Search

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.

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.

Peopling of Central Europe by Middle Pleistocene hominids is highly debatable, mainly due to the relatively harsh climatic and environmental conditions that require cultural and anatomical adjustments. At least several archaeological sites certify human occupation in the region dated back to MIS 13-11, but they represent open-air settlements. Based on the new fieldwork conducted in Tunel Wielki Cave, we can date the human occupation traces in the cave to MIS 14-12. Bipolar-on-anvil knapping technique prevails in the lithic assemblage, made exclusively in flint. The obtained results have given ground for studying the frontiers of human oikumene and the required cultural adaptive abilities.

Aim: The Expansion-Contraction model has been used to explain the responses of species to climatic changes. During periods of unfavourable climatic conditions, species retreat to refugia from where they may later expand. This paper focuses on the palaeoecology of red deer over the past 54 ka across Europe and the Urals, to reveal patterns of change in their range and explore the role of environmental conditions in determining their distribution. Location: Europe and western Asia to 63°E. Taxon: Red deer (Cervus elaphus). Methods: We collected 984 records of radiocarbon-dated red deer subfossils from the Late Pleistocene and the Holocene, including 93 original dates. For each deer sample we compiled climatic and biome type data for the corresponding time intervals. Results: During the last 54 ka changes in red deer range in Europe and the Urals were asynchronous and differed between western and eastern Europe and western Asia due to different environmental conditions in those regions. The range of suitable areas for deer during the Last Glacial Maximum (LGM) was larger than previously thought and covered vast regions not only in southern but also in western and eastern Europe. Throughout the period investigated the majority of specimens inhabited forests in the temperate climatic zone. The contribution of forests in deer localities significantly decreased during the last 4 ka, due to deforestation of Europe caused by humans. Mean January temperature was the main limiting factor for species distribution. Over 90% of the samples were found in areas where mean January temperature was above −10°C. Main conclusions: Red deer response to climatic oscillations are in agreement with the Expansion-Contraction model but in contradiction to the statement of only the southernmost LGM refugia of the species. During the last 54 ka red deer occurred mostly in forests of the temperate climatic zone. © 2020 John Wiley & Sons Ltd. European Social Fund, Grant/Award Number: UDA-POKL.04.01.01-00-072/09-00; University of Wroclaw, Grant/Award Number: 0410/2990/18; Institute of Environmental Biology, University of Wrocław, Grant/Award Number: 0410/2990/18; Mammal Research Institute Polish Academy of Sciences; Narodowe Centrum Nauki , Grant/Award Number: DEC-2013/11/B/NZ8/00888 and UMO-2016/23/B/HS3/00387; Romanian National Authority for Scientific Research, UEFISCDI, Grant/Award Number: PN-IIIP4-ID-PCE-2016-0676; National Centre for Atmospheric Science and the Centre for Environmental Data Analysis, UK; Faunal Database of the Stage Three Project; Leverhulme Trust, Grant/Award Number: F00568W.

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.

The present phylogeographic pattern of red deer in Eurasia is not only a result of the contraction of their distribution range into glacial refugia and postglacial expansion, but probably also an effect of replacement of some red deer s.l. mtDNA lineages by others during the last 50 000 years. To better recognize this process, we analysed 501 sequences of mtDNA cytochrome b, including 194 ancient and 75 contemporary samples newly obtained for this study. The inclusion of 161 radiocarbon-dated samples enabled us to study the phylogeny in a temporal context and conduct divergence-time estimation and molecular dating. Depending on methodology, our estimate of divergence between Cervus elaphus and Cervus canadensis varied considerably (370 000 or 1.37 million years BP, respectively). The divergence times of genetic lineages and haplogroups corresponded to large environmental changes associated with stadials and interstadials of the Late Pleistocene. Due to the climatic oscillations, the distribution of C. elaphus and C. canadensis fluctuated in north–south and east–west directions. Some haplotypes dated to pre-Last Glacial Maximum periods were not detected afterwards, representing possibly extinct populations. We indicated with a high probability the presence of red deer sensu lato in south-eastern Europe and western Asia during the Last Glacial Maximum.

