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Panthera gombaszoegensis was recorded over from 100 African and Eurasian localities, ranged between 2.5 and 0.3 Myr. Among them, seven cave sites, Żabia, Tunel Wielki, Kozi Grzbiet, Draby 3, Poludniowa, Biśnik, and Komarowa caves, dated in the range of 1.7–0.3 Myr, were those in Poland. They were located in Sudety Mts, Krakow-Czestochowa Upland, Wielun Upland and Holy Cross Mts. The material from Polish sites is not very numerous, being represented mostly by isolated teeth, metapodials and phalanges, however it is well informative diagnostically. From each site, except the Biśnik Cave, remains of a single large-sized individual, mostly of males, were found. The oldest record from Żabia Cave, dated on 1.7–1.5 Myr, because of lack of suifficient diagnostic features, was classified as P. gombaszoegensis. Bones from the other sites were assigned to Panthera gombaszoegensis gombaszoegensis, late Early and Middle Pleistocene chronoform widespread in Eurasia. Analysis of a large series of teeth size in the course of time does not showed any particular changes, and the size of the Eurasian jaguar was more or less comparatively large through its entire timespan occurrence. Panthera gombaszoegensis was ecologically flexible species able to hunt in a wide prey spectrum; its size, comparable with that of a small lion Panthera leo (Linnaeus, 1758) or island tiger Panthera tigris (Linnaeus, 1758), guaranteed this jaguar one of the leading position among large carnivores. This was a member of stabile by 1 Myr carnivore paleoguild, single, conical-toothed cat, and it was well adapted to changed environmental conditions initiated by glacial-interglacial cycles. It seems that climate changes do not affected so much on P. gombaszoegensis as previously thought. The main factor responsible for the jaguar decline was the arrival of the Pleistocene lion, which affected negatively and caused strong competition. Because to its flexibility, jaguar was able to survive relatively longer than other ancient carnivores. Gradually as the density and abundance of the lion increased, a compact and wide geographic range started to shrink up to the form of isolated areas. Areas with the last, relict jaguar survivors were located far away one from another and overwhelmed by populations of P. s. fossilis. The final extinction of the species was between 350 and 300 kyr, and among them Polish find from the layer 19ad of Biśnik Cave should be highlighted as one of the latest Eurasian record of Panthera gombaszoegensis.

The present is a palaeobiological and taphonomic analysis of a Marine Isotope Stages (MIS) 4–3 (Late Pleistocene) assemblage of animal remains and hominin artefacts from layers 7–5 of Biśnik Cave, Czestochowa Upland, Poland. The analysis indicates that the bone assemblage is the result of a time-averaged palimpsest of both biotic and episodic abiotic events, the former consisting of many successive generations of animals and hominins that frequented the cave, and the latter including hydraulic winnowing. In fact, the taphonomic history of the fossil assemblage from Biśnik Cave's layers 7–5 is partially obscured by the overprint of hydraulic winnowing, which purportedly removed a certain amount of the original specimens. Besides evidence of cave bear deaths from non-violent, hibernation-related mortality and of occupation by generations of denning wolves and hyaenas, there is a wealth of flint artefacts, alongside remains of a few fireplaces and of a structure built in the cave by hominins to partition the cave chambers. The studied layer contains an impressive number of shed antlers, primarily of the red deer Cervus elaphus. Crocuta crocuta spelaea is normally held responsible for such accumulations of shed antlers in various European caves; Biśnik Cave's layers 7–5 will therefore simply add to the list. However, the role of accumulator of shed antlers attributed to the Pleistocene spotted hyena does not match the behaviour of its modern counterpart and seems not accounted for metabolically. The only reasonable alternative is that the antlers were collected by hominins. From this alternative perspective the cave would have functioned as a warehouse, where naturally shed antlers were stored as raw material, potentially to be shaped into tools and/or employed as tools to make other tools. The palaeobiological and taphonomic analysis presented here provides new insights into the succession of pre- and postdepositional events that involved the bone remains accumulated in the cave, as well as into the interactions between the animals and hominins of the time. More importantly, if hominins, and not hyaenas, were responsible for the amassment of the shed antlers in Biśnik Cave, this study raises doubts as to the hyaenid or human origin of other similar cave accumulations of shed antlers throughout Europe.

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 recognized, with P. s. fossilis recorded from 7 sites, dated between 750 and 240 ka and P. s. intermedia only partially recognized in Wierzchowska Gorna 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.

