow V-shaped valley separating both cusps is less developed. There is no preparastyle on the blunt anterior wall of the parastyle. The paracone is high and long, with three thick ridges running from the apex to the cusp base. The metacone is shorter and lower than the paracone, with a well-developed saddle in the middle part and a blunt posterior wall. The posterior half of the P4 is buccally and lingually strongly broadened in the middle part, and reveals almost straight buccal margin. In occlusal view, the P4 is short and broad, while in buccal view it is rather low. The p3 are oval teeth, characterised by relatively large linear di­ mensions as well as a well-developed, proportionally high and rounded protoconid located in the centre. Both additional cusplets, the paraconid and the hypoconid, are weakly developed and low. The paraconid is shifted anterolingually, while the medially positioned hypoconid is only slightly raised posterolingually. These cusps are closely associated with the main cusp and moderately separated from the surrounding cingulum which is better developed only on the anterior- and posterior walls. In occlusal view, the buccal margin is slightly convex, and the lingual margin of the posterior half is moderately expanded. The transition between the anterior and posterior parts is not strongly marked. The p4 are large and long, with a short and high protoconid and a robust crown. The buccal margin is straight, the lingual margin is noticeably expanded and forms a broad and smooth area. The accessory cusplets are equal in size, large, rounded and high. The hypoconid is poorly separated from the strong posterior cingulum, whereas the paraconid is well separated from the main cusp by a broad, V-shaped valley. Behind the hypoconid, a small but distinctive metaconid is visible. The m1 are oval and robust, with the breadth to length index exceeding 50. The paraconid is long and low, with the anterior margin shifted far anteriorly, whereas the pro­ toconid is slightly longer and much higher than the paraconid and its posterior wall is visibly sloped. The notch between main cusps is deep, narrow and V-shaped. The height, measured on the lingual side from the bottom of the cingulum to that of the notch between the paraconid and the protoconid, is greater. The crown is high and more elongated in lateral view. A well-developed bulge is situated on the lingual margin in the middle part of the crown, between the paraconid and the protoconid. In many specimens, it rises into a small cusp-like structure. The strongly developed lower margin of the cingulum rises considerably upwards, buccally and lingually in distal direction under the protoconid. A zigzag enamel structure formed by the lower margin of the cingulum is placed on the border between the protoconid and the talonid both buccally and lingually. The talonid is elongated and well-developed. The Polish material of P. s. fossilis was split into two main chro­ nostratigraphical groups. The first and older one (correlated to MIS 19–12) from Południowa and Tunel Wielki Caves is represented by specimens of P. s. fossilis possessing classical morphology which is comparable with type specimens from Mauer and Mosbach 2 (Barycka, 2008; Marciszak et al., 2019b). Among them is the oldest Polish and one of the oldest European records from Kozi Grzbiet. Specimens from the second period, dated to MIS 11-9, found in Draby, layers 19ad–18 of Bi´snik Cave, as well as Deszczowa and Wierzchowska G´ orna Caves, show somewhat more advanced morphology, with some more evolved fea­ tures. They differ from lions from the first period (MIS 19–12) in (1) narrower crowns P3 with proportionally shorter paracone and more reduced accessory cusplets and (2) p4 comparable in length with m1, with short and low protoconid, and more reduced accessory cusplets.

by Argant and Brugal (2017) based on the material from a few French sites and described as that characterising by smaller size (as compared with P. s. fossilis), less massive muscle attachments, triangular and shorter mandible ramus with shallower and shorter masseteric fossa, narrower canines, narrower P4 with almost straight buccal margin, narrower and smaller m1 with less developed bulge and zigzag enamel, as well as more gracile metapodials and calcaneus. In general, it was described as form in half way between P. s. fossilis and P. s. spelaea, and linking them in the evolutionary lineage of P. spelaea. Its occurrence was dated on MIS 8–6, and the presence of lions with intermediate morphology from Poland was already signalised (Marciszak and Stefa­ niak, 2010; Marciszak, 2014; Sabol, 2014). But only the recently made a detailed revision of all available Polish material of P. spelaea showed that some specimens should be attributed as advanced P. s. fossilis, and ´rna Cave can be only mentioned above material from Wierzchowska Go partially assigned to this form. We critically revisited the features pro­ posed by Argant and Brugal (2017) in the following part of this section. The mandibles of P. spelaea are much better represented in the Polish material, and at least five mandibles were classified as P. s. intermedia and 34 mandibles – as P. s. spelaea in the studied sample (Fig. 4). Despite the uniform morphology, particular characteristics are highly variable and it is not an easy task to determine reliable morphometrical features distinguishing these lion chronosubspecies. For P. s. fossilis Barycka (2008: 71) proposed the presence of gentle curvature of the lower mandible margin under m1. Sotnikova and Foronova (2014: 517) add that “it is distinguished from P. s. spelaea by a heavy built mandibular corpus with rectangular profile in the cheek teeth area, a deep, welloutlined and narrow anterior section of the masseteric fossa, and a large р4 supported by the large unreduced anterior root”. Argant and Brugal (2017) add also particularly larger size, robust build and powerful muscles attachment, triangular and proportionally shorter ramus with the vertical projection of the coronoid apophysis in front of the condyle, strongly developed angular process, deep masseteric fossa clearly extending forward under the m1, with a marked constriction, and the presence of two mental foramens on the mandible body. Relatively small size, moderate massiveness and well-developed muscles attachments, elongated, relatively narrow ramus with two parallel edges, the vertical projection of the coronoid apophysis behind the condyle, with masseteric fossa extending far forward under the rear of the m1, poorly or moderately developed constriction and the occur­ ence of two mental foramens on the mandible body (their shape is highly variable) characterised the mandible of P. s. intermedia (Argant and Brugal, 2017). The mandible of P. s. spelaea is metrically and morphologically very variable, and its massiveness and the development of muscle attach­ ments are usually correlated with dimensions (Fig. 5). Ramus of the mandible is fairly elongated, relatively narrow, with two parallel edges and the masseteric fossa not extending under the m1, and with weakly developed constriction (Argant and Brugal, 2017). Barycka (2008: 71) added also a rather straight lower margin of the mandible (Figs. 4 and 5). We test these features on the Polish material. It is shown that most of those characteristics do not apply in the studied sample. Firstly the size, mandible body masiveness and the level of development of muscle at­ tachments. The revision of Polish records of P. s. fossilis showed that all analysed mandibles belonged to large individuals, sometimes of immense size (Marciszak et al., 2019b). However, all these lions were assigned as males, and great massiveness and large size of the specimen mentioned by previous authors, e.g. Sotnikova and Foronova (2014: 517): “heavy built mandibular corpus” or Barycka (2008: 117): “massive mandible with a convex lower board, suggesting Middle Pleistocene characteristics” should be rather correlated with the sexual dimorphism. Already in 1906 von Reichenau (1906: 305) pointed out that the mandible of lioness can be easily distinguished from that of male in its gracile build. The great sexual dimorphism is characteristic for large pantherinae

5.2.2. Panthera spelaea intermedia Argant and Brugal (2017) The second form which was recognized in the Polish material of P. spelaea is P. s. intermedia, whose occurrence was so far partially ´rna Cave. This form was erected confirmed only from Wierzchowska Go 7

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Fig. 4. Mandibles of the Pleistocene lion Panthera spelaea from the Silesian (southern Poland) sites. 1 - Panthera spelaea fossilis from Wschodnia Cave (JW/Pss/1). Panthera spelaea spalea: 2 - left from Nied´zwiedzia Cave (JNK/Pss/17x); 3 - right from Nied´zwiedzia Cave (JNK/Pss/18x, the same specimen as JNK/Pss/17x); 4 right from Nied´zwiedzia Cave (JNK/Pss/16); 5 - right from Naciekowa Cave (JNW/Pss/20); 6 - left from Sosnowiec-Milowice (MIL/M/Pss/1); 7 - right from Pyskowice (MUZ PIG 1742.II.1); 8, right from Węgry (WEG/M/Pss/1). Lions 1–5 were found in cave sediments, wheres those 6–8 came from open-air sites. All individuals are shown in the same scale, scale bar 50 mm; a - buccal view; b - lingual view; c - occlusal view.

cats, and it is even more pronounced in the Pleistocene lion than that in the extant lion (Hemmer, 2011). The relative percentage in particular examined groups, populations and sites may differ greatly. For this reason, the general analysis, means of samples including both sexes are apt to be misleading, and such comparisons should be made between groups of the same sex (Kurt´ en, 1985: 118). For this reason, mandibles of the Middle and Late Pleistocene lions were sexed using two factors: m1 length and height of mandible body measured after m1. For this purpose, only mandibles with preserved m1 of fully adult individuals were used, since extant lions could be sexed over two and more years old (Smuts et al., 1978; Turner, 1984). Metrical data showed similar values where ranges of variability of all three chronosubspecies generally overlapped. Slightly higher values of P. s. fossilis seem to be a result of the limited number of individuals and higher percentage of males in the analysed group. In turn, smaller values of P. s. intermedia males result from that this form was described on the French material as rather small lion (Argant and Brugal, 2017). Additionally, some populations are of interglacial age, so Bergman’s rule also should be taken into consider­ ation. In this context we agree wth Diedrich (2017) that glacials and interglacials may be among the main factors affecting the variability of Pleistocene lions, even if the proposed scenario is reconstructed too simply. However, terminology like “P. l. s. intermedia”, „P. l. s. brachyg­ nathus” or “P. l. s. maximus” used by this author is an unnecessary complication of taxonomy and confusing the terminology. All these new

“forms” should be considered as a synonimies for three widely used and accepted chronosubspecies established long time ago. Concluding, if any lion’s mandible exceeds 57 mm in height of body mandible measured after the m1 and any lion’s m1 exceeds 29.5 mm in crown length, almost exclusively belonged to male. A good example is here the male from Nied´zwiedzia Cave, where two mandibles (left and right) assigned to this individual have massive body mandible, with height measured after the m1 of ca. 70 mm, even that they are not particularly large (estimated total length ca. 275 mm). The next feature proposed by Argant and Brugal (2017) was fairly elongated, relatively narrow mandible ramus with two parallel edges, and the masseteric fossa which does not extend under the m1, and with weakly developed constriction. Observation of the large series of Polish Pleistocene lions showed that these features do not applied for this material (Figs. 4 and 5). Most of the Polish lions have relatively short and massive mandible body with rounded outline of the anterior margin of deep and long masseteric fossa which extended fairly anteriorly, and reached almost para- and protoconide border in Nied´zwiedzia individ­ ual (Fig. 4/2a and 3a). This specimen shows also another feature of the fully grown and mature lion, a noticeably large and massive symphysis which exceeds in size and massiveness the symphysis of similarly large another male mandible from Nied´zwiedzia Cave (Fig. 4/4a-c), which undoubtedly belonges to younger individual. This showed that the developmental stage of the aforementioned features is for sure 8

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Fig. 5. Mandibles of the Pleistocene lion Panthera spelaea from Wierzchowska G´ orna Cave. Panthera spelaea intermedia: 1 - left (MF/320/6801); 2 - left (MF/320/ 6793); 3 - left (MF/320/6796); 4 - left (MF/320/6795). Panthera spelaea spelaea: 5 - left (MF/320/6806); 6 - left (MF/320/6805); 7 - right (MF/320/6800); 8 - left (MF/320/6799); 9 - left (MF/320/6809); 10 - right (MF/320/6710); 11 - left (MF/320/6802); 12 - left (MF/320/6889; 13 - right (MF/320/6889); 14 - right (MF/ 320/6804). All individuals are shown in the same scale, scale bar 100 mm; a - buccal view; b - lingual view; c - occlusal view.

correlated with ontogenetic growth, and they are sharper pronounced and better developed in older individuals. In addition, Silesian lions also well illustrated a significant sexual dimorphism where the only found lioness from Sosnowiec-Milowice (Fig. 4/6a-c) is characterised by much smaller dimensions, narrower mandibular body and less massive sym­ physal part of mandible. ´rna Cave, the situa­ When analysed material from Wierzchowska Go tion is more complicated (Fig. 5). All four mandibles classified as P. s. intermedia belonged to large and robust individuals with relatively massive and short mandible body similar to those in the Silesian lions (Fig. 5/1–4). Between the individuals assigned to P. s. spelaea with proportionally long and massive mandibular body (Fig. 5/6a-c and 14ac), most lions have elongated and quite slim mandibular body with triangular outline of the anterior margin of the masseter fossa which reaching the m1 protoconid (Fig. 5/9–13). Most of the Polish Late Pleistocene lions have rather straight lower margin of the mandible (Figs. 4 and 5), which is in agreement with Barycka (2008). However, the slightly convex lower margin under the m1 is not uncommon, as can be seen in Silesian lions from Naciekowa Cave (Fig. 4/5a-b) or Węgry open-air site (Fig. 4/8a-b). Besides, the number (usually two large), shape and distances between the mental foramens are too variable for being more or less informative. Summa­ rising the metrical and morphological valuability of the mandible in taxonomical analysis, it should be noted that it is rather weak. Some previous researchers tried to analyse mandibular morphology, espe­ cially when the lack of the teeth excluded dental analysis, e.g. Sotnikova and Foronova (2014). However, analysis of the Polish material showed that in fact none of such features can be used for distinguishing different lion chronosubspecies. Features like height and thickness of the mandibular body, the shape of masseteric fossa and mandibular ramus, and the shape of lower mandibular margin should be regarded as an intraspecific variability. They are sexually dimorphic and ontogeneti­ cally highly variable, and their taxonomical usefulness is rather poor.

5.2.3. Panthera spelaea spelaea (Goldfuss, 1810) The last and more numerous lion form recorded from Polish sites is P. s. spelaea which was an integral member of the Late Pleistocene Eurasian “mammoth faunas”. It was recorded in 41 Polish sites, dated at MIS 5–3. Material of this form is also the most numerous and informative, and among them there are also the most impressive findings of this species (e.g. complete skulls). The cranium of P. s. spelaea differs from that of P. s. fossilis in a proportionally shorter and broader snout, larger orbits, better developed temporal and mastoid areas as well as less inflated tympanic bulla, broader incisors tooth row and narrower upper teeth with better cutting projection (P3 and P4). The index of rostrum breadth measured on canines to the total skull length for the Pleistocene lion is the following: MIS 19–12 – 25.3 (24.5–25.9, n = 3), MIS 11–7 – 27.2 (26.4–27.9, n = 6), MIS 6–5 – 28.5 (25.4–30.6, n = 8), and MIS 4–2 – 30.4 (27.2–31.5, n = 20). There are only two complete skulls in the studied Polish material (from Nied´zwiedzia Cave and San River), and both belong to large adult males (Fig. 6). In addition, there is also a second neurocranium of the large male from Nied´zwiedzia Cave (its size and proportions are almost identical to those mentioned above), left ´rna maxilla and a neurocranium of a lioness from Wierzchowska Go Cave. Since both complete skulls from Nied´zwiedzia Cave were previ­ ously studied and described (Wiszniowska, 1978; Barycka, 2008) and San River (Marciszak et al., 2014) we prepare only a short summary. Three among main morphological characters of the skull differs P. s. spelaea from P. s. fossilis: a broad and short rostrum, short and wide temporal region and strongly convex frontal profile (in lateral view) with well-marked concavity in the medium of frontal part. These fea­ tures are well marked in both lions (Figs. 6 and 7). It is especially well visible when compare two very similar in size lions from Az´e (MIS 11; Argant and Brugal, 2017) and Nied´zwiedzia Cave (MIS 4). The lion from Nied´zwiedzia Cave has proportionally broader snout (index of C1–C1 breadth to the total skull length in Nied´zwiedzia lion is 30.4, in Az´e lion is 26.9), wider incisors tooth row (the index of I3–I3 breadth to the total skull length in Nied´zwiedzia lion is 13.6, in Az´ e lion – 11.8), larger orbits (the index of maximal length of orbit to the total skull length in 9

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Fig. 6. Skulls of extinct and extant lions. Panthera spelaea spelaea: 1 - ♂ from San river (NKM-00664); 2 - ♂ from Nied´zwiedzia Cave (JNK/Pss/1); 3 - ♀ from Wierzchowska G´ orna Cave (MF/320/7059). Panthera leo: 4 - ♂ from East Africa; 5 - ♂ from North Africa. All individuals showed on the same scale, scale bar 120 mm; a - dorsal view; b - ventral view.

Fig. 7. Total skull length and the least breadth at the canine alveoli of Panthera spelaea male skulls. For citation see the chapter Comparative material.

Nied´zwiedzia lion is 21.0, and 19.3 in Az´e lion), as well as larger tym­ panic chamber. However, despite of Late Pleistocene age (ca. 60 ka BP), the skull from Nied´zwiedzia Cave still holds some primitive features similar to those in late Middle Pleistocene Az´ e calvarium. Among them, there are smaller and less convex tympanic bullae and moderately developed temporal and mastoid areas. Such an admixture of primitive and advanced features isn’t uncommon in Panthera spelaea. It is also a proof how dynamically this species evolved and how complicated was this process.