Muskox Ovibos moschatus is a Pleistocene relic, which has survived only in North America and Greenland. During the Pleistocene, it was widely distributed in Eurasia and North America. To evaluate its morphological variability through time and space, we conducted an extensive morphometric study of 217 Praeovibos and Ovibos skull remains. The analyses showed that the skulls grew progressively wider from Praeovibos sp. to the Pleistocene O. moschatus, while from the Pleistocene to the recent O. moschatus, the facial regions of the skull turned narrower and shorter. We also noticed significant geographic differences between the various Pleistocene Ovibos crania. Siberian skulls were usually larger than those from Western and Central Europe. Eastern Europeanmuskoxen also exceeded in size those from the other regions of Europe. The large size of Late Pleistocene muskoxen from regions located in more continental climatic regimes was probably associated with the presence of more suitable food resources in steppe-tundra settings. Consistently, radiocarbon-dated records of this species are more numerous in colder periods, when the steppe-tundra was widely spread, and less abundant in warmer periods.

Revision of the Silesian bear fauna, based on material from 152 sites, mainly cave and karstic localities, and also archaeological and open-air sites, shows the presence of 13 forms and species. These records encompass the last 16.5 Ma and may be divided into five main morphophyletic groups. The oldest bears, represented by the genera Ballusia and Ursavus , are dated to 16.5–11 Ma, and belonged to the stem forms of the subfamily Ursinae. After a break of 6 My, the earliest members of the genus Ursus appeared, and all known Pliocene bears in Silesia belong to the genus Ursus were identified as Ursus minimus . They represent one or two migration events. Additionally, between 3.6 and 3.2 Ma, a single occurrence of Agriotherium insigne was recorded from the Weze 1 site. A few Early Pleistocene bear records are represented by U. etruscus , which was a probable ancestor of both arctoid and spelaeoid bear lineages. The oldest representative of Ursus ex gr. arctos known so far and assigned to U. a. suessenbornensis is known from one latest Early Pleistocene (1.2–0.9 Ma) locality, while other Polish records of this form require confirmation. The first occurrence of U. deningeri , the oldest taxon within the U . ex gr. deningeri–spelaeus lineage, was recorded from ~700 ka deposits in Silesia. During the pronounced cold period of MIS 12, the Scandinavian ice sheet covered almost the entire modern territory of Poland, with the exception of the Sudetes and the Carpathians. The accompanying drastic faunal turnover led to the formation of the pan-Eurasian Mammoth Fauna at ~460 ka. At that time a characteristic member of this fauna, the steppe brown bear Ursus arctos priscus , a specific ecomorph adapted to live in open grasslands, appeared in this region. It survived until the beginning of MIS 1, when modern Ursus arctos arctos appeared in Silesia and survived to the present day. U. deningeri was the most common bear during the Middle Pleistocene, while the first records of U. spelaeus spelaeus appeared since MIS 7. The latter form was replaced by U. ingressus during the Late Pleistocene (~110–100 ka). Spelaeoid bears totally dominated the cave assemblage, and finally vanished between 27 and 24 ka.

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.

There is a widespread belief that the abrupt warming at 14.7 ka had a profound impact on the environment. However, the direct correlation between the global climatic event and changes in local environments is not obvious. We examined faunal succession in an intra-mountain basin of the Western Carpathians to assess the potential influence of the climatic change between Greenland Stadial-2a and Greenland Interstadial-1e on the local environment. We investigated three vertebrate assemblages (total number of identified specimens = 18,745; minimum number of individuals = 7515; 138 taxa) from Obłazowa Cave (western entrance) and a Rock overhang in Cisowa Rock, radiocarbon dated to the period before and after the global warming, between ca. 17.0 and 14.0 ka. Our data revealed that the major abrupt warming that occurred 14.7 ka had little impact on the local environment, which could suggest that ecosystems in Central Europe were resilient to the abrupt global climate changes. The increase in fauna population sizes and species diversities in local biotopes was gradual and began long before the temperature increase. This was supported by the analysis of ancient DNA of Microtus arvalis, which showed a gradual increase in effective population size after 19.0 ka. The results of palaeoclimatic reconstruction pointed out that the compared sites were characterized by similar climatic conditions. According to our calculations, the differences in the annual mean temperatures did not exceed 0.5 °C and mean annual thermal amplitude changed from 22.9 to 22.4 °C. The environmental changes before 14.7 ka had no impact on the activity of Final Palaeolithic hunters in the studied area.