Funeral and ritual practices in cave sites during the Late Bronze Age and Early Iron Age have been recog-nised in multiple sites south of the Carpathians. This paper presents the first evidence for the funeral and ritual use of cave sites with such chronology north of the Carpathians. Unburned human remains dated to Ha B and Ha C/D have been identified in two cave sites Zbójecka Cave and Bramka Rockshelter ) located 500 m apart, in the Polish Jura. Additionally, a pottery deposit dated to Ha B2-C has been found in a third cave (Ciasna Cave ) situated near the aforementioned sites. The paper analyses these finds in the context of the local Lusatian culture settlement and the already recognised traces of Lusatian cave site use in the studied karstic region. The results give ground to search for more evidence of ritual cave use in the Late Bronze Age and Early Iron Age. V mnoha jeskynních lokalitách jižně od Karpat byly rozpoznány doklady pohřebních a rituálních praktik z pozdní doby bronzové a starší doby železné. Tento článek představuje první důkaz takového jednání severně od Karpat. Nespálené lidské ostatky datované do Ha B a Ha C/D byly identifikovány ve dvou lokalitách Polské Jury (Zbójecka jeskyně a převis Bramka), vzdálených 500 m od sebe. Kromě toho bylo ve třetí jeskyni ( jeskyně Ciasna), která se nachází poblíž jmenovaných, nalezen depot keramiky datované do Ha B2-C. Příspěvek tyto nálezy začleňuje do kontextu místního osídlení lužické kultury a již známých dokladů využití jeskynních lokalit ve zkoumaném krasovém regionu. Výsledky poskytují podklad pro iden tifikaci dalších důkazů rituálního využití jeskyní v pravěku

Sackdilling Cave is a karst fissure filled with a fossiliferous breccia including abundant fossil remains, mostly snails and small mammals. The taxonomic status of mustelid remains from this cave was previously regarded as controversial and, consequently, different authors listed different species from the locality. A detailed revision of the material shows the presence of five mustelid species: Meles sp., Martes vetus, Mustela strandi, Mustela palerminea, and Mustela praenivalis. Among them, especially noteworthy is the presence of the type specimen of Martes vetus. The Sackdilling holotype resembles Martes foina and differs from that of Martes martes in a short and broad viscerocranium; wide and less extended forward incisor row; compressed, short and wide temporal region; large, strongly inflated and convex tympanic bullae; and narrow P3 with weak lingual bulge. Some dental characters are specific for Martes vetus and show some intermediate values between Martes foina than Martes martes. Among them are the P4 protocone length and the M1 trigon length. The revised material from Sackdilling Cave of Mustela palerminea and Mustela praenivalis was compared with that from other Early and early Middle Pleistocene sites of Europe and showed the presence of intermediate characters. Subsequently, based on the entire mustelid assemblage and other faunal elements (mainly rodents), the age of this fauna was estimated as ca. 0.9–0.7 Ma.

The study of past environments, their ecology, and detailed changes through time has become an important task of environmental science. Records of paleoenvironment usually vary between regions owing to different influences of local climate, vegetation, relief, and depositional processes. Therefore, studying local sequences thoroughly allows paleoenvironmental and paleoecological reconstructions for particular regions, but it also provides important data that complement the global record. The Żarska Cave is an exceptional site, which owing to its thick and undisturbed Holocene sediments with very rich paleobotanical and paleozoological materials has become the most complete example of the uppermost Upper Pleistocene and Holocene cave deposits in the Polish uplands. The aim of our study was to understand paleoenvironmental changes in southern Poland, from the late Glacial to the late Holocene, which has been targeted by use of a detailed geological analysis accompanied by analysis of a wide range of paleobotanical, paleozoological, and archeological assemblages. All the results obtained have permitted characterization of the paleoenvironmental changes occurring in the area of the Polish Jura during the last >15 ka years. A particularly well-represented sequence covers the Allerød interstadial, which revealed the presence of forests with associated shade-loving mollusks and rodents. The beginning of the Holocene was clearly identified with an increase of shaded forest habitats, which developed in a still relatively cold climate with continental features, and with the first appearance of mesophilous deciduous trees. During the middle Holocene, unusual evidence for maple forests is documented, which developed before the formation of beech forests, typical for the late Holocene. The obtained sequence has great significance not only for the Polish Jura region, but also has wider implications for southern Poland and the vast area of the northern Carpathian foreland.