5.3. Evolution of the cheeck teeth of P. spelaea from Poland P. s. spelaea has a narrower P3 with proportionally longer, lower and more oval main cusp, the paracone with straight buccal margin and less expanded convexity on the lingual posterior part (Fig. 8). The index of the posterior breadth to the total crown length is ca. 55 for P. s. fossilis and ca. 50 for P. s. spelaea. The size and shape of two smaller additional cusplets, anterior protocone and the posterior hypocone, are more var­ iable. The protocone is larger and more rounded and stronger shifted 10

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Fig. 8. P3 of the Pleistocene lion Panthera spelaea from the Polish late Middle and Late Pleistocene sites. Panthera spelaea intermedia (from Wierzchowska G´ orna Cave): 1 - MF/320/6808; 2 - MF/320/6794. Panthera spelaea spelaea: 3 - Wierzchowska G´ orna Cave (MF/320/6797); 4 - Wierzchowska G´ orna Cave (MF/320/6811); 5 - Wierzchowska G´ orna Cave (MF/320/6789); 6 - Bi´snik Cave, layer 9 (JB/Ps/294); 7 - San River (NKM-00664); 8 - San river (NKM-00664); 9 - Nied´zwiedzia Cave (JNK/Pss/1); 10 - Nied´zwiedzia Cave (JNK/Pss/1); 11 - Nied´zwiedzia Cave (JNK/Pss/200); 12 - Nied´zwiedzia Cave (JNK/Pss/236); 13 - Nied´zwiedzia G´ orna Cave (JNG/Pss/3). All individuals are shown in the same scale, scale bar 15 mm; a - buccal view; b - lingual view; c - occlusal view.

anterio-lingually, while the hypocone is smaller, higher and placed slightly more posterobuccally. The distinction between the paracone and the hypocone, as well as between the hypocone and the posterior cingulum, is well pronounced, and the posterior cingulum is weakly developed. The protocone is placed more centrally and varies strongly from the minute to small one, and totally disappeared in many Late Pleistocene specimens (Dawkins and Sandford, 1866; Schütt, 1969a). The hypocone is smaller and usually placed almost exactly in the middle of the crown, just after the paracone (Fig. 8). The P4 of P. s. spelaea possesses triangular, moderately-developed, relatively narrow and low protocone with centrally positioned top (Fig. 9). In occlusal view, the tooth is elongated and narrow, with convex buccal margin and sharply ended crown thickening only buccally. In buccal view, the tooth looks higher. More reduced and low protocone is shifted more posterolingually and has a less marked top. The parastyle is large, high and oval. It is also well separated from the paracone by a deep and sharply pointed valley, and the constriction between the par­ astyle and the paracone is notable in occlusal view. The paracone is shorter and lower, while the metastyle is slightly longer than the para­ cone (Fig. 10). The index of metastyle length to the paracone length is ca. 95 for P. s. fossilis and ca. 105 for P. s. spelaea. It is confirmed the general trend to shortening of the paracone and elongating of the

metastyle in course of time. According to Sotnikova and Nikolsky (2006), the P4 of P. s. fossilis is characterised by the absence of preparastyle which appeared in the late Middle Pleistocene and additionally P4 of P. s. spelaea tends to develop an incipient cingulum, anterior to the preparastyle. Contrary to that, Hanko (2007) proposed the lack of preparastyle in P. s. spelaea. He found that this small cusplet (if present) is always poorly developed in P. s. fossilis. Analysis of the large series of P4 from Polish sites showed that the presence/absence and/or the developmental stage of this cusplet shows a great variability (Fig. 9). The general trend is the lost of pre­ parastyle and the development of additional, incipient cingulum in front to the parastyle. Schütt (1969a) found a minor preparastyle (named by her as ectoparastyle) in the specimens from Rübeland, Wallertheim, Weinberg Cave, Vypustek Cave and in four (of five) specimens from Mauer, while it was absent in the individuals from Steiermark and Einchorn Cave. Schütt (1969a) also found that individuals from Einc­ horn Cave and both P4 from Mauer skull have well developed antero­ lateral wall of the parastyle which forms a well-visible bulging. In the case of Repolust Cave, she noted the presence of an incipient cingulum in ´ (2007) mentioned the presence of poorly front of the parastyle. Hanko ¨llo ¨s 2 and Solymar in­ developed preparastyle in case of Vertesszo dividuals (somewhat better developed in specimen from Solymar), while 11

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Fig. 9. P4 of the Pleistocene lion Panthera spelaea from the Polish late Middle and Late Pleistocene sites. Panthera spelaea intermedia from Wierzchowska G´ orna Cave: 1 - MF/320/6766, 2 - MF/320/6769; 3 - MF/320/6770; 4 - MF/320/6771. Panthera spelaea spelaea: 5 - San river (NKM-00664); 6 - San river (NKM-00664); 7 - Bi´snik Cave, layer 7 (JB/Ps/3-3); 8 - Bi´snik Cave, layer 7 (JB/Ps/302); 9 - Nietoperzowa Cave, layer 10 (MF/6253); 10 - Wierzchowska G´ orna Cave (MF/320/6797); 11 Wierzchowska G´ orna Cave (MF/320/5825); 12 - Komarowa Cave, layer F (KO/F/1); 13 - Komarowa Cave, layer F (KO/F/2); 14 - Nied´zwiedzia Cave (JNK/Pss/1); 15 - Nied´zwiedzia Cave (JNK/Pss/1); 16 - Nied´zwiedzia Cave (JNK/Pss/204); 17 - Nied´zwiedzia Cave (JNK/Pss/205); 18 - Nied´zwiedzia G´ orna Cave, layer 3 (JNG/Pss/ 4). All individuals are shown in the same scale, scale bar 20 mm; a - buccal view; b - lingual view; c - occlusal view.

Fig. 10. Graph showing the relationship between the paracone length of P4 and the metastyle length of P4 of Panthera spelaea. For citation see the chapter Comparative material.

the specimens from Kiskevely and Igric Pestere show the absence of this cusp. Sabol (2011a) also does not found the preparastyle in the case of Slovakian localities (Medvedia Jaskyna and Tmava Skala Jaskyna). In the case of Polish specimens, this cingulum swelling is well developed in those of Late Pleistocene age (Nied´zwiedzia, Naciekowa, Nietoperzowa, and Wierzchowska G´ orna Caves). Individuals from San River and Nied´zwiedzia Cave showed a very well-developed additional cingulum on P4, clearly separated from the parastyle; however, bulgening is also

well marked. Von Reichenau (1906) described an isolated P4 from Mauer as pro­ portionally short and wide, while Freudenberg (1914) noticed quite low crown. Hank´ o (2007) agreed with Freudenberg’s opinion, but according ´ 2007: 39): “Freudenberg’s observation seems to be to him (Hanko convincing, but cannot be verified by the measurements taken from the studied specimens because the adequate teeth of these two taxa have almost the same proportions”. Most authors also noticed a greater width 12

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

´nossy, 1990; Hanko ´, 2007; of the P4 measured at the protocone (Ja Barycka, 2008). This narrowing of the anterior part of P4 is correlated with the reduction in size and height of the protocone which became shorter from P. s. fossilis to P. s. spelaea, lower and more rounded instead of more triangular like in the case of P. s. fossilis. In P. s. spelaea, the protocone is shifted more posterolingually, and the transition between anterior walls of the parastyle and protocone is better pronounced. Posterior part of the crown in P. s. fossilis is strongly thickened buccaly and lingually, while in P. s. spelaea the thickening is visible only on the buccal side. Therefore, the P4 of P. s. spelaea look narrower and more slender in occlusal view than that of P. s. fossilis. Middle Pleisto­ cene lions revealed also more straight buccal margin of the porterior part, while it is slightly convex buccaly in the case of P. s. spelaea. Great variability of p3 does not provide any significant features allowing to distinguish different lion forms. Size, height and the level of development/reduction of main and additional cusps as well as

massiveness of the p3 crown do not differ and strongly overlap in different lion chronosubspecies (Fig. 11). Schütt (1969a) noticed that distinctions in the dentition between the Middle and Late Pleistocene lions are the most visible on p4, and the mean difference are significant even though the ranges of variability generally overlap. She found that P. s. spelaea has a proportionally longer and lower protoconid of the p4. Kurt´en (1960) described more massive build of the p4 in P. s. fossilis as a subspecies feature. Also, Schütt (1969a) and Hemmer (1974) found narrower p4 for P. s. spelaea. Two smaller additional cusps, anterior paraconid and posterior hypo­ conid (metaconide sensu Barycka, 2008) are in most of the specimens (all from Mosbach and mostly from the Late Pleistocene localities) equal in ´, 2007; size (Kabitzsch, 1960; Schütt, 1969a; Hemmer, 1974; Hanko Barycka, 2008). In two individuals, the paraconid is very small, while both specimens from Einchorn Cave and two Late Pleistocene have the paraconid larger than hypoconid (Schütt, 1969a). Generally, the trend

Fig. 11. Comparison of p3 of the Pleistocene lion Panthera spelaea from the Polish late Middle and Late Pleistocene sites. Panthera spelaea intermedia (from Wierzchowska G´ orna Cave): 1 - left MF/320/6792; 2 - left MF/320/6796. Panthera spelaea spelaea: 3 - left from Wierzchowska G´ orna Cave (MF/320/6806); 4 - right from Wierzchowska G´ orna Cave (MF/320/6804); 5 - left from Komarowa Cave, layer F (KO/F/19); 6 - right from Bi´snik Cave, layer 8 (JB/Ps/306); 7 - right from Zegar Cave, layer 13 (JZg/Pss/10); 8 - left from Nied´zwiedzia Cave (JB/Pss/17); 9 - right from Nied´zwiedzia Cave (JB/Pss/18); 10 - right from Nied´zwiedzia Cave (JB/Pss/16). All individuals are shown in the same scale, scale bar 10 mm; a - buccal view; b - lingual view; c - occlusal view. 13

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

to reduction both additional cusps can de detect, but this process was diminished by a great variability, although even extreme reduction up to the lost of the hypoconid in case of the specimen from Molodova 5 has been noted (Sotnikova and Foronova, 2014). Schütt (1969a) also noticed that the posterior cingulum is less developed in P. s. spelaea and the border between hypoconid and cingulum in less marked. The p4 of Polish lions from the late Middle and Late Pleistocene sites also showed a tendency of crown narrowing and elongating of the main ´rna cusp. Specimens classified as P. s. intermedia from Wierzchowska Go Cave still showed some primitive features, like large size, broad crowns, and large additional cusplets (Fig. 12/1–4). The p4 of P. s. spelaea are on average smaller, their main cusp is longer and higher and additional cusplets are more reduced (Fig. 12/5–16). The posterior part of the crown is less expanded and the transition between the anterior and the posterior halves of the crown is less marked (Fig. 12). The p4 of P. s. ´rna Cave are especially massive as intermedia from Wierzchowska Go compared with p4 of P. s. intermedia from French sites (Fig. 13). The m1 is probably the best studied tooth, and most authors mentioned a pronounced and broader crown of the Middle Pleistocene lions. The general trend to narrowing of the crown of m1 is observed in the cave lion lineage evolution and generally for P. s. spelaea m1 width to the length ratio is ca. 50. However, in the large series from Europe lower carnassials with ratio higher than 50 (usually 51–53) also appeared. Most of those lions are dated on the last interglacial and the early part of the last glacial (Crusafont-Pairo and Casajuana, 1950; Rakovec, 1951; Schütt and Hemmer, 1978; Dufour, 1989; Aleekseva, 1990; Baryshnikov and Boeskorov, 2001). They represent so called “primitive cave lion”, P.

s. spelaea with some still existed primitive features (Fig. 14). No differences in length between the paraconid and protoconid distinguishing P. s. fossilis and P. s. spelaea have been found, while they differ in height of both main cusps. The paraconid in P. s. fossilis is long and low cusp, with anteriory margin shifted far anteriorly, while the protoconid is much higher and its posterior wall is visibly sloped. The difference in height between both cusps in P. s. spelaea is less pro­ nounced, the notch between them is stronger developed, and the angle is more opened. The anterior margin of the paraconid does not rise so anteriorly, and the posterior wall of the protoconid falls almost verti­ cally. It makes better cutting surface of protoconid exposition and strongly correlated with the evolution to meat slicing process. Also the third measurement (so called “incisors height”, taking on the lingual side from the bottom of the cingulum to that of the notch between the paraconid and protoconid) indicates proportionally higher index for mid Middle and late Middle Pleistocene lions. In general, the m1 crown in the case of P. s. spelaea looks more compact on lateral view and lower than that of P. s. fossilis. The next distinction is a development of lingual bulgae located in the middle part of the crown between the paraconid and protoconid. This cingulum swelling is well developed in P. s. fossilis and in most of the Middle Pleistocene specimens it rises even in small cusp-like formation. In the course of time, bulgening decreased and it is less marked in late Middle Pleistocene lions. In P. s. spelaea, this bulge is in most cases poorly developed and the lingual margin in many individuals is almost straight (Fig. 14). Strongly developed lower margin of cingulum in P. s. fossilis rises

Fig. 12. The p4 of the Pleistocene lion Panthera spelaea from the Polish late Middle and Late Pleistocene sites. Panthera spelaea intermedia (from Wierzchowska G´ orna Cave): 1 - MF/320/6785; 2 - MF/320/6786; 3 - MF/320/6796; 4 - MF/320/6783. Panthera spelaea spelaea: 5 - Wierzchowska G´ orna Cave (MF/320/6788); 6 Wierzchowska G´ orna Cave (MF/320/6791); 7 - Wierzchowska G´ orna Cave (MF/320/6808); 8 - Wierzchowska G´ orna Cave (MF/320/6804); 9 - Naciekowa Cave (JNW/Pss/111); 10 - Naciekowa Cave (JNW/Pss/1); 11 - Węgry (WEG/M/Pss/1); 12 - Nied´zwiedzia Cave (JNK/Pss/17); 13 - Nied´zwiedzia Cave (JNK/Pss/18); 14 Nied´zwiedzia Cave (JNK/Pss/16); 15 - Nied´zwiedzia Cave (JNK/Pss/198); 16 - Sosnowiec-Milowice (MIL/M/Pss/1). All individuals are shown in the same scale, scale bar 20 mm; a - buccal view; b - lingual view; c - occlusal view. 14

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Fig. 13. Graph showing the relationship between the crown length of p4 and the posterior breadth of p4 crown of Panthera spelaea. For citation see the chapter Comparative material.

number of the material of this form should be kept in mind. Similarly, slight differences in the morphology of muscle joints and articular sur­ faces should be assigned as an intraspecific variability. It can be concluded that metapodials, carpals and tarsals are actually almost undistinguishable morphologically. Metapodials from the Polish sites are also a good example of great sexual dimorphism and general dwarfing size in the evolution of P. spelaea (Fig. 16). During the Middle Pleistocene, the size of the Pleis­ tocene lion varied only slightly, while the real size changes began since the Late Pleistocene (Marciszak et al., 2019b). 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”. This gradual process shrank accordingly since the last glacial, which espe­ cially middle part is characterised by dramatic populations declination and their genetic variability reduction (Barnett et al., 2009). Somewhere between 48 and 45 ka, Pleistocene lion populations rapidly declined and their genetic variability was dramatically reduced, which affected in a genetic bottleneck (Barnett et al., 2009). This period, due to a massive stadial within MIS 3, affected significantly in haplotype loss, strongly pronounced size declination and massiveness decreasing. It lasted by MIS 3 and 2 and within it size drops sharply. Among mod­ erate sized specimens, like those from Siegsdorf or Zoolithen Cave (Dietrich, 1968; Gross, 1992; Diedrich, 2008; Marciszak et al., 2014, 2019b), comparable in the largest recent lions, also very small (some­ times even regarded as dwarf forms) lions also appeared for the first time. Apart of smaller dimensions, they were also characterised by much less stouter posture and in bone dimensions they are generally closer to the African lion than to the Late Pleistocene form (Altuna, 1972; Clot et al., 1984; Pautret-Homerville et al., 2011). Even against the fact that most of those extremely small lions were lionesses, their dimensions are mostly comparable medium-sized specimens of modern African lions (Fig. 16). The presence of smaller, lightly build lions is regarded as a local response for the harsh climatic conditions, lower than in other ´k, 2009). regions prey abundance and competition pressure (Lupta The last feature of P. s. fossilis, pronounced by previous authors, e.g. Kurt´en (1960, 1968), Schütt (1969a), Schütt and Hemmer (1978), Argant (1988, 1991, 2000), Argant et al. (2007) and many others, was the greater size. The question of timing and the nature of size fluctua­ tions of Pleistocene lions was investigated by comparing the dental, cranial and postcranial dimensions from a number of sites by means of percentage diagram (Fig. 17). This method allows comparing each

considerably upwards distally under the protoconid buccaly and lingually, while this enamel curve in P. s. spelaea is poorly developed and present usually only on the buccal side. It does not rise on the lingual side and creates almost straight line. Additionally, Schütt (1969a) noticed the presence of a zigzag enamel structure forming by lower margin of the cingulum placed on the border between the protoconid and talonide in P. s. fossilis. This feature has been found in all specimens from Mosbach and Mauer, but it is also present in some late Middle ¨llo ¨s 2, Dechen Cave, Hunas, Bi´snik Pleistocene like those from Vertesszo ´rna Cave. Among Cave and two specimens from Wierzchowska Go others, it also has been found (although less developed) in specimens from Wallertheim, Repolust Cave and Gross-Bieberau. This structure in P. s. spelaea is strongly reduced and present only as a minute on the lingual side (Schütt, 1969a; Barycka, 2008). ´ (2007: 34) claimed: “The talonide of P. s. fossilis is the most As Hanko developed of all examined fossil species”. In P. s. spelaea, the stage of development of the talonide slightly varies, but this part of the tooth is always much more reduced than in the case of P. s. fossilis. Minute metaconid was noticed in a single specimen from Gross-Bieberau and Zoolithen Cave (Schütt, 1969a) as well as in a single individual from Wierzchowska G´ orna and Bi´snik caves. Occasionally the presence of this rudimentary tooth element should be considered as anomaly and should not be regarded as a distinguishing feature. ´rna Cave, determined The Polish specimens from Wierzchowska Go as P. s. intermedia, still hold many primitive features like the large size, moderately reduced lingual bulge and the talonid (Fig. 14/1–8). Their crowns are still broad, and compared to other Polish lions are particu­ larly wide, similarly to those in French specimens (Fig. 15). In addition, especially Late Pleistocene specimens are characterised by an elongated and narrower crown which looks higher in lateral view. The evolu­ tionary tendency to the elongating of m1 is visible (Fig. 15). 5.4. Size changes over time of P. spelaea from Poland Analysis of more numerous postcranial bones provided less valuable information, and showed a greater stage of variability than those of cranial or dental material (Fig. 16). The massiveness of metapodials, counted as a minimal median shaft breadth to the total bone length index, showed no particular differences between P. s. fossilis and P. s. spelaea, except the metatarsal II, III and V, where the metapodials of P. s. fossilis are more robust (Marciszak et al., 2019b). However, the limited 15