The woolly mammoth (Mammuthus primigenius) was widespread in almost all of Europe during the late Pleistocene. However, its distribution changed because of population fluctuations and range expansions and reductions. During Marine Oxygen Isotope Stage 2 (MIS 2), these processes were highly dynamic. Our analyses of 318 radiocarbon dates from 162 localities, obtained directly from mammoth material, confirmed important changes in mammoth range between ~28.6 and ~14.1 ka. The Greenland stadial 3 interval (27.5–23.3 ka) was the time of maximum expansion of the mammoth in Europe during MIS 2. The continuous range was soon fragmented and reduced, resulting in the disappearance of Mammuthus during the last glacial maximum from ~21.4 to ~19.2 ka in all parts of the North European Plain. It is not clear whether mammoths survived in the East European Plain. The mammoth returned to Europe soon after ~19.0 ka, and for the next 3–4 millennia played an important role in the lifeways of Epigravettian societies in eastern Europe. Mammoths became extinct in most of Europe by ~14.0 ka, except for core areas such as the far northeast of Europe, where they survived until the beginning of the Holocene. No significant correlation was found between the distribution of the mammoth in Europe and human activity.

The saiga antelope ( Saiga tatarica ), which today is a specialist steppe herbivore adapted to long-distance migrations in massive herds, appeared at irregular intervals in the Late Pleistocene Europe. Its special ecological requirements limited the number and timing of dispersal events. In the Eemian the saiga was absent on the continent. During MIS 5d-a, MIS 4 and MIS 3 its range was restricted to steppe areas north of the Black Sea. In MIS 2 and the Late Glacial (MIS 1) this species was able to colonize Central Europe at least three or four times and Western Europe at least twice. The most successful second migration dates to part of GS-2c, GS-2b and part of GS-2a, from ca. 19.5 to 15.3 cal ka BP when the saiga reached southern France. During GI-1d (Older Dryas), ca. 14.0 cal ka BP, the saiga reached England. The first migration during the GI-2 warm phase (ca. 23.7–23.3 cal ka BP) and the last one (ca. 13.5 cal ka BP) in the GI–1c 2 cool phase were probably restricted to Central Europe.

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 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.

In Central Europe, a major gap in the radiocarbon chronology of Mammuthus primigenius has resulted from the absence of dated records in most of the area north of the Sudetes and Carpathian mountains. This paper presents almost 60 directly dated mammoth remains, based both on new research and published sources from the whole of Poland. The dates, ranging from ca. 54 to 15 cal. ka BP (51.8–12.6 ka BP), are correlated with Marine Isotope Stage 3 (MIS 3) and MIS 2 of the Late Pleistocene. The mammoth was able to colonise this region at least three to four times. The number of mammoths was probably reduced during the middle of MIS 3, between ca. 43.2 cal. ka BP (39.0 ka BP) and 40.6 cal. ka BP (35.9 ka BP), as well as between ca. 34.8 cal. ka BP (30.5 ka BP) and 32.1 cal. ka BP (27.4 ka BP). Also, at the beginning of MIS 2 ca. 27.5 cal. ka. BP (ca. 22.8 ka BP) woolly mammoth probably became very rare in the studied area until ca. 24 cal. ka BP (ca. 20.2 ka BP), the beginning of H2 cold event. In the middle of MIS 2, between ca. 24.1 cal. ka BP to ca. 18.3 cal. ka BP (ca. 20.2 ka BP to ca. 14.6 ka BP), Mammuthus withdrew from Poland entirely. From ca. 18.3 cal. ka BP (ca. 14.6 ka BP) to 15.0 cal. ka BP (12.6 ka BP) the mammoth reoccupied part of its former range, the south-eastern part of Poland (Lublin Upland) and the easternmost part of Sudetes foothills and the upper Odra river valley. The marked loss of open habitats at the beginning of the Lateglacial Interstadial was followed by the retreat and extinction of M. primigenius in southern Poland around 15.0 cal. ka BP (12.6 ka BP).

A new Greek sequence of early Upper Palaeolithic, Aurignacian, Epigravettian, and Mesolithic assemblages, which differs from the sequences of Franchthi and Kephalari caves, was uncovered during the excavations in Cave 1 in Klisoura Gorge (Western Peloponnese). This is the first case of Middle Palaeolithic deposits immediately covered by an early Upper Palaeolithic assemblage. The long Middle Palaeolithic in this site underlies a long sequence of Upper Palaeolithic layers. Most interesting is the Early Upper Palaeolithic industry which contains numerous arched backed blades and other lithics demonstrating morphological affinities to the Italian Uluzzian, a resemblance that raises questions concerning the potential makers of this industry. Above it, several Aurignacian levels dated from 24 to 34 ka bp were exposed. This is the first well-dated sequence of Aurignacian occupations in Greece in which a number of basin-like hearth structures were exposed, lined with a clay that had been brought in and specially prepared. The Aurignacian sequence is covered by Epigravettian layers. The unconformity between the Epigravettian and the underlying Aurignacian corresponds to the Last Glacial Maximum. A Mesolithic layer caps the prehistoric sequence.