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The remains of 12 canid species that date back ca. 4.9 myr have been found at 116 paleontological localities. Among these localities, eight are dated to the Pliocene age, 12 are dated to the Early Pleistocene age, 12 are from the Middle Pleistocene age, while the most numerous group includes 84 sites from the Late Pleistocene–Holocene age. Some, especially older forms such as Eucyon odessanus and Nyctereutes donnezani, have only been found at single sites, while the remains of species from the genus Lycaon, Canis and Vulpes have been recorded at numerous sites from the last 2 myr. Ancient canids such as Eucyon and Nyctereutes had already vanished from Poland in the Earliest Pleistocene, between 2.5 and 2.2 myr ago. Poland’s extant canid fauna is characterised by the presence of two new species, which spread into the territory due to a human introduction (Nyctereutes procyonoides) or natural expansion (Canis aureus). Research indicates a strong competition between dogs, especially between Lycaon, Canis and Cuon, with a strong lycaon-limiting effect on the wolf between 2.5 and 0.4 myr ago. After the extinction of Lycaon lycaonoides, Canis lupus evolved rapidly, increasing in number and size, and taking over the niche occupied by Lycaon. In order to reduce competition, the body size of Cuon alpinus gradually reduced, and it became an animal adapted to the forest, highland and mountain environments. Generally, the history of canids in Poland is similar to that known of Eurasia with some noteworthy events, such as the early occurrence of Canis cf. etruscus from Węże 2 (2.9–2.6 myr ago), Lycaon falconeri from Rębielice Królewskie 1A or one of the latest occurrences of L. lycaonoides from Draby 3 (430–370 kyr). Predominantly lowland or upland in the southern part and devoid of significant ecological barriers, Poland is also an important migration corridor in the East–West system. This 500–600 km wide corridor was the Asian gateway to Europe, from where species of an eastern origin penetrated the continent’s interior. In colder periods, it was in turn a region through which boreal species or those associated with the mammoth steppe retreated.

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.

The Pleistocene history of the leopard (Panthera pardus) in Europe has been documented by the material obtained from 312 localities, with the last dated ∼1.1 Myr. The relatively small and gracile form of the leopard was very rare during the late Early and Middle Pleistocene. Only after the disappearance of the jaguar (Panthera gombaszoegensis) did P. pardus spread widely in Europe, increasing in size and ecologically substituting P. gombaszoegensis. The number of late Middle Pleistocene localities with leopard remains, younger than 300 kyr, increased considerably. The leopard reached the maximum extension of its geographical range in the Late Pleistocene. The Iberian Peninsula was the last European refuge for this cat. Six sites, the Naciekowa, Obok Wschodniej, Radochowska, and Wschodnia Caves from the Sudety Mountains and the Biśnik and Dziadowa Skała Caves from the Kraków-Częstochowa Upland, have documented the presence of the leopard in Poland between MIS 10/9 and MIS 3. These records are from rocky regions with rugged terrain and are located in the territory of Silesia (southern Poland). A newly obtained radiocarbon date (43–42 kyr) from the Radochowska Cave directly confirms the occurrence of P. pardus in the Sudety Mountains in the middle part of MIS 3.

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.

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.

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.

Climate changes that occurred during the Late Pleistocene have profound effects on the distribution of many plant and animal species and influenced the formation of contemporary faunas and floras of Europe. The course and mechanisms of responses of species to the past climate changes are now being intensively studied by the use of direct radiocarbon dating and genetic analyses of fossil remains. Here, we review the advances in understanding these processes by the example of four mammal species: woolly mammoth (Mammuthus primigenius), cave bear (Ursus spelaeus s. l.), saiga antelope (Saiga tatarica) and collared lemmings (Dicrostonyx ssp.). The cases discussed here as well as others show that the migrations, range shifts and local extinctions were the main responses to climate changes and that the dynamics of these climate driven processes were much more profound than it was previously thought. Each species reacted by its individual manner, which depended on its biology and adaptation abilities to the changing environment and climate conditions. The most severe changes in European ecosystems that affected the largest number of species took place around 33–31 ka BP, during the Last Glacial Maximum 22–19 ka BP and the Late Glacial warming 15–13 ka BP.

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.