Evidence of mobiliary art and body augmentation are associated with the cultural innovations introduced by Homo sapiens at the beginning of the Upper Paleolithic. Here, we report the discovery of the oldest known human-modified punctate ornament, a decorated ivory pendant from the Paleolithic layers at Stajnia Cave in Poland. We describe the features of this unique piece, as well as the stratigraphic context and the details of its chronometric dating. The Stajnia Cave plate is a personal 'jewellery' object that was created 41,500 calendar years ago (directly radiocarbon dated). It is the oldest known of its kind in Eurasia and it establishes a new starting date for a tradition directly connected to the spread of modern Homo sapiens in Europe.

The Micoquian is the broadest and longest enduring cultural facies of the Late Middle Palaeolithic that spread across the periglacial and boreal environments of Europe between Eastern France, Poland, and Northern Caucasus. Here, we present new data from the archaeological record of Stajnia Cave (Poland) and the paleogenetic analysis of a Neanderthal molar S5000, found in a Micoquian context. Our results demonstrate that the mtDNA genome of Stajnia S5000 dates to MIS 5a making the tooth the oldest Neanderthal specimen from Central-Eastern Europe. Furthermore, S5000 mtDNA has the fewest number of differences to mtDNA of Mezmaiskaya 1 Neanderthal from Northern Caucasus, and is more distant from almost contemporaneous Neanderthals of Scladina and Hohlenstein-Stadel. This observation and the technological affinity between Poland and the Northern Caucasus could be the result of increased mobility of Neanderthals that changed their subsistence strategy for coping with the new low biomass environments and the increased foraging radius of gregarious animals. The Prut and Dniester rivers were probably used as the main corridors of dispersal. The persistence of the Micoquian techno-complex in South-Eastern Europe infers that this axis of mobility was also used at the beginning of MIS 3 when a Neanderthal population turnover occurred in the Northern Caucasus.

Fossil bear material from the lowermost deposits of the Jasna Strzegowska cave is described and its taxonomic status assessed. Comparison with bear remains from other Early and early Middle Pleistocene localities of Eurasia shows the presence of two bear species: Ursus etruscus and U. deningeri , based on morphological and size characters of the teeth and postcranial bones. The teeth of U. deningeri from the Jasna Strzegowska cave are larger, wider and have much more complicated occlusal surfaces when compared with the teeth of U. etruscus . Both bear species are characterized by relatively short and robust metapodials, although those of U. etruscus are on average less massive. Some differences in muscle attachments and articular surfaces have also been found. Metapodials of primitive arctoid bears, that co-occurred with U. etruscus and U. deningeri , are much longer and slimmer. U. etruscus and U. deningeri probably did not live in the Jasna Strzegowska cave at the same time. Based on its time range in Eurasia and morphological characteristics, the age of U. etruscus is estimated as Early Pleistocene, between 1.8–1.3 Ma. This is its first description from Poland, and the remains described are in size and morphology indistinguishable from material from other European localities. For U. deningeri , an early Middle Pleistocene age is suggested.

Lincombian-Ranisian-Jerzmanowician (LRJ) sites are sparse, and Koziarnia Cave in Poland is one of only few such sites situated at the eastern fringe of LRJ. The aim of the recent study was to obtain new chronostratigraphic data for the LRJ industries due to their extreme scarcity in Central Europe. Although the new fieldworks did not bring newfossil directeursuch as bifacial leafpoints, a detail debitage analysis enabled identifying a presence of the ventral thinning chips in layer D, which could be identified as the LRJ assemblage-containing stratum. Besides the LRJ assemblage, strata with traces of Late Middle Palaeolithic and Early Gravettian occupation were found at the site. The radiocarbon dates of Koziarnia samples show that the archaeological settlement represent one of the oldest Gravettian stays north to Carpathians. What is more, these dates demonstrate that the cave had been alternately occupied by humans and cave bears. Additionally the radiocarbon dates indicate rather young chronology of the Jerzmanowician occupation in Koziarnia Cave (c.a. 39-36 ky cal. BP). The results confirm the possibility of long chronology of the LRJ technocomplex, exceeding the Campanian Ignimbrite event.