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Fig. 14. The m1 of the Pleistocene lion Panthera spelaea from the Polish late Middle and Late Pleistocene sites. Panthera spelaea intermedia (from Wierzchowska G´ orna Cave): 1 - MF/320/6801; MF/320/6795; 3 - MF/320/6773; 4 - MF/320/6775; 5 - MF/320/6781; 6 - MF/320/6782; 7 - MF/320/6780; 8 - MF/320/6779. Panthera spelaea spelaea: 9 - Wierzchowska G´ orna Cave (MF/320/6776); 10 - Wierzchowska G´ orna Cave (MF/320/6809); 11 - Wierzchowska G´ orna Cave (MF/320/6806); 12 Wierzchowska G´ orna Cave (MF/320/6804); 13 - Wierzchowska G´ orna Cave (MF/320/6805); 14 - Wierzchowska G´ orna Cave (MF/320/6799); 15 - Komarowa Cave, layer F (KO/F/3); 16 - na Gołąbcu Cave (MF/2233); 17 - Hureczko (KP/389); 18 - Węgry (WEG/M/Pss/1); 19 - Sosnowiec-Milowice (MIL/M/Pss/1); 20 Nied´zwiedzia Cave (JNK/Pss/17); 21 - Nied´zwiedzia Cave (JNK/Pss/18); 22 - Nied´zwiedzia Cave (JNK/Pss/206); 23 - Nied´zwiedzia Cave (JNK/Pss/207); 24 Nied´zwiedzia Cave (JNK/Pss/208). All individuals are shown in the same scale, scale bar 20 mm; a - buccal view; b - lingual view; c - occlusal view.

measurement of the fossil sample with a standard set of measurements. In this case, it is a sample of measurements of the recent African lion, compiled from Gross (1992) and our own measurements. The percent­ age diagram is advanced because it allows to compare even too small samples. The samples from the available sites with remains of the steppe lion are arranged in a temporal series spanning the Middle Pleistocene. The diagram shows that measurements actually overlap and did not changed significantly throughout the whole Middle Pleistocene (Fig. 17) (Barycka, 2008; Marciszak et al., 2014). Summarising the analysis of P. spelaea material from the Polish sites, most of trends previously mentioned by many other authors, e.g. Reichenau (1906); Schütt (1969a); Hemmer (1974); Kurt´en and Pou­ lianos (1977, 1981); Schütt and Hemmer (1978); Bishop (1982); Riedel ´nossy (1990); (1982); Toepfer (1983); Argant (1988, 1991, 2010); Ja Sala (1990); Guˇzvica (1998); Garcia (2003); Bona (2006); Testu (2006); ´ (2007); Barycka (2008); Lewis et al. (2010); Argant et al. (2007); Hanko

Marciszak et al. (2014); Sabol (2014) have been confirmed. The most valuable data for the biochronological analysis were found mostly on dental material, where multiple evolutionary changes were found. For P3, narrowing of the crown, especially the anterior part, paracone tends to be longer but lower, and the reduction of additional cusplets and cingulum was observed. In P4, the crown narrowing, paracone tends to be shorter and lower, while the metastyle is elongated, enlargening of the parastyle and reduction of the protocone, development of the pre­ parastyle and incipient cingulum in front of the parastyle were noticed. In p3, decreasing the overall tooth size, narrowing of the crown and reduction of additional cusplets was found, alongside with protoconid tending to be lower and shorter. In p4, analysis showed shortening of the total length, elongating of the protoconid which tends to be lower, and narrowing of the crown. In m1, narrowing and lengthening of the crown was observed together with the paraconid tending to be shorter and lower; median bulge, talonid and zigzag enamel structure tend to be lost. 16

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Fig. 15. Graph showing the relationship between the crown length of m1 and the breadth of m1 crown of Panthera spelaea. For citation see the chapter Compar­ ative material.

Fig. 16. Example of great sexual dimorphism in the Cave lion P. s. spelaea metacarpals and metatarsals. Mtcp III: 1 - Wierzchowska G´ orna Cave (MF/320/5665); 2 Nied´zwiedzia Cave (JNK/Pss/194); 3 - Naciekowa Cave (JNW/Pss/51). Mtcp V: 4 - Oborniki (MRR-687/G; Ob/2013/8.F3); 5 - P´ ołnocna Du˙za Cave (JPD/Pss/1); 6 Naciekowa Cave (JNW/Pss/63). Mtts II: 7 - Wierzchowska G´ orna Cave (MF/320/5660); 8 - Wierzchowska G´ orna Cave (MF/320/4649); 9 - Nied´zwiedzia Cave (JNK/ Pss/197). Mtts V: 10 - Skarszyn; 11 - Wierzchowska G´ orna Cave (MF/320/4656); 12 - Nied´zwiedzia Cave (JNK/Pss/166). All individuals are shown in the same scale, scale bar 60 mm. Specimens 4–6 and 10–12: a - dorsal view; b - ventral view. Specimens 1–3 and 7–9: a - outer view; b - inner view.

17

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Fig. 17. Measurement comparison of P. spelaea from Poland with recent P. leo (according to Gross, 1992 and own measurements). The fossil individuals are plotted as a percentage deviation from the mean of the modern sample (vertical line), and with the localities arranged in an approximate temporal sequence (dots mean cranial and dental material, circles mean postcranial material). For citation see the chapter Material.

Finally, the general size decreasing was also confirmed.

Lewis et al., 2010; Marciszak et al., 2014, 2019b; Sabol, 2014). But the Asiatic records are very scarce. However, among them one is the most important in the reconstruction of the evolutionary history of P. spelaea. Sotnikova and Foronova (2014) described a left hemimandible of particularly large lion from the Kuznetsk Basin dated to the late Early Pleistocene. The age was previously questioned by Hemmer (2011), however the explanation made by authors may confirm the true age of this find. If so, it is the oldest record of P. spelaea outside Africa, and the proof for the expansion of this species from Africa to Eurasia via the Asian continent (Sotnikova and Foronova, 2014). The eastern migration into Europe can be additionally documented by a new find from the Polish site Kozi Grzbiet which is now the oldest European mainland lion record, and one of the oldest in Europe. The finds (C1 and ph III) came from the unit 2, layers 2a-c, bone-bearing layer composed principally of yellowish-brown to dark brown sandy loams. The age of this horizon was established on the basis of the fluorochloro-apatite (FCL/P) and collagene methods at ca. 750–700 ka

6. Discussion 6.1. The earliest polish record of P. spelaea P. spelaea has a complex history, with three subspecies described within the lineage and some characteristic local populations. It is widely accepted that this species originated in Africa and dispersed northward into Eurasia, but the time of this migration event is not so clear. In the past it was thought that it took place during the early Middle Pleisto­ cene, before 0.75 Ma, as was documented from the British finds from Pakefield (Lewis et al., 2010; Hemmer, 2011). This early Middle Pleis­ tocene and later mid Middle Pleistocene history of the species is well documented in Europe (Hemmer, 1974; Kurt´en and Poulianos, 1977, 1981; Schütt and Hemmer, 1978; Bishop, 1982; Sala, 1990; Argant, 1991; Garcia, 2003; Testu, 2006; Argant et al., 2007; Barycka, 2008; 18

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

(Głazek et al., 1976). It is assigned to interstadial warming (Małopola­ nian interglacial sensu Ber, 2005). Paleomagnetic investigation of the sediment revealed positive magnetic polarity referred to Brunhes epoch of normal polarity (Głazek et al., 1977; Lindner, 1982). The composition of the rodent fauna documents the uppermost Biharian of the Tem­ plomhegy phase (Nadachowski, 1990). Mimomys savini, the index spe­ cies for the upper Biharian in Central Europe, is abundant and shows some advanced morphological features similar to Arvicola Lac´ep` ede, 1799, believed to be its descendant (Nadachowski, 1985; Fostowicz-­ Frelik, 2008). The fauna is dominated by the Early and earliest Middle Pleistocene faunal elements such as Pliomys episcopalis M´ehely, 1914, P. coronensis, Allocricetus bursae Schaub, 1930, and A. ehiki Schaub, 1930, Glis sackdillingensis (Heller, 1953), Canis mosbachensis Soergel, 1925; Vulpes praeglacialis; early form of Ursus deningeri von Reichenau, 1904; Gulo schlosseri Kormos, 1914; Martes vetus Kretzoi, 1942; Mustela stromeri Kormos, 1934; Mustela strandi Kormos, 1934; Mustela palerminea; Mus­ tela praenivalis Kormos, 1934; Praemegaceros verticornis; Capreolus sues­ senbornensis Kahlke, 1956; Cervalces latifrons (Wiszniowska, 1989; Wolsan, 1989a, b; Nadachowski, 1990; Marciszak et al., 2011, Wagner ˇ ´k, 2012; Stefaniak, 2015). On the other hand, the lack of and Cerm a earlier faunal elements like Allophaiomys Kormos, 1933, also confirms the late dating. Additionally, the occurrence of forms like Microtus arvalidens Kretzoi, 1958, M. gregaloides and Clethrionomys glareolus (Schreber, 1780), and morphology of some forms of large mammals, like U. deningeri, M. vetus, M. stromeri, M. palerminea, M. praenivalis, P. ver­ ticornis, C. suessenbornensis or C. latifrons narrows this period to the early Middle Pleistocene (Nadachowski, 1990; Fostowicz-Frelik, 2008; Mar­ ciszak et al., 2011; Stefaniak, 2015). This earliest Polish lion record fits well into the eastern migration of the P. spelaea, which is also supported by the changes within the evolutionary lineage, which was already discussed by Barycka (2008) and Marciszak et al. (2014, 2019b). However, since the records from Great Britain (Pakefield, West Runton) and Poland (Kozi Grzbiet) are probable contemporary, more migration waves from Africa into Eurasia should be taken into consid­ eration. The first eastern way was via Asia Minor, Caucasus and the European part of Russia, throught the Central Europe and further into the Western Europe. For this direction appeal the finds from Kuznetsk Basin (Sotnikova and Foronova, 2014), Kozi Grzbiet and earlier appearance of lions with progressive dental characteristics in the eastern than in Western Europe. Vast flat plains of Central Europe, full of large ungulates, played an important role as a corridor for migrating animals which moved from east to west and in the opposite direction. This area also played a major role in north-south migrations, especially during glacials and stadials, where more thermophilous species were retreated to the south. Such migrations were already documented for Silesia, a part of southern Poland (Marciszak et al., 2016, 2019a, 2020). The route they took to the territory of the present-day Czech and Slovak Republics led through some gaps in the Carpathian Arch, such as ‘The Moravian Gate’ situated between the Sudeten in the west and the Carpathians in the east. The second migration route is the south-eastern direction, via Bal­ kans and partially also the Apennine Peninsula. This way is even more poorly documented, since the earliest findings are dated on MIS 19–16 (Sala, 1990; Kahlke et al., 2011). Finally, the last, third, way via the Iberian Peninsula, taking into account the British records, was also possible. Althougth, until now there are no known lion records from the latest Early and earliest Middle Pleistocene, and it cannot be ruled out that the lion was not present in this area at that time. The presence of lion-like felid in the North Africa (Morocco) was documented already in the Early Pleistocene (Geraads, 2008). The distance from Africa to Europe in the nearest place is recently ca. 14 km, and such a distance for extant large cats is not a problem (Hemmer and Heidtke, 2013). During the cold periods, the lovered sea level caused, therefore the distance could have been even shorter. Additionally, in the case of many species, especially carnivores, the role of natural barriers should not be overestimated. Lion is regarded as a

species avoiding water, but it is also very flexible animal and tend to adapt to local, sometimes very specific, habitat conditions. A good example are populations from Okavango Delta, where there are prides adapted to live and hunt in water (Kotze et al., 2018). In this context, the Gibraltar Strait can be regarded as a conceivable gate from Africa to Eurasia, and one of the waves which initiated the Pleistocene lion speciation. A similar scenario was already discussed for the Pleistocene jaguar Panthera gombaszoegensis (Hemmer and Heidtke, 2013). 6.2. The relationships of P. spelaea with speleoid bears P. spelaea is commonly represented in Polish sites, and among them one of the most extensive collection from Nied´zwiedzia Cave with more than 350 bones belonging to 10 individuals including 9 males and 1 female. A similar scenario with the predominance of large, massive males also in other Sudeten caves may suggest that, among healthy specimens, they were regarded as an especially predicted to such hunts. Anyonge (1996) noted that the long bones of Panthera atrox were much thicker than those of similar-sized modern Panthera leo. Being very similar, robustly build and large as P. atrox, European Pleistocene lions in general (Hemmer, 1974, 2011; Argant, 1991, 2000; Sabol, 2011b; Marciszak et al., 2014) and Polish lions in particular, also had extremely powerful forelimbs. It might had given a great advantage when hunting a cave bear in narrow and dark cave corridors. Speleoid bears were atypical prey for the Pleistocene lion, but when the food was scarce, lions entered deeply into caves and hunted bears (Argant, 1988, 1991, 2000; Diedrich, 2013a-b, 2012, 2014, 2017; Marciszak et al., 2014, 2019a, b, 2020). It was risky business facing this powerful animal in total darkness, but their strong forelimbs might have helped subdue prey due to a lethal bite. During such fights, lions also lost their battles, especially with large bears, but their carcasses were mostly not scav­ enged by herbivorous cave bear and left inside the caves. This behaviour might be interpreted as the main reason for the accumulation of lion bones, which was not typical cave dweller as well (Diedrich, 2011a; b, 2012, 2014, 2017). Evidences of such activity were found on the numerous bear bones belonged mostly to cubs and young specimens, less than one-year-old. But also many bones of adult animals hold bite marks, chewing and scratches, clear proof of predators’ activity. One of the most spectacular examples is a neurocranium of the young female of Ursus ingressus, which holds a few holes, with two main on the frontal and right parietal bones (Nowakowski and Stefaniak, 2015). The blurred edges of the external lamellae and sclerotic lines confirmed that the cave bear female had survived the blows, however because of illness caused by them it died in the next few months. They were interpreted as remains of a powerful bite from the canines of another carnivore (Nowakowski and Stefaniak, 2015). Possible explanation is an attack of P. s. spelaea, since the hunting specialisation by this species on bears is well known due to nitrogen isotope signs (Bocherens et al., 2011a, b; Bocherens, 2015). The skull was found in the trench K III in Lion’s Hall of Nied´zwiedzia Cave, where also a skull of P. s. spelaea of considerable size (43 cm) was found (Wiszniowska, 1978; Barycka, 2008; Marciszak et al., 2014). The dis­ tance between two main holes (ca. 85 mm) fits well with the canines spacing in the lion specimen. Additionally, the cranium has scraches, sharp-edged longitudinal scars indicating the use of sharp-edged tool which was interpreted as an effect of skinning and possible indirect proof of the human’s existence (Nowakowski and Stefaniak, 2015). However, in our opinion, another explanation is possible. Recently, when the Siberian tiger Panthera tigris tigris (Linnaeus, 1758) hunts on brown bear, it jumps on a bear from above, and tried to hold and overpowered its head in frontal paws, and simultaneously biting through its cervical vertebrae (Ognev, 1935; Maz´ ak, 1979; 1981; Kir­ illova and Tesakov, 2008; Kirillova et al., 2009). During such fights, tiger claws usually left sharp-edged longitudinal scars on bear’s head, the same as observed on the skull of the cave bear female from the Nied´zwiedzia Cave. There will be no doubt that it will be correct to 19

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

extrapolate such a hunting behaviour on P. s. spelaea (Kirillova et al., 2009; Diedrich, 2017). Another example of such activity is known from the layer III of Deszczowa Cave, where gnawing mark was found on the cave bear radius (Wojtal, 2007: 39).

existed in low density. Based on palaeontological data, it might be said that these areas were split in the entire territory of Europe (e.g. partially British Isles, vicinity of the North Sea or the Carpathian Arch). All these areas were more or less restricted to small habitats, and maybe even called “refugees” for the scimitar cat. And although being extremely rare in the late Middle and Late Pleistocene, H. latidens survived as long as to the Late Pleistocene (Reumer et al., 2003), and finally disappeared somewhere in the latest part of the last glacial (probably for the same reasons as the cave lion does). The direct and indirect negative role in the vanishing process and final disappearance of P. gombaszoegensis and H. latidens is well documented and was discussed by many previous ´n et al., 2005; García authors (Hemmer, 2001, 2003, 2004, 2011; Anto ´s, 2007; Bona and Sardella, 2014; Marciszak et al., 2014). and Virgo

6.3. The role of P. spelaea in European palaeguilds The appearance of a huge newcomer, P. s. fossilis, into Eurasia, together with the cave hyena C. c. spelaea and probably also leopard Panthera pardus Linnaeus, 1758, was a dramatic event and turnover of the carnivore paleoguild. Dominated so far carnivores, like Homotherium crenatidens, Lycaon lycaonoides (Kretzoi, 1938), Pachycrocuta brevirostris, Megantereon cultridens , Acinonyx pardinensis, and P. gombaszoegensis, faced to a new inteligent, social (apart the leopard) and ecologically flexible species. Until newcomers were quite rare, and restricted to some areas, the impact was not so high (Hemmer, 2001, 2003, 2004; García ´s, 2007). However, just after lion and hyena widespread into and Virgo Europe, their impact was much stronger. Some of older forms like M. cultridens, A. pardinensis and P. brevirostris vanished very early or became extremely rare. Other survived, but their density was low, and were probably pushed into a less comfortable habitats. Only thanks to the extreme ecological adaptability, P. gombaszoegensis was one of the longest survived species from the ancient paleoguild, and its last Eurasian records are dated on MIS 10–9 (Baryshnikov, 2011; Marciszak et al., 2014). The fossil material from many early and Middle Pleistocene Euro­ pean localities indicate enormous large and robust lion, which presence within the carnivore guild would have been an especially valuable for the cats’ palaeohierarchy (Hemmer, 2001, 2003, 2004). Its enormous size (body mass 350–500 kg), and probably social life guaranteed him a leading position among other carnivores. By virtue of its impressive posture, it is likely that the early lion was able to displace any European predator from its rightful kill. Even the scimitar cat, despite its fearsome sabres, does not stand a chance to defend its kill against the cow-sized rival (Hemmer, 2004). It seems, however, that different species might suffered less or more, which mostly depend on their ecological adapt­ ability. When analysed a large amount of Middle Pleistocene sites of various age, it states clear what carnivores were able to survive in a new situation. The first vanishing species from was A. pardinensis, which disappeared in the mid Middle Pleistocene (Hemmer, 2001, 2003, 2004; ´s, 2007). Among other large carnivores, giant cheetah García and Virgo was probably the most delicate species vulnerable for the competition, which simultaneously held relatively (in comparison to its size) low position in paleohierarchy. L. lycanoides and P. gombaszoegensis survived longer, probably even up to the late Middle Pleistocene, until MIS 10–9 ´nossy, 1990; Marciszak et al., 2014). (Ja Relationships between the Pleistocene lion and the scimitar cat where more complicated, since Homotherium still existed in the Late Pleistocene. Members of the genus Homotherium are among the most successful carnivore species and are present in the fossil material since the late Pliocene (Diedrich and McFarlane, 2017). In Europe, species was widely distributed and regarded as dominant carnivore species through the Early Pleistocene (Hemmer, 2001, 2003, 2004). But in the latest part of the Early and in the early Middle Pleistocene, Homotherium decreasing in size and in massiveness, which caused on average twice lesser weight in Middle Pleistocene individuals that those from the Early Pleistocene. It is not clear what factor was mainly responsible for that (most probably it was a multiple situation), because these lesser, gracile homotheres appeared in the same time in many different European re­ gions. We could not resolve this problem, but locally lion pressure might play a significant role (even if the negative influence of other factors cannot be entirely rule out). It is virtually unknown how hard the competition level was, but it might be cautiously assume that both species of equal size, lion and homotherium, compete, and Homotherium found a way to reduce a competitive pressure. After long time of co-existence, it found favorable habitats where lion was absent or

6.4. Extinction of P. spelaea in Poland and Europe Finally, the basic question is about the reasons of the extinction of the Pleistocene lion in Poland. Their respective disappearance is treated as an extinction, in view of their taxonomical independence, and part of a major global episode of megafaunal extinction that occurred within the late part of the Last Glacial (Martin and Steadman, 1999; Barnosky et al., 2004; Koch and Barnosky, 2006; Stuart and Lister, 2011). In Poland, this species vanished much earlier than in other European areas, and the reasons of such situation can be diverse. The youngest radio­ carbon date from Poland came from Raj Cave and the specimen is dated just before the Last Glacial Maximum (LGM) sensu Marks et al. (2016). What interesting, similar dated specimens are known so far only from Kostenki (Russia) (Stuart and Lister, 2011). Firstly, despite the species is recently known from 60 sites, only the limited number of dates was done and it is possible that later survivals may yet be discovered in some areas. This explanation seems to be recently the most reliable. Climatic changes might play the main role, even if the species showed a great ecological flexibility, and this predominantly open-habitat felid may also have occurred in more open woodland (Stuart and Lister, 2011; Marciszak et al., 2020). Such situation in the Late Pleistocene of Poland was already discussed for Sudeten caves for the cave hyena and wolf. Such massive animal, with relatively short legs and narrow feets might have found a substantial disadvantage to move and hunt in a thick snow cover. For the snowing periods, this short-legged animal was probably not capable of moving efficiently and fastly. Almost the same in linear dimensions, but ca. 30% lighter (on average 60–70 kg) C. l. spelaeus with longer legs and broader feet would have an advantage. Therefore, it was suggested the main cursorial predator in the Sudeten area (Marciszak et al., 2020). Since C. c. spelaea vanished from Poland even before the cave lion, C. l. spelaeus and Ursus arctos priscus should be regarded as main competitors of P. s. spelaea in this area. Analysis of isotope values documented that wolf seems to have outcompeted cave lion from access to other prey species (Bocherens et al., 2011a; Bocherens, 2015). Wolf as smaller, but agile, highly mobile and ecologically extremely adapted species, could roam much broad areas and run over much larger dis­ tances in search of food. Such an indirect pressure might cause that herds or large ungulates, like rendeers, horses and bovids, were fright­ ened and chased away by packs of hunting wolves. It might caused that the potential prey was virtually absent, or very limited in number. Lion, despite their great mobility, was also a territorial felid, and seems to be at a disadvantage in this respect. When co-existing with collective and gregarious wolf, cave lion seems to have limited their prey choice to those not taken by these other predators, essentially reindeer and cave bears (Bocherens et al., 2011a; Bocherens, 2015). As in the case of hyena-wolf relationships in Sudetenland, climatic condition like deep snow cover also played an important role (Marciszak et al., 2020). Beside the indirect competition, some level of direct competition between wolf and lion might also exist, although its level is hard to reconstruct. Cave lion was a solitary hunter or operated in very small prides (Hemmer, 2011; Sabol et al., 2018). Recently, wolves tend to avoid direct competition with the Siberian tiger, which high density may 20

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

localy lowered wolves’ number to make them a functionally insignifi­ cant component of the ecosystem (Miquelle et al., 2005). Obervations on interactions between the two species indicate that tigers occasionally chase wolves from their kills, while wolves will scavenge from tiger kills. There are records of tigers killing wolves without consuming them (Heptner and Naumov, 1967; Miquelle et al., 2005). However, in the second part of the Late Pleistocene large areas were overhelmed by packs of large, robust wolves which totally dominated in the environ­ ment. It cannot be excluded that in great number they will be able to attack and even kill a solitary lion. As an example of wolf dominance are the Carpathians and Sudety Mts, together with Bohemian, Moravian and Silesian lowlands adjacent to them. Cave hyena was absent in Sudety Mts, and the cave lion and wolf were very frequent. But between 35 and 30 ka lion disappeared, and the wolf became the sole, dominant, active carnivore, together with the brown bear. The relationships between P. s. spelaea and U. a. priscus in the late Middle and Late Pleistocene are hardly to reconstruct, in view of scarce direct evidences. As was discussed above, the Pleistocene lion attacked and killed cave bears, and smaller brown bears also falled victim of lion attacks. However, beside U. a. arctos, metrically and morphologically similar to extant Central European bears in the Middle and Late Pleis­ tocene immense steppe brown bear also existed, which on average was as larger or larger than even the largest extant coastal brown bears (Thenius, 1956; Musil, 1964; Marciszak et al., 2016, 2017, 2019a, b, c; Ersmark et al., 2019). It is known from the isotope analysis, that the pre-LGM brown bears were highly carnivorous and their diet strongly overlap with other carnivores like cave lion (Bocherens et al., 2011a, b, 2015; Bocherens, 2015; Ersmark et al., 2019). The steppe brown bear became so large, and being formidable kleptoparasite rather than active hunter, went into the direct confrontation with the lion and other car­ nivores (Marciszak et al., 2016, 2019a, b; Musil, 2018). Recently the Siberian tiger regularly kills brown bear, and the latter can constitute up ´k, 1981; Sabol et al., 2018). However, almost all to 8% of its diet (Maza these bears are the specimens under 200 kg (sometimes rearely exceeding this value), cubs, young individuals and females. Hunting on the brown bear is a risky buissnes for the tiger, as was shown by sum­ marising studies by Seryodkin et al. (2018: 859–860): “The brown bear is capable of not only showing resistance to the tiger that attacked it, but also of emerging victorious out of the fight. Of the 45 cases of encounters of tigers with brown bears (…), the tiger was the initiator in 13 cases, the bear started eight fights, and in the other cases, the attacker was not established. In 51.1% of cases, the fights ended with the death of the bear, in 26.7% with the death of the tiger, and in 22.2% the animals broke up”. The adult male brown bear class appears instead virtually invulnerable from tiger predation, as pretty much straightforward when considering size, mass, power and strength of healthy enough in­ dividuals and their very tough and aggressive behaviour. Actually the opposite situation of adult male brown bear potentially posing a threat to the Siberian tiger, even adult males, seems prevalent as assessed from kill disputes which appear invariably totally dominated by the large adult male brown bears weighing in excess of 300 kg (Miquelle et al., 2005; Tkachenko, 2012). Large brown bears are known regularly tracking tigers and usurping their kills or even killed them during aggressive encouters (Sysoev, 1960, 1966; Rakov, 1965, 1970; Gor­ okhov, 1973; Kostoglod, 1977, 1981; Tkachenko, 2012; Seryodkin, 2016; Seryodkin et al., 2018). Bear in such size class can even remain virtually completely unchallenged if already tested by any tiger in a previous experience ended up without drastic consequences for the tiger able to break the fight and run away (Warsaw, 2018). Largest males of the Polish specimens of P. s. spelaea reached up to 400 kg, which is double of the weight of the average Siberian tiger. However, U. a. priscus was truly gigantic bear with an average weight of adult males ca. 600–800 kg, and it can reach even up to 1500 kg in the largest in­ dividuals (Musil, 2018; Marciszak et al., 2019c). Such immense in­ dividuals are known among other from Nied´zwiedzia Cave, where they co-occurred with the cave lion. In direct confrontation with the

horse-size bear, even the largest cat, hyena clan or wolf pack was without any chance and usually left their rightfull kill for the competitor (Bocherens et al., 2011a, b, 2015; Bocherens, 2015) (Fig. 18). There is also no doubt that P. spelaea encountered and came into a conflict with human, and so far known direct evidence for such behav­ iour from the Polish territory are cut marks on lion metatarsal bone from layer 12 of Nietoperzowa Cave (Wojtal, 2007; Nadachowski et al., 2015). Summarising, extinction of the Pleistocene lion is broadely correlated with the general collapse of the “mammoth steppe” ecosystem, and was resulted of multiple reasons, like climatic changes, re-building of herbivore guilds, competition and human pressure (Stuart and Lister, 2011). There is no doubt on the direct human impact on the Holocene extinction of the lion in Eurasia, e.g. Masseti and Mazza (2013). But the level of human pressure on last populations of the cave lion in Central Europe, as in Poland, needs further verification. More work concentrating on the timing and extent of human population changes as well as dynamic of key carnivore species like cave hyena, brown bear and wolf in Poland is needed. It is especially important in order to test if there is any correlation with P. spelaea extinction, and if so, how this process runs. 7. Conclusions A detailed review of the Polish P. spelaea material documented the occurrence of this species from 60 localities including 18 open-air and 42 cave sites dated in the range of 750–28 ka. Most of the records came from southern Poland (Silesia) and adjacent areas, with few located more northward. Among them, the oldest record from Kozi Grzbiet, mentioned for the first time in this paper, is also with the finds from British localities, among the oldest European occurences of the species. Most of the dates are concentrated on the second half of MIS 3 and showed that the species disappeared more or less synchronously across Poland. However, given the limitations of the data, it is possible that later survivals may yet be discovered in some areas. Records from openair sites are mostly accidental discoveries during field works, while cave finds are mostly connected with archaeological research in caves and exploitation of their sediments for fertilizers. Only few juvenile speci­ mens were found, and the rest material is overhelmed by adult, but not senile individuals, and the strong dominance of males. Their predomi­ nance can be partially explained by their specialisation in hunting on cave bears during the times when the other, more typical food sources like ungulates were scarce. Spelaeoid bears were atypical prey for the Pleistocene lion, but when food was scarce, lions entered deeply into caves and hunted them. Many lions were killed during such incidents, but not eaten by herbivorous bears. Another explanation is that larger and stronger male lions more often tend to hunt very big preys and went into the conflicts with other carnivores than females do, which resulted in higher injuries and mortality. In Poland three chronosubspecies of the Pleistocene lion were distinguished, where P. s. fossilis was recorded from 7 sites, dated between 750 and 240 ka, P. s. intermedia only ´rna Cave and P. s. spelaea, typial faunal partially from Wierzchowska Go element of the Late Pleistocene paleoguilds, from 48 localities. Except of four sites, Bi´snik, Naciekowa, Nied´zwiedzia and Wierz­ ´rna Caves, the Polish finds of P. spelaea throughout its chowska Go Pleistocene history are generally low in abundance at sites where they occur. Furthermore, fossil records of the species are in the entire Polish territory, suggesting the country as a fragment of the Eurasian meta­ population. In this situation, relatively subtle changes in the animals’ habitat could have impacted its survival and extirpated local pop­ ulations. The preferred habitat for P. spelaea was a productive envi­ ronment – tundra steppe. It is therefore plausible that the extirpation of Polish populations was driven by the loss of suitable habitat. The pos­ sibility of changes in adaptation of P. spelaea should also be considered. The species had survived durable glacials and interglacials, so why not the Holocene? However, a particular loss of mitochondrial lineages taken place between 48 and 45 ka, created a bottleneck phenomenon. 21

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Fig. 18. Two scenes illustrating the complexity of relationships between Panthera spelaea spelaea and different bears in Nied´zwiedzia cave. At the top hunting scene where lion using powerful forelimbs subdue cave bear Ursus ingressus before delivering the lethal bite. Below lion loses battle for carcass with immense steppe brown bear Ursus arctos priscus. Drawn by W. Gornig.

Dwarf, gracile specimens are noted since that time in the European population of P. spelaea, also in the Polish material. Whatever the precise adaptive significance of these changes, they may point to a shift in the adaptive niche of the species during its LGM refugial phase, or possibly the spread of a differently adapted population. This might explain the restricted Late Pleistocene range and also the species’ heightened intolerance of some environments. However, there are also coincidences in timing between terminal dates of P. spelaea and the spread of human populations. People were present in Europe at the time of extirpation of the species. However, in Poland the archaeological evidence suggests low human density with limited environmental impact during the time when the Pleistrocene lion vanished. Landscape alterations were not a significant factor in threatening populations for the last European Pleistocene lions, while

hunting remains a possible, but unproven, factor. Body-size reduction in last and terminal populations has been suggested as a marker for envi­ ronmental deterioration driving extinction in the Late Pleistocene, but its absence does not necessarily implicate hunting and impact by default. The apparent maintenance of normal body size in the late P. spelaea may be due to countervailing selective forces. Summarising all the data presented above, it seems clear that environmental factors, cumulatively over thousands of years, reduced the Pleistocene lion populations to a highly vulnerable state. In this situation, even relatively low-level hunting by small human populations could have contributed to its extinction. In worsening habitat conditions, when Pleistocene lion reproductive rates dropped, even low levels of competition pressure may have been unsustainable. It is alternatively plausible that in this situa­ tion, with the species reduced to small vulnerable populations, its 22

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

extinction was inevitable even without human impact, analogous to the extinction debt posited for many threatened species today. Unstable climatic conditions, low density of favorable prey spectrum and growing competition with wolf may be decisive for the extinction of lion in Poland. The post-LGM records of the lion are unknown from this country, and it is one of the earliest vanished carnviores, pre-dated only by a cave hyena.