Marine Isotope Stage 3 is considered a period with several climate oscillations that drove the environments to rapid changes. To understand how these stadial-interstadial cycles affected southern Poland, we combined the results of eight proxies analysed in the samples from the old excavations and a new 2017 trench of Koziarnia Cave (Ojcow National Park, Krakow-Czestochowa Upland, Poland) in layers related to Middle Palaeolithic, Jerzmanowician, and Early Gravettian. Among the studied proxies were charcoals, pollen record, remains of malacofauna, and vertebrates (including rodents, birds and large mammals, and ZooMS analysis of fragmented bones). Moreover, sediment samples have been analysed for lipid composition (by Gas Chromatography-Mass Spectrometry, GC–MS). Despite several taphonomic issues, it was possible to recognise two oscillations. The first one, reflected in pollen record and lipid analysis, took place during Dansgaard-Oeschger (DO) 14 to 8 and included Heinrich Stadial (HS) 4. The second one, recorded by rodents and bird proxies, was related to DO 8/7 to DO 6 and included HS 3. Charcoal and large mammal proxies provided the broad context of our study. The Jerzmanowician occupation was connected with a relatively cold episode in a landscape characterized mainly by grassland and periglacial environments, while the Late Middle Palaeolithic and Early Gravettian groups settled the cave during milder climatic conditions, where environments were open with sparse boreal woodlands. Such trends provide additional arguments in a broad discussion on Middle-Upper Palaeolithic transition in Central Europe.

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

The deposits of Biśnik Cave yielded numerous bear remains from sediments dated to the MIS 8–MIS 3 interval. This situation allowed examination of the paleoecology of Ursus spelaeus over time, compared with two other species of bears: Ursus deningeri and Ursus arctos, also present in fossil material from Biśnik Cave. The stable isotope ratios (δ13C, δ18O) in tooth enamel were applied in combination with examination of seasonal deposition of dental cementum. The δ13C values showed the diet of each bear species stayed in the range of C3 plants. Results for U. spelaeus showed a wider range of δ13C values than for U. deningeri or U. arctos. The values are relatively negative in comparison to fossil bears from other European sites. Low δ13C values observed in bears from Biśnik Cave could be linked to a canopy effect and document a forested environment for these bears. No correlation between δ18O and δ13C values was noted. The cementum analysis revealed that season of death of all species was winter and all studied specimens died during hibernation. The highest mortality concerned mostly individuals below 4 years old and the highest individual aged was U. deningeri.

The Pleistocene lion Panthera spelaea was not a homogenous taxon; it was represented by three chronoforms. The oldest one, from the Early and Middle Pleistocene, was P. s. fossilis, the youngest was the Late Pleistocene cave lion P. s. spelaea. The intermediate form between them was P. s. intermedia. The scarcity of stratigraphically old lion made it difficult to understand the evolution of this carnivore. Therefore, we revised the fossil material of the Pleistocene lion from Polish excavation sites. The analysis revealed the presence of P. s. fossilis in 7 sites: Tunel Wielki, Poludniowa, Wschodnia, Draby, Biśnik, Deszczowa and Wierzchowska Gorna caves. The remains from these localities significantly differed in their average size and teeth proportions from the nominate younger chronoform, i.e. the cave lion. Using this material and all other available data, we compared the lion remains grouped into four palaeoclimate periods: MIS 17-12, MIS 11-9, MIS 8-6 and MIS 5-2. We found significant time-related trends in many morphometric features of dentition and limb bones. Between the first two periods, the lion became larger but from MIS 8, it gradually decreased in size. The teeth showed a much greater rate of changes than the metapodial bones, which in turn were characterised by more heterogeneous and mosaic evolution. The changes observed in the fossil material can result from in situ evolution of lion populations occupying Europe or immigration and replacement of the native forms by those from other regions, for example Asia.

The Stajnia Cave is one of the most important archaeological sites due to the finds of the first remains of Neanderthals in Poland, and several tens of thousands of flint artefacts from the Middle Palaeolithic. Based on geological, geochemical, palaeobotanical, palaeozoological, archaeological and isotope analyses, coupled with absolute age determinations (OSL, U-Th and C 14 ), 15 lithostratigraphic layers were distinguished and palaeoenvironmental conditions during the sedimentation of these beds were determined. The cave loams accumulated through weathering, aeolian and fluvial processes. Their age may be correlated with an interval from the Early Glacial to the Late Pleniglacial of the Visulian (Weichselian) Glaciation. Archaeological relics related to the Neanderthals have been discovered in layer D with an absolute age of about 52,000–45,000 years BP and correlated with MIS 3 – the Middle Plenivistulian (Interplenivistulian). Climate oscillations in the Vistulian are reflected by the type of the sediments and their physical-chemical features, allowing determining warmer interstadial and colder stadial periods. Generally, the climate was cold, characteristic of tundra areas with a typical vegetation and fauna, and with the mean temperature of the warmest month not exceeding 12°C. Based on multi-proxy studies it can be concluded that from layer E1 upwards, the climate conditions became progressively drier. At that time, the climate was cold with continental features enhancing tundra domination. This conclusion is confirmed by palaeontological investigations and the record of stable oxygen isotopes in the teeth of reindeer. The studies have also indicated seasonal migration of reindeer on the tundra that surrounded the cave. Probably, short-term slight climate warmings occurred during the Middle Plenivistulian (Interplenivistulian).