References ´ Abelov´ a, M., 2005. Doprovodn´ a fauna vel’kých cicavcov medvedej jaskyne “Za H´ ajnovnou”, Javoˇriˇcsky kras. Pˇrirodovˇedne˙ studie muzea Prostˇejovska 8, 171–184. Adam, K.D., 1975. Die mittelpleistoz¨ ane S¨ augetier-Fauna aus dem Heppenloch bei Gutenberg (Württemberg). Stuttgarter Beitr¨ age zur Naturkunde. Serie B Geologie und Pal¨ aontologie 3, 1–247. Aleekseva, L.I., 1990. Fauna Pozdnepleystotsenovykh Mlekopitayushchikh Vostochnoy Yevropy [Late Pleistocene Mammal Fauna of Eastern Europe]. Nauka Press, Moscow (in Russian). Altuna, J., 1972. Fauna de Mamíferos de los yacimientos prehist´ oricos de Guipúzcoa. Munibe 24, 1–464. Altuna, J., 1981. Fund eines Skeletts des H¨ ohlenl¨ owen (Panthera spelaea Goldfuss) in Arrikrutz, Baskenland. Bonn. Zool. Beitrage 32, 31–46. Anelli, F., 1954. Contibutio alla conoscenza della fauna diluviale della caverna Pocala di Aurisina (Trieste). Memoires Carta Geologie Italiane 11, 1–57. Ant´ on, M., Galobart, A., Turner, A., 2005. Co-existence of scimitar-toothed cats, lions and hominins in the European Pleistocene. Implications of the post-cranial anatomy of Homotherium latidens (Owen) for comparative palaeoecology. Quat. Sci. Rev. 24, 1287–1301. https://doi.org/10.1016/j.quascirev.2004.09.008. Anyonge, W., 1996. Locomotor behavior in Plio-Pleistocene sabre-tooth cats: a biomechanical analysis. J. Zool. 238, 395–413. https://doi.org/10.1111/j.14697998.1996.tb05402.x. Arambourg, C., 1960. Les faunes mammalogiques du Pl´eistoc`ene circumm´ editerran´ een. Quaternaria 6, 97–109. Argant, A., 1988. Etude de l’exemplaire de Panthera spelaea (Goldfuss, 1810) (Mammalia, Carnivora, Felidae) du gisement Pl´ eistoc` ene moyen recent de la grotte d’Az´ e (Saˆ oneet-Loire). Rev. Pal´eobiol. 7, 449–466. Argant, A., 1991. Carnivores quaternaires de Bourgogne. Doc. Lab. Geol. Facul. Sci. Lyon 115, 1–301. Argant, A., 2000. Les sites pal´ eontologiques du Pleistoc`ene moyen en Mˆ aconnais. Bull. Soc. Prehist. Fr. 97, 609–623. Argant, A., 2010. Carnivores (Canidae, Felidae et Ursidae) de Romain-la-Roche (Doubs, France). Rev. Pal´eobiol. 29, 495–601 doi: halshs-00586673. Argant, A., Argant, J., Jeannet, M., Erbajeva, M., 2007. The big cats of the fossil site Chˆ ateau Breccia Northern Section (Saˆ one-et-Loire, Burgundy, France): stratigraphy, palaeoenvironment, ethology and biochronological dating. Cour. Forschungsinst. Senckenberg 259, 121–140 doi: hal-01778652f. Argant, A., Brugal, J.-P., 2017. The cave lion Panthera (Leo) spelaea and its evolution: Panthera spelaea intermedia nov. subspecies. Acta Zool. Cracov. 60 (2), 59–104. https://doi.org/10.3409/azc.60_2.59. Astruc, J.-G., Legenre, S., Marandat, B., Montuirem, S., Sudre, J., Sige, B., 1994. Red´ecouverte du Lion de Cajarc, Panthera spelaea (Goldfuss, 1810), d’ˆ age pl´eistoc` ene sup´erieur, sur le Causse m´eridional du Quercy, en context de karst polyphas´e. Bulletin Societe Histoire Naturelle de Toulouse 130, 79–83. ´ Ballesio, R., 1975. Etude de Panthera (Leo) spelaea (Goldfuss) nov. subsp. (Mammalia, Carnivora, Felidae) du gisement Pl´eistoc` ene moyen des Abimes de la Fage ` a Noailles (Corr`eze). Nouvelles Archives du Mus´ eum d’Histoire Naturelle de Lyon 13, 47–55. Ballesio, R., 1980. Le gisement pl´ eistoc` ene sup´erieur de la grotte de Jaurens ´ a Nespouls, Corr`eze, France: les carnivores (Mammalia, Carnivora). II Felidae. Nouvelles Archives du Mus´ee d’Histoire naturelle Lyon 18, 61–102. Barnett, R., Shapiro, B., Barnes, I., Ho, S.Y., Burger, J., Yamaguchi, N., Higham, T.F., Wheeler, H.T., Rosendahl, W., Sher, A.V., Sotnikova, M., Kuznetsova, T., Baryshnikov, G.F., Martin, L.D., Harington, C.R., Burns, J.A., Cooper, A., 2009. Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity. Mol. Ecol. 18, 1668–1677. https://doi. org/10.1111/j.1365-294X.2009.04134.x. Barnett, R., Zepeda Mendoza, M.L., Rodrigues Soares, A.E., Ho, S.Y.W., Zazula, G., Yamaguchi, N., Shapiro, B., Kirillova, I.V., Larson, G., Gilbert, M.T.P., 2016. Mitogenomics of the extinct cave lion, Panthera spelaea (Goldfuss, 1810), resolve its position within the Panthera cats. Open Quat. 2, 1–4. https://doi.org/10.5334/ oq.24. Barnosky, A.D., Koch, P.L., Feranec, R.S., Wing, S.L., Shabel, A.B., 2004. Assessing the causes of Late Pleistocene extinctions on the continents. Science 306, 70–75. https:// doi.org/10.1126/science.1101476. Barycka, E., 2008. Middle and late Pleistocene felidae and hyaenidae of Poland. Fauna Pol. 2, 1–228. Baryshnikov, G.F., 2011. Pleystotsenovyye felidae (mammalia, carnivora) iz paleoliticheskikh peshchernykh stoyanok kudaro na kavkaze [Pleistocene felidae (mammalia, carnivora) from the kudaro paleolithic cave sites in the Caucasus]. Proceedings of the Zoological Institute Russian Academy of Science 315 (3), 197–226 (in Russian). doi: 569.742.7:551.791(479.2). Baryshnikov, G.F., 2016. Ostanki pozdnepleystotsenovykh felidae (mammalia, carnivora) iz peshchery geograficheskogo obshchestva na dal’nem vostoke rossii [late Pleistocene felidae remains (mammalia, carnivora) from geographical society cave in the Russian far east]. Proceedings of the Zoological Institute Russian Academy of Science 320 (1), 84–120 (in Russian). doi: 569.742.7 (571.63). Baryshnikov, G., Boeskorov, G., 2001. The Pleistocene cave lion, Panthera spelaea (carnivora, felidae) from yakutia, Russia. CRANIUM 18, l–18. Baryshnikov, G.F., Petrova, E.A., 2008. Cave lion (Panthera spelaea) from the Pleistocene of chuvashiya, European Russia. Russ. J. Theriol. 7, 33–40. Baryshnikov, G.F., Tsoukala, E., 2010. New analysis of the Pleistocene carnivores from petralona cave (Macedonia, Greece) based on the collection of the thessaloniki aristotle university. Geobios 43, 389–402. https://doi.org/10.1016/j. geobios.2010.01.003.

Availability of the data All information, which provide readers where they can obtain the research data required to reproduce the work reported in the manuscript like status of the material, its location, collection numbers etc are pro­ vided within the manuscript and both appendixes. All material, if currently present, is available to study in particular museums and pri­ vate collections. Authors’ contributions All authors have made substantial contributions to this submission. The individual contributions of each co-author is as follow: AM: intro­ duction, morphological and metrical studies, description of bones and partially of sites, morphological and metrical results and partially in discussion, Figs. 2–18, Tables 1 and 2, and Appendixes; GL: partially description of sites and materials, and partially in metrical and morphological results and discussion, Tables 1 and 2, and Appendixes; KP: partially description of sites and materials, and partially metrical and morphological results and discussion, Tables 1 and 2, and Appen­ dixes; GJ: partially description of sites and materials, and partially dis­ cussion; UR-S: Fig. 1, and partially metrical and morphological results and discussion; KZ-S: partially metrical and morphological results and discussion; AN: partially metrical and morphological results and discussion. Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Acknowledgements We are very grateful to O. Kovalchuk for linguistic improvements of the manuscript, and two anonymous reviewers for their helpful critical comments and suggestions. We are very grateful to Mrs Robert Szybiak, Tatiana Woroncowa-Marcinowska, Waldemar Wiązek and Jerzy Dąb­ rowski for granting us access to the material and Prof. Piotr Wojtal for sharing unpublished AMS date. This work was supported by the Ministry ´ of Science and Higher Education of Poland (project 1076/S/IBS/2017)

and internal grant for young scientists by the Faculty of Biological Sci­ ´ ences (project no. 2224/M/IBS/17). Our research was also supported by (1) the grant of KBN (State Committee for Scientific Research) project 303 078 32/2589 awarded to A. Nadachowski; (2) the grant of Ministry of Sciences and Higher Education of Poland, by the Institute of Sys­ tematics and Evolution of Animals, Polish Academy of Sciences, project No. 2 P04C 081 30 awarded to P. Wojtal; (3) the grant from the Ministry of Science and Higher Education of Poland (N N307 050139), project “Biochronology of the Pleistocene lion Panthera spelaea (Goldfuss, 1810) from Poland in overall view of the lineage evolution” awarded to G. Lipecki; (4) grant from the Ministry of Science and Higher Education (N N307 050139) awarded to K. Pawłowska. Appendix A. Supplementary data Supplementary data to this article can be found online at https://doi. org/10.1016/j.quaint.2020.12.018.

23

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Ber, A., 2005. Polish Pleistocene stratigraphy – a review of interglacial stratotypes. Neth. J. Geosci. 84, 61–76. https://doi.org/10.1017/S0016774600022964. Bishop, M.J., 1982. The mammal fauna of the early Middle Pleistocene cavern infill site of Westbury-sub-Mendip. Somerset. Special Papers in Palaeontology 28, 1–108. Bocherens, H., 2015. Isotopic tracking of large carnivore palaeoecology in the mammoth steppe. Quat. Sci. Rev. 117, 42e71. https://doi.org/10.1016/j. quascirev.2015.03.018, 2015. Bocherens, H., Drucker, D.G., Bonjean, D., Bridault, A., Conard, N.J., Cupillard, C., Germonpr´e, M., H¨ oneisen, M., Münzel, S.C., Napierrala, H., Patou-Mathis, M., Stephan, E., Uerpmann, H.-P., Ziegler, R., 2011a. Isotopic evidence for dietary ecology of cave lion (Panthera spelaea) in North-western Europe: prey choice, competition and implications for extinction. Quat. Int. 245, 249–261. https://doi. org/10.1016/j.quaint.2015.09.091. Bocherens, H., Stiller, M., Hobson, K.A., Pacher, M., Rabeder, G., Burns, J.A., Tütken, T., Hofreiter, M., 2011b. Niche partitioning between two sympatric genetically distinct cave bears (Ursus spelaeus and Ursus ingressus) and brown bear (Ursus arctos) from Austria: isotopic evidence from fossil bones. Quat. Int. 245, 238–248. https://doi. org/10.1016/j.quaint.2010.12.020. Bocherens, H., Drucker, D.G., Germonpr´e, M., L´ azniˇckov´ a-Galetov´ a, M., Naito, Y., Wissing, C., Brůˇzek, J., Oliva, M., 2015. Reconstruction of the Gravettian foodweb at Pˇredmostí I using isotopic tracking of bone collagen. Quat. Int. 359–360, 211–268. https://doi.org/10.1017/qua.2018.40. Bona, F., 2006. Systematic position of a complete lion-like cat skull from the Eemian ossiferous rubble near Zandobbio (Bergamo, north Italy). Rivista Italiana e Stratigrafia 112, 157–166. https://doi.org/10.13130/2039-4942/5855. Bona, F., Sardella, R., 2012. The middle Pleistocene large felids (mammalia) from brecce di Soave (verona, N-E Italy). Riv. Ital. Paleontol. Stratigr. 118 (1), 193–199. https:// doi.org/10.13130/2039-4942/5999. Bona, F., Sardella, R., 2014. Co-ccurrence of a sabertoothed cat (homotherium sp.) with a large lion-like cat (Panthera sp.) in the middle Pleistocene karst infill from nuova cava zanola (paitone, brescia, lombardy, northern Italy). Boll. Soc. Paleontol. Ital. 53 (2), 81–88. https://doi.org/10.4435/BSPI.2014.08. Boule, M., 1906a. Les grand chats des caverns. Ann. Palaontol. 1, 69–95. Boule, M., 1906b. Les grottes de Grimaldi (Baouss´ e-Rouss´ e). Geologie et Pal´eontologie 1, 70–362. Bourguignat, J.-R., 1879. Description des ossements du grand Felis d´ecouvertes dans une caverne, pr` es Vence. Gustave Retaux, Abbeville. Boeskorov, G.G., Lazarev, P.A., 1997. New finds of late Pleistocene lions, Panthera (leo) spelaea in yakutia. Acta Zool. Cracov. 40, 223–227. Bronk Ramsey, C.H., Higham, T., Bowles, A., Hedges, R., 2004. Improvements to the pretreatment of bone at Oxford. Radiocarbon 4 (1), 155–163. Caloi, L., Palombo, M.R., 1978. Anfibi, rettili e mammiferi di Torre del Pagliaccetto (Torre in Pietra, Roma). Quaternaria 20, 315–328. Casta˜ nos, P., 2005. Estudio paleontol´ ogico de un esque leto de le´ on (Panthera leo) de la Sima de Azoleta (Gorbeia, Alava). Muibe. Antropologia-Arkeologia 57, 123–129. Cervera, J., 1992. Nuevos hallazgos de carnivoros en el complejo de yacimientos mesopleistocenos de la Sierra de Atapuerca, Burgos. Revista Espa˜ nola de Paleontologia Volumne extraordinare 1992, 21–27. Clot, A., 1980. La grotte de la Carriere (Gerde, Hautes-Pyrenees). Stratigraphie et Paleontologie des Carnivores, vol. 3. Travaux Laboratoire Geologie Toulouse, pp. 1–502. Clot, A., Brochet, G., Chaline, J., Desse, G., Evin, J., Granier, J., Mein, P., MourerChauvire, C., Omnes, J., Rage, J.C., 1984. Faune de la grotte pr´ehistorique du bois de Cantet (Esp` eche, Hautes-Pyr´ en´ees, France). Munibe 36, 33–50. Conwentz, H., 1895. XVI. Amtlicher Bericht über die Verwaltung der naturhistorischen, archaeologischen und ethnologischen Sammlungen des Wespreussischen ProvinzialMuseums für das Jahr 1895. Danzig. Cr´egut, E., 1979. La faune de mammif`eres du Pl´ eistoc` ene moyen de la Caune de l’Arago. Universit´e de Provence, Aix-Marseille I. Tautavel (P.-O.). PhD Thesis. ´ 2011. Sur la pr´ Cr´egut-Bonnoure, E., esence du lion des caverns Panthera (Leo) spelaea Goldfuss en provence (sud-est de la France). Quaternaire, Hors-S´ erie 4, 237–244. ´ Boulbes, N., Gu´ Cr´egut-Bonnoure, E., erin, C., Pernaud, J., Tavoso, A., Cammas, R., 2010. Le contexte g´eomorphologique et faunique de l’homme de Montmaurin (HauteGaronne). Pr´ehist. M´ editerran´eennes 1, 33–85. http://journals.openedition. org/pm/543. Crusafont-Pair´ o, M., Casajuana, M.T., 1950. Primer hallazgo del leon de las cavernas, en el Pleistoceno de Cataluna. Pirineos, vols. 17–18. Publicaciones de Instituto Pirenaico de Ecologia et Consejo superior de investogaciones cientificas, pp. 521–543. Cyrek, K., Nadachowski, A., Madeyska, T., Boche´ nski, Z., Tomek, T., Wojtal, P., Miękinia, B., Lipecki, G., Garapich, A., Rzebik-Kowalska, B., Stworzewicz, E., Wolsan, M., Godawa, J., Ko´sci´ ow, R., Fostowicz-Frelik, Ł., Szyndlar, Z., 2000. Excavation in the Deszczowa cave (kroczyckie rocks, częstochowa Upland, Central Poland). Folia Quat. 71, 5–84. Daxner-Hӧck, G., 1975. Caniden-, Feliden- und Musteliden-Funde aus dem Travertin von Weimar-Ehringsdorf. Abh. Des. Zent. Geol. Inst. 23, 485–500. Dawkins, W.B., Sandford, W.A., 1866. The British Pleistocene mammalia. Parts I-II. British Pleistocene felidae. Felis spelaea, Goldfuss. Palaeontogr. Soc. Monogr. 1866, 1–124. Del Campana, D., 1954. Carnivori quaternari della tecchia e della caverna di Equi nelle Alpi Apuane (Mustelidi, Canidi, Felidi). Palaeontogr. Ital. 114, 1–42. Dietrich, W.O., 1968. Fossile L¨ owen im europ¨ aischen und afrikanischen Pleistoz¨ an. Pal¨ aontologische Abhandlungen Abteilung A 3, 323–366. Diedrich, C.G., 2007a. Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeleton remains from Praha-Podbaba and the other lion finds from loess and river terrace

sites in Central Bohemia (Czech Republic). Bull. Geosci. 82 (2), 99–117. https://doi. org/10.3140/bull.geosci.2007.02.99. Diedrich, C., 2007b. The fairy tale about the “cave lions” Panthera leo spelaea (Goldfuss 1810) of Europe – late Ice Age spotted hyenas and Ice Age steppe lions in conflicts – lion killers and scavengers around Prague (Central Bohemia). Scripta Facultatis Scientiarum Naturalium Universitatis Masarykianae Brunensis Geology 35, 107–112. Diedrich, C., 2008. The rediscovered holotypes of the Upper Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) and the steppe lion Panthera leo spelaea (Goldfuss, 1810) and taphonomic discussion to the Zoolithen Cave hyena den at Gailenreuth (Bavaria, South-Germany). Zoological Journal of the Linnaean Society London 154, 822–831. https://doi.org/10.1111/j.1096-3642.2008.00425.x. Diedrich, C., 2009a. Upper Pleistocene Panthera leo spelaea (Goldfuss 1810) remains from the Bilstein Caves (Sauerland Karst) and contribution to the steppe lion taphonomy, palaeobiology and sexual dimorphism. Ann. Paleontol. 95, 117–138. https://doi. org/10.1016/j.annpal.2009.07.001. Diedrich, C., 2009b. Steppe lion remains imported by ice age spotted hyenas into the late Pleistocene perick caves hyena den in northern Germany. Quat. Res. 71 (3), 361–374. https://doi.org/10.1016/j.yqres.2008.12.006. Diedrich, C., 2010. Ein Skelett einer kranken L¨ owin Panthera leo spelaea (Goldfuss 1810) und andere m¨ annliche L¨ owenreste aus Neumark-Nord. Arch¨ aologie in SachsenAnhalt Sonderband 62, 432–441. Diedrich, C., 2011a. The largest European lion Panthera leo spelaea (Goldfuss) population from the Zoolithen Cave, Germany-specialized cave bear predators of Europe. Hist. Biol. 23 (2–3), 271–311. https://doi.org/10.1080/08912963.2010.546529. Diedrich, C., 2011b. Cave bear killers and scavengers from the last Ice Age of Central Europe-feeding specialization as reaction on the mammoth steppe fauna absence in mountainous regions. Quat. Int. 255, 59–78. https://doi.org/10.1016/j. quaint.2011.06.048. Diedrich, C., 2011c. The Late Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeletons from the Sloup and Srbsko caves in Czech Republic (central Europe) and contribution to steppe lion cranial pathologies and postmortally damages as results of interspecies fights, hyena antagonism and cave bear attacks. Bull. Geosci. 86 (4), 817–840. https://doi.org/10.3140/bull.geosci.1263. Diedrich, C., 2011d. Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss, 1810) footprints and bone remains from open air sites in northern Germany-evidence of hyena-lion antagonism in Europe. Quat. Sci. Rev. 30, 1883–1906. https://doi.org/ 10.1016/j.quascirev.2011.03.006. Diedrich, C.G., 2011e. A diseased Panthera leo spelaea (Goldfuss 1810) lioness from a forest elephant graveyard in the Late Pleistocene (Eemian) interglacial lake at Neumark-Nord, central Germany. Hist. Biol. 23 (2), 195–217. https://doi.org/ 10.1080/08912963.2010.507814. Diedrich, C., 2011f. Pleistocene Panthera leo spelaea (Goldfuss, 1810) remains from the Balve Cave (NW Germany) – a cave bear, hyena den and Middle Palaeolithic human cave, and review of the Sauerland Karst lion sites. Quaternaire 22 (2), 105–127. https://doi.org/10.4000/quaternaire.5897. Diedrich, C.G., 2012. Cave bear killers and scavengers from the last ice age of central Europe: feeding specializations in response to the absence of mammoth steppe fauna from mountainous region. Quat. Int. 255, 59–78. https://doi.org/10.1016/j. quaint.2011.06.048. Diedrich, C., 2013. Late Pleistocene Eemian hyena and steppe lion feeding strategies on their largest prey-Palaeoloxodon antiquus Falconer and Cautley 1845 at the straighttusked elephant graveyard and Neanderthal site Neumark-Nord Lake 1, Central Germany. Archaeological and Anthropological Sciences 6 (3), 271–291. https://doi. org/10.1007/s12520-013-0150-7. Diedrich, C., 2014. Holotype skulls, stratigraphy, bone taphonomy and excavation history in the Zoolithen Cave and new theory about Esper’s “great deluge”. Quaternary Science Journal 63 (1), 78–98. https://doi.org/10.3285/eg.63.1.05. Diedrich, C.G., 2017. Hermann’s Cave (Germany) – a Late Pleistocene Cave Bear Den. Bentham Science Publishers, Szard˙za. Diedrich, C.G., McFarlane, D.A., 2017. Homotherium from Middle Pleistocene archaeological and carnivore den sites of Germany - taxonomy, taphonomy and a revision of the Sch¨ oningen, West Runton and other saber-tooth cat sites. Quat. Int. 436, 76–83. Diedrich, C.G., Rathgeber, T., 2012. Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss 1810) skeleton remains of the Upper Rhine Valley (SW Germany) and contributions to their sexual dimorphism, taphonomy and habitus. Hist. Biol. 24, 1–28. https://doi.org/10.1080/08912963.2010.549943. Dubois, A., Stehlin, H.G., 1932. La grotte de Cotencher, station moust´erienne. Memoires de la Soci´ et´e pal´eontologique Suisse 52–53, 1–178. Dufour, R., 1989. Les carnivores pl´eistoc` enes de la caverne de Malarnaud (Ari`ege), (Collection E. Harle, Mus´eum d’Histoire Naturelle de Bordeaux). Universit´e de Bordeaux, Bordeaux. Edinger, T., 1931. Über jungdiluviale S¨ augetiere aus dem Emschergebiet. Pal¨ aontol. Z. 13, 119–133. Erdbrink, D.P.B., 1981. Some more cave lion remains. Zoology, Proceedings C 84 (1), 21–45. Erdbrink, D.P.B., 1983a. Eleven bones: more fossil remains of cave lions and cave hyenas from the North Sea area. Bijdr. Dierkd. 53 (1), 1–12. Erdbrink, D.P.B., 1983b. Still more cave lion remains. Palaeontology, Proceedings B 86 (2), 113–123. Erdbrink, D.P.B., 1986. Lions, leopards and a hyena from deposits along the Meuse and the Waal Netherlands. Proc. Koninklijke Nederl. Akademie Wetenschappen - Ser. B Palaeontol. Geol. Phys. Chem. Anthropol. 89 (1), 1–14. Erdbrink, D.P.B., 1988. Contribution to the record of fossil mammals from the Meuse levels. Proc. Koninklijke Nederl. Akademie Wetenschappen - Ser. B Palaeontol. Geol. Phys. Chem. Anthropol. 91 (4), 321–338.

24

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Ersmark, E., Orlando, L., Sandoval-Castellanos, E., Barnes, I., Barnett, R., Stuart, A., Lister, A., Dal´ en, L., 2015. Population demography and genetic diversity in the Pleistocene cave lion. Open Quat. 1 (1), 1–4. https://doi.org/10.5334/oq.aa. Ersmark, E., Baryshnikov, B., Higham, T., Argant, A., Casta˜ nos, P., D¨ oppes, D., Gasparik, M., Germonpr´ e, M., Lid´ en, K., Lipecki, G., Marciszak, A., Miller, R., Moreno García, M., Pacher, M., Robu, M., Rodriguez Varela, R., Rojo Guerra, M., Sabol, M., Spassov, N., Storå, J., Valdiosera, Ch, Villaluenga, A., Stewart, J.R., Dal´ en, L., 2019. Genetic turnovers and northern survival during the last glacial maximum in European brown bears. Ecology and Evolution 9 (10), 5891–5905. https://doi.org/10.1002/ece3.5172. Fabiani, R., 1919. I mammiferi quaternary della regione Veneta. Mem.gli Ist. Geol. Mineral. Univ. Padova 5, 1–174. Filhol, E., Filhol, H., 1871. Description des ossements de Felis spelaea decouverts dans la caverne de Lherm (Ariege). Annales Science Naturelle, Serie Zoologie 14, 1–120. Fischer, K.-H., 1994. Neufunde von jungpleistoz¨ anen H¨ ohlenl¨ owen Panthera leo spelaea (Goldfuss, 1810) in Rübeland (Harz). Brauschweiger Naturkundliche Schriften 4 (3), 455–471. Foronova, I.V., 2001. Chetvertichnyye mlekopitayushchiye yugo-vostoka Zapadnoy Sibiri (Kuznetskiy basseyn): filogeniya, biostratigrafiya i paleoekologiya [Quaternary mammals of the south-east of western Siberia (Kuznetsk basin): phylogeny, biostratigraphy, and palaeoecology]. Transactions of the Russian Academy of Sciences, Siberian Branch, United, Institute of Geology, Geophysics and Mineralogy 848, 1–243 (in Russian). Fostowicz-Frelik, Ł., 2008. First record of Trogontherium cuvieri (Mammalia, Rodentia) from the middle Pleistocene of Poland and review of the species. Geodiversitas 30 (4), 765–778. Fourvel, J.B., Fosse, P., Fernandez, P., Antoine, P.O., 2015. Large mammals of fouventsaint-andoche (Haute-Saˆ one, France): a glimpse into a late Pleistocene hyena den. Geodiversitas 37 (2), 237–266. https://doi.org/10.5252/g2015n2a5. Frech, F., 1904. Lethea Geognostica. III Teil. Band 2. Quart¨ ar, Stuttgart. ¨lteren Quart¨ Freudenberg, W., 1914. Die S¨ augetiere des a ars von Mitteleuropa mit besonderer Berücksichtigung der Fauna von Hundsheim und Deutschaltenburg in Nieder¨ osterreich etc. Geologische und Pal¨ aontologische Abhandlungen. Neue Folge 12, 1–219. Garci´ a, N.G., 2003. Osos y otros carnivoros de la Sierra de Atapuerca. Fundacion Oso de Asturias, Asturia. García, N., Virg´ os, E., 2007. Evolution of community composition in several carnivore palaeoguilds from the European Pleistocene: the role of interspecific competition. Lethaia 40, 33–44. https://doi.org/10.1111/j.1502-3931.2006.00004.x. Geraads, D., 2008. Plio-Pleistocene Carnivora of northwestern Africa: a short review. Comptes Rendus Palevol 7, 591–599. Głazek, J., Lindner, L., Wysocza´ nski-Minkowicz, T., 1976. Interglacial mindel I/mindel II in fossil-bearing karst at Kozi Grzbiet in the holy cross Mts. Acta Geol. Pol. 26, 377–393. Głazek, J., Lindner, L., Wysocza´ nski-Minkowicz, T., 1977. Old Pleistocene cave deposits z fauna at Kozi Grzbiet (holy cross Mts., Central Poland). Kras Speleol. 10, 13–28. Goldfuss, G.A., 1810. Die Umgebung von Muggendorf, ein Taschenbuch für Freunde der Natur- und Alterthumskunde. Johan Jacob Palm, Erlangen. Gorokhov, G., 1973. Tigr i drugiye khishchniki, Okhota Okhot [The tiger and other predators, Okhota Okhot]. Khozotyi 9, 16–17 (in Russian). Groiß, J., 1992. Neue Funde von quart¨ aren Gross¨ augern aus der Moggaster H¨ ohle bei Ebermannstadt (Ofr.). Archaeopteryx 10, 31–49. Groiß, J.T., 2002. Großkatzen-Reste aus der Zoolithenh¨ ohle. Geol. Blatter NordostBayern Angrenzende Geb. 52 (1–4), 85–108. Gromov, W.I., 1935. About the occurrence of the cave lion in connection with some archeological sites. Historical problems of neighbourhood areas 1–2, 165–168. Gromova, B.I., 1928. Nakhodka peshchernogo l’va v Kostromskoy gubernii [Find of a cave lion in Kostroma guberniya]. Priroda 9, 844–845 (in Russian). Gross, C., 1992. Das Skelett des H¨ ohlenl¨ owen (Panthera leo spelaea Goldfuss, 1810) aus Siegsdorf/Ldkr. Traunstein im Vergleich mit anderen Funden aus Deutschland und den Niederlanden. PhD Thesis. Tier¨ arztliche Fakult¨ at der Maximilians-Universit¨ at, München, Deutschland. Guˇzvica, G., 1998. Panthera spelaea (Goldfuss 1810) from north-western Croatia. Geol. Croat. 51, 7–14. Gürich, G., 1885. Quart¨ arfauna von Schlesien. Jahresberichte der Schlessischen Gesellschaft für Vaterl¨ andische Cultur 62, 261–270. Hagmann, G., 1899. Die diluviale Wirbeltierfauna von V¨ oklinshofen (Ober-Elsass). I. Raubtiere und Wiederk¨ auer. Straussburger Druckerei and Verlagsanstalt, Strassburg. Hank´ o, E.P., 2007. A revision of three Pleistocene subspecies of Panthera, based on mandible and teeth remains stored in Hungarian collections. Fragm. Palaeontol. Hung. 24–25, 25–43. Heller, F., 1953. Ein Sch¨ adel von Felis spelaea Goldf. aus der Frankenalb (zugleich ein Beitrag zum L¨ owe-Tiger-Problem der diluvialen Grobkatze). Erlanger Geol. Abh. 7, 1–24. Hemmer, H., 1974. Untersuchungen zur Stammesgeschichte der Pantherkatzen (Pantherinae). Teil III. Zur Artgeschichte des L¨ owen, Panthera (Panthera) leo (Linnaeus 1758). Veroeff. Zool. Staatssamml. (Muench.) 17, 167–280. Hemmer, H., 1977. Die Carnivorenreste (mit Ausnahme der Hy¨ anen und B¨ aren) aus den jungpleistoz¨ anen Travertinen von Taubach bei Weimar. Quartarpalaontologie 2, 379–387. Hemmer, H., 2001. Die Feliden aus dem Epivillafranchium von Untermaßfeld bei Meiningen (Thüringen), Teil 3. In: Kahlke, R.-D. (Ed.), Das Pleistoz¨ an von Untermaßfeld bei Meiningen (Thüringen), Teil 3. Monographien des R¨ omischGermanischen Zentralmuseums Mainz, vol. 40, pp. 699–782. Hemmer, H., 2003. Pleistoz¨ ane katzen europas-eine übersicht. CRANIUM 20 (2), 6–22.

Hemmer, H., 2004. Notes on the ecological role of European cats (Mammalia: felidae) of the last two million years. Miscelanea en homenaje a Emiliano Aquirre, Paleontologia 2, 214–232. Hemmer, H., 2011. The story of the cave lion - Panthera leo spelaea (Goldfuss, 1810) – a review. Quaternaire, Hors-s´erie 4, 201–208. Hemmer, H., Heidtke, H.J., 2013. Die Pfalz – eine frühe Station auf dem langen Weg des Jaguars, Panthera onca (Linnaeus, 1758), nach Südamerika. Jahresberichte und Mitteilungen des Oberrheinischen Geologischen Vereins. Neue Folge 95, 373–390. https://doi.org/10.1127/jmogv/95/2013/373. Hemmer, H., Keller, T., 2011. Die L¨ owin von Rockenberg (Wetterau, Hessen) im Kontext von 200 Jahren Forschung am H¨ ohlenl¨ owen, Panthera leo spelaea (Goldfuss, 1810). Geologischen Jahresberichte Hessen 137, 55–64. ¨ Hemmer, H., Schütt, G., 1970. K¨ orpergr¨ osse und Extremit¨ atenmasse Altest-und Altpleistoz¨ aner Europ¨ aischer Pantherkatzen (Genus Panthera). Mainz. Naturwissenschaftliches Arch. 9, 132–146. Hensel, R., 1852. Über die fossile S¨ augetiere Schlesiens. Jahresberischte des Schlesien Gesellschaft für Vaterl¨ andische Cultur 30, 37–38. Hensel, R., 1853. Übersicht der fossilen und lebenden S¨ augethiere Schlesiens. Denkschrift 50-j¨ ahrige Bestehens der Schlesien Gesellschaft für Vaterl¨ andische Cultur 1853, 239–250. Heptner, V.G., Naumov, N.P., 1967. Mlekopitayushchiye Sovetskogo Soyuza. Tom II, Chast’ 2. Plotoyadnyye (Giyeny I Koshki) [Mammals of the Soviet Union]. Volume II, Part 2. Carnivora (Hyenas and Cats). Vysshaya Shkola Publishers, Moscow (in Russian). Hermann, R., 1911. Rhinoceros merckii J¨ ager, im Diluvium Westpreussens und seine Beziehungen zu norddeutschen Diluvialfauna. Zeitschrift der Deutschen Geologischen Gesellschaft zu Berlin 63 (1), 13–35. Hermann, R., 1913. Die Rhinocerosarten des Westpreussischen Diluviums. Morphologisch-anatomische und biologische Untersuchungen. Schriften der Naturforschenden Gesellschaft in Danzig, Neue Funde 13 (3–4), 110–174. Hilzheimer, M., 1922. Die systematische Stellung von Felis spelaea Goldfuss. Sitzungsberichte Ges. Naturforschender Freunde Berl. 1924, 11–65. Hilzheimer, M., 1927. Zwei Radien von Felis spelaea aus der Mark Brandenburg. Zeitschrift der Geschiebeforschung 3, 79–81. Hoyer, 1937. Fauna dyluwialna polski. Kosmos 62 (3), 181–210. Jaeckel, O., 1927. Pr¨ ahistorische L¨ owen aus dem Formenkreis der Felis spelaea. Zool. Anz. 120 (9–10), 225–236. J´ anossy, D., 1969a. Stratigraphische Auswertung der europ¨ aischen mittelpleistoz¨ anen Wirbeltierfauna. Teil I. Berichte der deutschen Gesellschaft für geologische Wissenschaft. Abteilung A, Geologie und Pal¨ aontologie 4, 367–438. J´ anossy, D., 1969b. Stratigraphische Auswertung der europ¨ aischen mittelpleistoz¨ anen Wirbeltierfauna. Teil II. Berichte der deutschen Gesellschaft für geologische Wissenschaft. Abteilung A, Geologie und Pal¨ aontologie 5, 573–643. J´ anossy, D., 1990. Vertebrate fauna of site II. In: Kretzoi, M., Dobosi, V.T. (Eds.), Vertessz¨ ol¨ os. Site, Man and Culture. Akademiai Kiado, Budapest, pp. 187–219. Kabitzsch, J.F., 1960. Die Verwandtschaft von L¨ owe und Tiger dargestellt in ihrem Gebib unter Berücksichtigung der Gebisse von Jaguar und den zwei pleistoz¨ anen Grobkatzen Felis spelaea und Felis atrox. S¨ augetierkundliche Mitteilungen 8, 103–140. Kafka, J., 1903. Fossile und Recente Raubthiere B¨ ohmens (Carnivora). Archiv der Naturwissenschaftlichen Landesdurchforschung von B¨ ohmem 10 (6), 1–120. Kahlke, R.-D., Garcia, N., Kostopoulos, D.S., Lacombat, F., Lister, A.M., Mazza, P.P.A., Spassov, N., Titov, V.V., 2011. Western Palaearctic palaeoenvironment conditions during communities, and implications for hominin dispersal in Europe. Quat. Sci. Rev. 30, 1368–1395. https://doi.org/10.1016/j.quascirev.2010.07.020. ˇ a, V., Roblíˇckov´ K´ an a, M., 2013. Svrchnˇ e pleistocenní fauna Pruˇzinsk´e Dúpn´ e jeskynˇe ve svˇ etle nových n´ alezů. Spravodaj Slovenskej Speleologickej Spoloˇcnosti, Prieskum a výskum 4, 4–15. Kempe, S., D¨ oppes, D., 2009. Cave bear, cave lion and cave hyena skulls from the public collection at the Humboldt Museum in Berlin. Acta Carsol. 38 (2–3), 253–264. https://doi.org/10.3986/ac.v38i2-3.126. Kirillova, I.V., Tesakov, A.S., 2008. New mammalian elements of the ice age assemblage on the sakhalin island ice age mammals from sakhalin. Mamm. Stud. 33, 87–92. https://doi.org/10.3106/1348-6160(2008)33[87:NMEOTI]2.0.CO;2. Kirillova, I.V., Zelenkov, N., Tesakov, A.S., 2009. Master and visitors of the cave ostantsevaya (sakhalin island, Russian far east). Acta Carstologica Slovaca 47 (1), 57–66. Kl¨ ahn, H., 1922. Ein Fund von Felis leo im L¨ oss von Heitersheim i.B. nebst einer variationsstatistischen Untersudiung bezüglich der Stellung des Fossils zu anderen Feliden. Mitteilungen des Grossherzogtums der Badenischen Geologischen Landesanstalt 9 (1), 353–366. Kleczko, S., 1999. Position du gisement de la Belle-Roche dans le contexte de l’´evolution des faunes mammaliennes continentales en Europe. M´emoire de Licence de biologie animale. Universit´e de Li` ege, Liege. ` l’`etude de Felis spelaea Goldf. Verhandlungen der Koby, F.E., 1941. Contribution a Naturforschenden Gesellschaft Basel 52, 168–188. Koch, P.L., Barnosky, A.D., 2006. Late Quaternary extinctions: state of the debate. Annu. Rev. Ecol. Evol. Syst. 37, 215–250. https://doi.org/10.1146/annurev. ecolsys.34.011802.132415. Kotze, R., Keith, M., Winterbach, ChW., Winterbach, H.E.K., Marshal, J.P., 2018. The influence of social and environmental factors on organization of African lion (Panthera leo) prides in the Okavango Delta. J. Mammal. 99 (4), 845–858. https:// doi.org/10.1093/jmammal/gyy076. Kostoglod, V.E., 1977. Otnosheniya amurskogo tigra s burym i chernym medvedem v Primor’ye [Relations between the Amur tiger with brown and black bear in