The Crimean Mountains are well known from the abundance of Middle and Late Palaeolithic sites and palaeontological remains recovered from cultural layers in caves and rockshelters. The fossil-bearing deposits of Emine-Bair-Khosar Cave, located at the elevation of 1000 m on the Chatyrdag Plateau, yielded a very diverse and numerous vertebrate remains that widen the knowledge of Late Pleistocene faunal diversity in the Crimea. The assemblage comprised in total almost 50 species of vertebrates. Studies included geomorphological, geological and stratigraphic analyses as well AMS 14C dating. Faunal remains were present in ten palaeontological sites. The main bone accumulation (section Ba2) was deposited during Middle Valdai or Vytachiv (MIS 3) interstadial, and including a long time gap, to the end of the Pleistocene and the Holocene. Comparison of the Emine-Bair-Khosar fauna with vertebrate faunas of other Crimean sites showed a remarkable stability in the faunal composition and frequency during the whole MIS 3 interstadial. Steppe and other open-country species dominated in the compared assemblages, while boreal-tundra species were far less numerous. Inhabitants of forests, including red deer and some rodents, were stable members of fossil assemblages.

The Borsuka Cave is located in the southern part of Krakow-Czestochowa Up- land, about 20 km west of Krakow. During excavations conducted in 2008-2010 a very rare and interesting faunal assemblage from layer VI was found, dating to the Upper Pleniglacial. Among cold steppe-tundra or taiga species such as Rangifer tarandus, Vulpes alopex, Equus sp. or Coelodonta antiquitatis taxa adapted to forest environment were also found. Associated with them we found snails, such as Ena montana, Aegopi- nella pura and Perforatella incarnata; insectivores, like Sorex araneus, carnivores, such as Martes martes, Meles meles and Lynx cf. lynx; ungulates, like Alces alces and Bos primigenius and, among rodents, Clethrionomys glareolus, Apodemus sylvaticus/flavicol- lis and Castor fiber. This assemblage is the first from southern Poland during this time pe- riod to comprise such relatively rich material, and indicates the presence of forest adapted species at the end of the Upper Plenivistulian. No similar assemblages are known from other caves from the Krakow-Czestochowa Upland, and this suggests that during Late Pleistocene in a limited area of Poland, short episodes of forest formations could appear.

The fossil-bearing deposits of Żabia Cave (Czestochowa Upland, Poland) yielded a diverse vertebrate fauna (75 taxa in total) of middle Early Pleistocene age. The assemblage comprises 5 amphibian, 10 reptilian and 6 bird taxa. Small mammals are represented by13 insectivorous species, 6 taxa of bats, 2 lagomorphs and 18 rodents. Large mammals include 15 species of carnivores, an unidentified horse, and 3 even-toed ungulates. The assemblage contains few Pliocene relict forms (e.g. Mioproteus wezei, Asoriculus gibberodon, Beremendia fissidens, Mimomys pitymyoides, Pannonictis ardea) and numerous Early Pleistocene newcomers (e.g. Sorex runtonensis, Ochotona zabiensis, Prolagurus ternopolitanus, early Microtus, Gulo schlosseri, Lutra simplicidens, Cervalces carnutorum). The assemblage documents migration of species associated with cool climate into Central Europe in the middle Early Pleistocene.

The first leopard (Panthera pardus) from the Pleistocene of Poland was discovered in the sediments of Biœnik Cave. Bones show morphological and metric similarity to large specimens of the modern leopard. This finding provides evidence for the existence of the leopard in late Middle and early Late Pleistocene in Europe north of the Carpathian arch.

Two forms of cave lion Panthera spelaea GOLDFUSS, 1810: Middle Pleistocene Panthera spelaea fossilis REICHENAU, 1906 and Upper Pleistocene Panthera spelaea spelaea GOLDFUSS, 1810 are reported from the Bisnik Cave (Czþ estochowa Upland, Southern Poland). A detailed examination of tooth (particularly carnassials) and mandible morphology provides a basis to discuss the inferred size trend from the earlier, bigger P. s. fossilis to the smaller and more recent chronosubspecies P. s. spelaea. The original acquisition labels and detailed stratigraphy make it possible to place these finds within an updated stratigraphic and biochronological framework. The cave lion remains from the Bisnik Cave show that the large, primitive form fossilis was replaced by the more specialized form spelaea.