25

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Primorye]. In: Rare Species of Mammals and Their Conservation. Nauka Press, Moscow, pp. 131–133 (in Russian). Kostoglod, V.E., 1981. Opyt mnogoletnego rozyska ne zimuyushchego burogo medvedya v Sikhote-Aline [An experience of long-term tracing of a non-hibernating brown bear in the Sikhote-Alin]. Bulletin of Moscow Society of Naturalists Biological series 86 (1), 3–12 (in Russian). Kowalski, K., 1959. Katalog Ssak´ ow Plejstocenu Polski. Pa´ nstwowe Wydawnictwo Naukowe, Warszawa, Wrocław. Kowalski, K., 1972. Fossil fauna. Folia Quat. 41, 45–49. Kurt´ en, B., 1960. The age of the Australopithecinae. Stock. Contrib. Geol. 6, 9–22. Kurt´ en, B., 1968. Pleistocene Mammals of Europe. Weidenfeld and Nicolson, London. Kurt´ en, B., 1985. The Pleistocene lion of beringia. Ann. Zool. Fenn. 22 (1), 117–121. Kurt´ en, B., Poulianos, N., 1977. New stratigraphic and faunal material from Petralona Cave with special reference in the Carnivora. Anthropos 4, 47–130. Kurt´ en, B., Poulianos, A.N., 1981. Remarks on the middle and lower Pleistocene vertebrate fauna in the petralona cave. Anthropos 8, 57–72. Lehmann, U., 1954. Die Fauna des „Vogelherds” bei Stetten. Neues Jahrbuch Geol. Palaontol. Abhand. 99, 33–146. Leonardi, P., 1935. Nuovi resti di mammiferi pleistocenici della caverna Pocala (carso triestino). Atti Mus. Civico Storia Nat. Trieste 13, 1–26. Lewis, M., Pacher, M., Turner, A., 2010. The larger carnivora of the west Runton freshwater bed. Quat. Int. 228, 116–135. https://doi.org/10.1016/j. quaint.2010.06.022. Lindner, L., 1982. South-polish glaciations (nidanian, sanian) in southern Central Poland. Acta Geol. Pol. 32, 163–177. Lipecki, G., Miękinia, B., Wojtal, P., 2001. Fauna kopalna jaskini Łokietka. In: Partyka, J. (Ed.), Badania Naukowe W Południowej Czę´sci Wy˙zyny Krakowsko-Częstochowskiej. Materiały Konferencyjne, Referaty, Postery, Sesje Terenowe. Ojc´ ow, 10-11 Maja 2001, pp. 277–280. Lipecki, G., Wolsan, M., 2003. Carnivorans (carnivora). In: Valde-Nowak, P., Nadachowski, A., Madeyska, T. (Eds.), Obłazowa Cave. Human Activity, Stratigraphy and Palaeoenvironment. Institute of Archaeology and Ethnology, Polish Academy of Sciences, Krak´ ow, pp. 125–129. Lisiecki, L.E., Raymo, M.E., 2005. A Pliocene-Pleistocene stack of 57 globally distributed 18 benthic δ O records. Paleoceanography 20, PA1003. https://doi.org/10.1029/ 2004PA001071. Longin, R., 1971. New method of collagen extraction for radiocarbon dating. Nature 230, 241–242. Lubicz-Niezabitowski, E., 1925. Les restes du Lion des Cavernes (Felis leo var. spelaea Goldfuss) en Pologne. Bulletin de la Soci´ et´e des amis des sciences de Pozna´ n, S´erie В 1925 (1), 1–8. Lubicz-Niezabitowski, E., 1938a. O kilku ciekawszych szczątkach kopalnych zwierząt ssących Polski. Kosmos 63 (4), 191–198. Lubicz-Niezabitowski, E., 1938b. O kilku ciekawszych szczątkach kopalnych zwierząt ssących Polski. Sprawozdania Pozna´ nskiego Towarzystwa Przyjaci´ oł Nauk 11, 77–78. Lumley, H.d., Kahlke, H.-D., Moigne, A.-M., Moull´e, P.-E., 1988. Les faunes de grands mammif` eres de la Grotte du Vallonnet, Roquebrune-Cap-Martin, Alpes Maritimes. L’Anthropologie 92 (2), 465–496. Lupt´ ak, P., 2009. Extern´ a variabilita a taxon´ omia súcasných a vyhynutých poddruchov leva (Panthera leo). Gazella 36, 33–150. Malez, M., 1963. Kvartarna fauna pecine veternice u medvednici. Palaeontol. Jugosl. 5, 1–193. Manuel, deM., Barnett, R., Sandoval-Velasco, M., Yamaguchi, N., Vieira, F.G., Mendoza, M.L.Z., Liu, S., Martin, M.D., Sinding, M.-H.S., Mak, S.S.T., Carøe, C., Liu, S., Guo, C., Zheng, J., Zazula, G., Baryshnikov, G., Eizirik, E., Koepfli, K.-P., Johnson, W.E., Antunes, A., Sicheritz-Ponten, T., Gopalakrishnan, S., Larson, G., Yang, H., O’Brien, S.J., Hansen, A.J., Zhang, G., Marques-Bonet, T., Gilbert, M.T.P., 2020. The evolutionary history of extinct and living lions. Proc. Natl. Acad. Sci. Unit. States Am. 117 (20), 10927–10934. https://doi.org/10.1073/pnas.1919423117. Marciszak, A., Gornig, W., Stefaniak, K., 2017. Large mammals (carnivores, artiodactyls) from solna jama cave (bystrzyckie Mts, southwestern Poland) in the context of faunal changes in the postglacial period of central Europe. Palaeontol. Electron. 20 (3A), 1–37. https://doi.org/10.26879/581. Marciszak, A., Kotowski, A., Przybylski, B., Badura, J., Wi´sniewski, A., Stefaniak, K., 2019a. Large mammals from historical collections of open-air sites of Silesia (southern Poland) with special reference to carnivores and rhinoceros. Hist. Biol. 31 (6), 696–730. https://doi.org/10.1080/08912963.2017.1388377. Marciszak, A., Schouwenburg, Ch, Darga, R., 2014. Decreasing size process in the cave (Pleistocene) lion Panthera spelaea (Goldfuss, 1810) evolution – a review. Quat. Int. 339 (340), 245–257. https://doi.org/10.1016/j.quaint.2013.10.008. Marciszak, A., Schouwenburg, Ch, Gornig, W., Lipecki, G., Mackiewicz, P., 2019b. Morphometric comparison of Panthera spelaea (Goldfuss, 1810) from Poland with the lion remains from Eurasia over the last 700 ka. Quat. Sci. Rev. 223, 105950. https:// doi.org/10.1016/j.quascirev.2019.105950. Marciszak, A., Schouwenburg, Ch, Lipecki, G., Talamo, S., Shpansky, A., Malikov, D., Gornig, W., 2019c. Steppe brown bear Ursus arctos “priscus” from the late Pleistocene of Europe. Quat. Int. 534, 158–170. https://doi.org/10.1016/j.quaint.2019.02.042. Marciszak, A., Socha, P., Nadachowski, A., Stefaniak, K., 2011. Carnivores from Bi´snik cave. Quaternaire. Hors-s´ erie 4, 101–106. Marciszak, A., Stefaniak, K., 2010. Two forms of cave lion: middle Pleistocene Panthera spelaea fossilis Reichenau, 1906 and upper Pleistocene Panthera spelaea spelaea Goldfuss, 1810 from the Bi´snik cave, Poland. Neues Jahrbuch Geol. Palaontol. Abhand. 258, 339–351. https://doi.org/10.1127/0077-7749/2010/0117. Marciszak, A., Sobczyk, A., Kasprzak, M., Gornig, W., Ratajczak, U., Wi´sniewski, A., Stefaniak, K., 2020. Taphonomic and paleoecological aspects of large mammals from

Sudety Mts (Silesia, SW Poland), with particular interest to the carnivores. Quat. Int. 546, 42–63. https://doi.org/10.1016/j.quaint.2019.11.009. Marciszak, A., Stefaniak, K., Gornig, W., 2016. Fossil theriofauna from the Sudety Mts (SW Poland). The state of research. CRANIUM 33 (1), 31–41. Marks, L., Dzier˙zek, J., Janiszewski, R., Kaczorowski, J., Lindner, L., Majecka, A., Makos, M., Szymanek, M., Tołoczko-Pasek, A., Woronko, B., 2016. Quaternary stratigraphy and palaeogeography of Poland. Acta Geol. Pol. 66 (3), 403–427. https://doi.org/10.1515/agp-2016-0018. Martin, R., 1968. Les Mammif` eres fossiles du gisement quaternaire de Villereversure ´ (Ain). Etude des carnivores, des cervid´es et des ´ equid´es. Documents des Laboratoires de G´ eologie de la Faculte de Lyon 27, 57–69. Martin, P.S., Steadman, D.W., 1999. Extinctions in near time. In: MacPhee, R.D.E. (Ed.), Causes, Contexts, and Consequences. Kluwer Academic Plenum, New York, pp. 17–55. Masseti, M., Mazza, P.P.A., 2013. Western European Quaternary lions: new working hypotheses. Biol. J. Linn. Soc. 109, 66–77. https://doi.org/10.1111/bij.12032. Maz´ ak, V., 1979. Die Tiger. A. Ziemsen Verlag, Wittenberg Lutherstadt, p. 228. Maz´ ak, V., 1981. Panthera tigris. Mamm. Species 152, 1–8. Miquelle, D., Nikolaev, I., Goodrich, J., Litvinov, B., Smirnov, E., Suvorov, E., 2005. Searching for the co-existence recipe: a case study of conflicts between people and tigers in the Russian Far East. In: Woodroffe, R., Thirgood, S. (Eds.), People and Wildlife: Conflict or Co-existence? Cambridge University Press, Cambridge, pp. 305–322. Mostecký, V., 1969. Jungpleistoz¨ ane S¨ augetiere aus der “Chlup´ aˇc-H¨ ohle” auf dem Hügel “Kobyla” bei Konˇeprusy (B¨ ohmischer Karst). Acta Musei Nationalis Pragae 25 (1), 1–56. Mottl, M., 1949. Die pleistoz¨ ane S¨ augetierfauna des Frauenlochs im R¨ atschgraben bei Stübing. Verh. Geol. Bundesanst. 1947, 94–120. Musil, R., 1960. Die pleistoz¨ ane fauna der Barov´ a h¨ ohle. Anthropos 10 (3), 1–37. ˇ Musil, R., 1964. Die Braunb¨ aren aus dem Ende des lezten Glazials. Casopis Moravsk´eho Musea, Vˇ edy pˇrirodni 49, 83–102. Musil, R., 1969. Eine karstspalte mit mittelpleistoz¨ anen Funden im Kalksteinbruch ˇ ˇ Zernav´ a. Casopis moravsk´eho musea. Vˇedy pˇrírodní 54, 85–96. ˇ Musil, R., 1996. Certova pec a jeji fauna. Slov. Kras. 34, 5–56. Musil, R., 2018. Phenological analysis of the Last Glacial vertebrates from the territory of Moravia (the Czech Republic). Continuity and change in faunistic communities. Fossil Imprint 74 (3–4), 199–236. https://doi.org/10.2478/if-2018-0015. Nadachowski, A., 1985. Biharian voles (arvicolidae, rodentia, mammalia) from Kozi Grzbiet (Central Poland). Acta Zool. Cracov. 29, 13–27. Nadachowski, A., 1990. Review of fossil rodentia from Poland (mammalia). Senckenberg. Biol. 70, 229–250. Nadachowski, A., Marciszak, A., Ridush, B., Stefaniak, K., Wilczy´ nski, J., Wojtal, P., 2015. Ekspoatacja zasob´ ow fauny przez paleolityczne społeczno´sci łowieckozbierackie na przykładzie strefy pery- i metakarpackiej. In: Łanczont, M., Madeyska, T. (Eds.), Paleolityczna Ekumena Strefy Pery- I Metakarpackiej. Wydawnictwo UMCS, Lublin, pp. 837–910. ˙ Nadachowski, A., Zarski, M., Urbanowski, M., Wojtal, P., Miękina, B., Lipecki, G., Ochman, K., Krawczyk, M., Jakubowski, G., Tomek, T., 2009. Late Pleistocene Environment of the Częstochowa Upland (Poland) Reconstructed on the Basis of Faunistic Evidence from Archaeological Cave Sites. Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krak´ ow, p. 112. Nagel, D., 1997. Panthera pardus und Panthera spelea (Felidae) aus der H¨ ohle von Merkenstein/Nieder¨ osterreich. Wissenschaftliche Mitteilungen aus dem Nieder¨ osterreichischen Landesmuseum. Neue Funde 417 (10), 215–224. Nielbock, R., 1987. Holoz¨ ane und jungpleistoz¨ ane Wirbeltierfaunen der Einhornh¨ ohle bei Harz. PhD thesis. Department of Geology, University of Technology, Clausthal. Nowakowski, D., Stefaniak, K., 2015. Pathological changes of the cranium of a young female cave bear (Ursus spelaeus R.) – a case study (the Sudety Mts, Poland). Int. J. Osteoarchaeol. 25 (1), 119–125. https://doi.org/10.1002/oa.2271. Ognev, S.I., 1935. Mlekopitayushchiye SSSR I Sopredel’nykh Stran. Chast’ 3. Plotoyadnyye Zhivotnyye (Fissipediya I Lastonogiye). [Mammals of U.S.S.R. And Adjacement Countries]. Part 3. Carnivores (Fissipedia and Pinnipedia). Nauka Press, Moscow (in Russian). Ovodov, N.D., Tarasov, A.Y., 2009. Bol’shoy kot (Panthera sp.) i malen’kiy peshchernyy medved’ v Sibiri, Yeniseyskaya oblast’ [Big cat (Panthera sp.) and a small cave bear in Siberia, Yenisej Province]. Paleontology and Speleology 4, 86–92 (in Russian). Ovodov, N.D., Zaika, A.I., 2008. Cherep peshchernogo l’va (Panthera spelaea Goldfuss, 1810) iz yuzhnoy chasti Sredney Sibiri [The cranium of the cave lion (Panthera spelaea Goldfuss, 1810) from south part of the central Siberia region]. Faunae and Florae of Northern Eurasia in the Late Cenozoic 18, 158–164 (in Russian). Ossowski, G., 1887. Fouilles de la Caverne de Wierzchowska-Gorna en Pologne. Antiqua 2 (2–5), 37–47. Pacher, M., 2006. Large mammal remains from the Mladeˇc Caves and their contribution to site formation processes. In: Teschler-Nicola, T. (Ed.), Early Modern Humans at the Moravian Gate. The Mladeˇc Caves and Their Remains. Springer-Verlag, Wien, pp. 99–148. https://doi.org/10.1007/978-3-211-49294-9_6. Pacher, M., 2018. Prowling the mountains – alpine cave lion (Panthera spelaea) distribution and metrics. Slov. Kras. 56 (1), 57–84. Parfitt, S.A., Barendregt, R.W., Breda, M., Candy, I., Collins, M.J., Coope, G.R., Durbridge, P., Field, M.H., Lee, J.R., Lister, A.M., Mutch, R., Penkman, K.E.H., Preece, R.C., Rose, J., Stringer, C.B., Symmons, R., Whittaker, J.E., Wymer, J.J., Stuart, A.J., 2005. The earliest record of human activity in northern Europe. Nature 438, 108–1012. https://doi.org/10.1038/nature04227. Pasa, A., 1947. I mammiferi di alcune antiche brecce veronesi. Mem. Mus. Civ. Stor. Nat. Verona 1, 1–111.

26

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx Seryodkin, I.V., 2016. Behavior of brown bears during feeding in the Sikhote-Alin. Achievements in the Life Sciences 10 (1), 38–47. https://doi.org/10.1016/j. als.2016.05.003. Seryodkin, I.V., Miquelle, D.G., Goodrich, J.M., Kostyria, A.V., Petrunenko, Y.K., 2018. Interspecific relationships between the Amur tiger (Panthera tigris altaica) and brown (Ursus arctos) and asiatic black (Ursus thibetanus) bears. Bulletin of Biology 45, 853–864. https://doi.org/10.7868/S0044513417120029. Siegfried, P., 1983. Fossilien westfalens. Eiszeitliche s¨ augetiere. Eine osteologie pleistoz¨ aner großs¨ auger. Münstersche Forschungen zur Geologie und Pal¨ aontoologie 60, 1–163. Smuts, G.L., Anderson, J.L., Austin, J.C., 1978. Age determination of the Africa lion (Panthera leo). J. Zool. 185 (1), 115–146. Sonntag, P., 1919. Geologie von Westpreussen. Borntraeger, Berlin. Sotnikova, M.V., Foronova, I.V., 2014. First asian record of Panthera (leo) fossilis (mammalia, carnivora, felidae) in the early Pleistocene of western siberia, Russia. Integr. Zool. 9, 517–530. https://doi.org/10.1111/1749-4877.12082. Sotnikova, M., Nikolskiy, P., 2006. Systematic position of the cave lion Panthera spelaea (Goldfuss) based on cranial and dental characters. Quat. Int. 142–143, 218–228. https://doi.org/10.1016/j.quaint.2005.03.019. Stefaniak, K., 2015. Neogene and Quaternary Cervidae from Poland. Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krak´ ow. Stuart, A.J., Lister, A.M., 2011. Extinction chronology of the cave lion Panthera spelaea. Quat. Sci. Rev. 30 (17–18), 2329–2340. https://doi.org/10.1016/j. quascirev.2010.04.023. Suire, C., 1970. Contribution ` a l’´ etude des dents de Felis spelaea Goldf. Bull. Assoc. Fr. ´ Etude Quat. 7, 243–252. Sysoev, V.P., 1960. Okhota V Dal’nevostochnoi Taiga [Hunting in the Far Eastern Taiga]. Khabarovsk. Knizhnaya Izba, Khabarovsk (in Russian). Sysoev, V.P., 1966. Po medvezh’im sledam [On the bear trail]. Knizhnaya Izba, Khabarovsk (in Russian). Terzea, E., 1965. Panthera spelaea Goldf. Pleistocenol superior din Romania. Lucraile Institutului de speologie “Emil Rakovita” 4, 251–283. Testu, A., 2006. Etude paleontologique et biostratigraphique des Felidae et Hyaenidae pleistocenes de l’Europe mediterraneenne. PhD. thesis, Facult´ e des Lettres et Sciences Humaines. Universit´ e de Perpignan, Perpignan, France. Thenius, E., 1956. Zur Kenntnis der fossilen Braunb¨ aren (Ursidae, Mammalia). ¨ Sitzungsberichte der Osterreichische Akademie der Wissenschaften. Mathematisch Naturwissenschaftliche Klasse, Abteilung 1: Biologie, Mineralogie, Erdkunde und Verwandte Wissenschaften 165 (1), 153–172. Thenius, E., 1972. Die Feliden (Carnivora) aus dem Pleistoz¨ an von Str´ ansk´ a Sk´ ala. Anthropos 20, 121–135. Tkachenko, K.N., 2012. Specific features of feeding of the Amur tiger Panthera tigris altaica (Carnivora, Felidae) in a densely populated locality (with reference to Bol’shekhekhtsirskii Reserve and its environs). Biol. Bull. 39 (3), 279–287. https:// doi.org/10.1134/S1062359012030120. Toepfer, V., 1983. Ein Oberkieferfragment des L¨ owen aus dem Travertinkomplex von Bilzingsleben, Kr. Artern, und die Fundstellen pleistoz¨ aner L¨ owen im Gebiet der DDR. Ver¨ offentlichungen des Landesmuseums für Vorgeschichte in Halle 36, 163–173. Toˇskan, B., 2007. Ostanki velikih sesalcev iz Divjih bab I: stratigrafija, taksonomija in biometrija. Opera Instituti Archaeologici Sloveniae 13, 221–279. Turner, A., 1984. Dental sex dimorphism in European lions (Panthera leo L.) of the Upper Pleistocene: palaeoecological and palaeoethological implications. Ann. Zool. Fenn. 21 (1), 1–8. Turner, A., 1997. Bears and lions of Bilzingsleben: paleoecological implications. In: Mania, D. (Ed.), Bilzingsleben V, Homo erectus, seine Kultur und Umwelt; zum Lebensbild des Urmenschen. Bad Homburg and Leipzig, Leipzig, pp. 189–196. Turner, A., 1999. Larger carnivores (mammalia, carnivora) from westbury cave. In: Andrews, P., Cook, J., Currant, A., Stringer, C. (Eds.), Westbury Cave: the Natural History Museum Excavations 1976-1984. Western Academic and Specialist Press, Bristol, pp. 175–193. Turner, A., Ant´ on, M., 1997. The Big Cats and Their Fossil Relatives. An Illustrated Guide to Their Evolution and Natural History. Columbia University Press, New York. Valensi, P., 1994. Les grands mammif`eres de la grotte du Lazaret (Nice, AlpesMaritimes). PhD. thesis. Mus´eum National d’Histoire Naturelle, Lyon, France. Vereshchagin, N.K., 1971. Peshchernyy lev i yego istoriya v Golarktike i na territorii SSSR [The cave lion and its history in the Holarctic and on the territory of the USSR]. Proceedings of the Zoological Institute Russian Academy of Science 49, 123–199 (in Russian). ˇ Wagner, J., Cerm´ ak, S., 2012. Revision of the early Middle Pleistocene bears (Ursidae, Mammalia) of Central Europe, with special respect to possible co-occurrence of spelaeoid and arctoid lineages. Bull. Geosci. 87, 461–496. https://doi.org/10.3140/ bull.geosci.1354. Warsaw, 2018. https://carnivora.net/siberian-tiger-interactions-with-brown-bea rs-asiat-t101.html access 14.05.2020. Wiszniowska, T., 1978. Panthera spelaea (Goldfuss) z jaskini nied´zwiedziej w kletnie. Acta Universitatis Wratislaviensis 24, 113–141. Wiszniowska, T., 1989. Middle Pleistocene carnivora (mammalia) from Kozi Grzbiet in the swiętokrzyskie Mts, Poland. Acta Zool. Cracov. 32, 589–630. Wojtal, 2007. Wojtusiak, K., 1953. Szczątki lwa jaskiniowego (Felis spelaea Goldfuss) z jaskini wierzchowskiej g´ ornej. Acta Geol. Pol. 3, 573–592. Woldˇrich, J.N., 1893. Fossilní zvíˇrena tursk´e maˇstale u berouna v cech´ ach a rozsedliny louv´ernesk´e ve francii. Schriftenreihe der B¨ ohmischen Akademie 2 (1), 1–16. Woldˇrich, J.N., 1897. Fossilni fauna stepní z Koˇsíˇrsk´e Bulovky u Prahy a její geologickofysiografický význam. Schriftenreihe der B¨ ohmischen Akademie 4 (1), 1.

Pautret-Homerville, C., Argant, A., Combier, J., G´ ely, B., Argant, J., 2011. La grotte aux ours de Chˆ ateaubourg (Ard`eche, France): synth` ese sur un gisement d´ecouvert il y a un si` ecle et demi. Quaternaire, Hors-s´erie 4, 185–199. Pawłowska, K., 2015b. Elephantids from Pleistocene Poland: state of knowledge. Quat. Int. 379, 89–105. https://doi.org/10.1016/j.quaint.2015.05.014. Petrbok, J., 1954. Lev jeskynní v Chlup´ aˇcovˇ e sluji na Kobyle. Ceskoslovensky Kras. 7, 193. Piccoli, G., Franco, F., Mior, S., Bacchin, M.L., Maretto, P., Taruna, M.F., 1979. Grandi carnivori fossili quaternary conservati nel museo universitario padovano di geologia e paleontologia (macherodonti, leoni, iene, orsi cavernicoli). Mem.gli Ist. Geol. Mineral. Univ. Padova 32, 4–21. Pidoplichko, I.G., 1956. Vklad v poznaniye ischeznuvshey fauny Ukrainskoy SSR [Contributions to knowledge extinct fauna of Ukrainian SSR]. In: Proceedings of Ukrainian Academy of Science. Kiev (in Russian). Piotrowska, N., Goslar, T., 2002. Preparation of bone samples in the Gliwice radiocarbon laboratory for AMS radiocarbon dating. Isot. Environ. Health Stud. 38, 1–9. https:// doi.org/10.1080/10256010208033272. Rakov, N.V., 1965. Current distribution of tigers in the Amur-Ussuri region. Zoological Zhournal 44 (3), 433–441. Rakov, N.V., 1970. Faktory smertnosti kabana i yego otnosheniya s khishchnikami v Amurskoy oblasti [Factors of mortality of the wild boar and its relations with predators in the Amur Region]. Zoological Zhournal 49 (8), 1220–1228 (in Russian). Rakovec, I., 1951. Jamski lev (Felis spelaea Goldf.) iz Postojnske jame. Slovenska Akademia Znanosti i Umetnasti 1, 127–172. ˇ Rakovec, I., 1958. Plejstocenski sesalci izj pri Crnem kalu. Razprave Slovenske Akademije Znanosti in Umetnosti 4, 365–433. Rakovec, I., 1965. Pleistocenska sesalska favna iz Risov˘ ace pri Arandjelovcu. Slovenska akademija znanosti in umetnosti. Razred za prirodoslovne in medicinske vede. Oddelek za prirodoslovne vede 8, 223–317. Reichenau, W.v., 1906. Beitr¨ age zur n¨ aheren Kenntnis der Carnivoren aus den Sanden von Mauer und Mosbach. Abhandlungen der Großherzoglichen Hessischen Geologischen Landesanstalt zu Darmstadt 4, 1–125. Reimer, P.J., Bard, E., Bayliss, A., Beck, J.W., Blackwell, P.G., Ramsey, ChB., Buck, C.E., Cheng, H., Edwards, R.L., Friedrich, M., Grootes, P.M., Guilderson, T.P., Haflidason, H., Hajdas, I., Hatt´ e, Ch, Heaton, T.J., Hoffmann, D.L., Hogg, A.G., Hughen, K.A., Kaiser, K.F., Kromer, B., Manning, S.W., Niu, M., Reimer, R.W., Richards, D.A., Scott, E.M., Southon, J.R., Staff, R.A., Turney, ChS.M., Plicht, J.v.d., 2013. IntCal13 and Marine13 radiocarbon age calibration curves, 0-50,000 years cal BP. Radiocarbon 55, 1869–1887. https://doi.org/10.2458/azu_js_rc.55.16947. Reumer, J.F.W., Rook, L., Van der Borg, K., Post, K., Moll, D., De Vos, J., 2003. Late Pleistocene survival of the saber-toothed cat homotherium in northwestern Europe. J. Vertebr. Paleontol. 23 (1), 260–262. https://doi.org/10.1671/0272-4634(2003) 23[260:LPSOTS]2.0.CO;2. Riabinin, A., 1932. O ostankakh peshchernogo l’va i giyeny iz chetvertichnykh otlozheniy Sibiri [On the remains of the cave lion and hyena from Quaternary deposits of Siberia]. Journal of Information Presidium for Association Study of Quaternary Deposits 2, 1–14 (in Russian). Riedel, A., 1982. The lion of the Tilde cave. Atti Museo Civico Storia Naturale di Trieste 34, 9–47. R¨ omer, F., 1883. Die knochenh¨ ohlen von Ojc´ ow in polen. Paleontographica 29, 1–41. Ryziewicz, Z., 1933. Ssaki dyluwialne z piaskowni pod Miłowicami. Wydawnictwo ´ Muzeum Sląskiego w Katowicach 6 (3), 49–56. Ruprecht, L., 1987. Lew jaskiniowy, Panthera spelaea (Goldfuss, 1810) nowym gatunkiem w faunie ssak´ ow kopalnych Białostocczyzny. Przeglad Zool. 31 (2), 203–208. Sabol, M., 2011a. A record of Pleistocene lion-like felids in the territory of Slovakia. Quaternaire, Hors-s´ erie 4, 215–228. Sabol, M., 2011b. Masters of the lost world: a hypothetical look at the temporal and spatial distribution of lion-like felids. Quaternaire, Horse-s´erie 4, 229–236. Sabol, M., 2014. Panthera fossilis (Reichenau, 1906) (felidae, carnivora) from za h´ ajovnou cave (moravia, the Czech republic): a fossil record from 1987–2007. Acta Musei Natl. Pragae, Ser. B - Hist. Nat. 70 (1–2), 59–70. https://doi.org/10.14446/ AMNP.2014.59. Sabol, M., Gull´ ar, J., Horv´ at, J., 2018. Montane record of the late Pleistocene Panthera spelaea (Goldfuss, 1810) from the z´ apadn´e tatry mountains (northern Slovakia). J. Vertebr. Paleontol. 38 (1), 1467921. https://doi.org/10.1080/ 02724634.2018.1467921. Sala, B., 1990. Panthera leo fossilis (V. Reich., 1906) (Felidae) de Iserna la Pineta (Pl´ eistocene moyen inf´erieur d’Italie). Geobios 23 (2), 189–194. https://doi.org/ 10.1016/S0016-6995(06)80051-3. Samson, P., Kovacs, A., 1967. Felis spelaea Goldfuss in pleistocenul superior al Bazinului Sf. Gheorghe (depresiunea Brasov). Lucraile Institutului de speologie “Emil Rakovita” 6, 211–220. Schmid, E., 1940. Variationsstatistische Untersuchungen am Gebiß pleistoz¨ aner und rezenter Leoparden und anderer Feliden. Zeitschrift für S¨ augetierekunde 15, 1–179. Schmidtgen, O., 1922. Felis pardus spec. L. aus dem Mosbacher Sand. Jahrbuch der Nassauischen Ver¨ offentlichungen für Natuurkunde 74, 51–58. Schütt, G., 1969a. Untersuchungen am Gebiß von Panthera leo fossilis (v. Reichenau 1906) und Panthera leo spelaea (Goldfuss 1810). Ein Beitrag zur Systematik der pleistoz¨ anen Großkatzen Europas. Neues Jahrbuch Geol. Palaontol. Abhand. 134, 192–220. Schütt, G., 1969b. Die jungpleistoz¨ ane Fauna der H¨ ohlen bei Rübeland im Harz. Quartar 20, 1–125. Schütt, G., Hemmer, H., 1978. Zur Evolution des L¨ owen (Panthera leo L.) im europ¨ aischen Pleistoz¨ an. Neues Jahrbuch Geol. Palaontol. Monatsh. 1978, 228–255.

27

A. Marciszak et al.

Quaternary International xxx (xxxx) xxx

Wolsan, M., 1989a. Zajęczaki-lagomorpha. Folia Quat. 59–60, 145–151. Wolsan, M., 1989b. Drapie˙zne-carnivora. Folia Quat. 59–60, 177–197. ˙ Woroncowa-Marcinowska, T., Zarski, M., Pawłowska, K., Urban, J., 2013. Rewizja I Naukowe Opracowanie Ko´sci Ssak´ ow Zgromadzonych W Kolekcjach Muzeum Geologicznego PIG-PIB. Narodowe Archiwum Geologiczne, Warszawa. Wurm, A., 1912. Beitr¨ age zur Kenntnis der diluvialen S¨ augetierfauna von Mauer a. d. Elsenz (bei Heidelberg). 1. Felis leo fossilis. Jahresberichte und Mitteilungen des Oberrheinischen Geologischen Vereines. Neue Folge 2, 77–102.

Zapfe, H., 1966. Die übrigen Carnivoren (außer H¨ ohlenhy¨ ane und H¨ ohlenb¨ ar). ¨ Denkschriften der Osterreichischen Akademie der Wissenschaften, mathematisch naturwissenschaftliche Klasse 112, 1–158. Z´ azvorka, V., 1954. Paˇzní kost lva jeskynního – Panthera (Leo) spelaea, Goldfuss – z ˇ Chlup´ aˇcovy sluje na Kobyle u Konˇ eprus (Berounsko). Casopis N´ arodního muzea 123 (2), 168–175. Zotz, L.F., 1939. Die Altsteinzeit in Niederschlesien. Kabitsch Verlag, Leipzig. ˇ Zelizko, J.V., 1918. Eine kleine L¨ owenart aus dem südb¨ ohmischen Diluvium von Wolin. Jahrbuch der Kaiserischen K¨ oniglichen Geologischen Reichsanstalt 68, 113–118.

28