History of Polish Canidae (Carnivora, Mammalia) and Their Biochronological Implications on the Eurasian Background.

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 Weze 2 (2.9-2.6 myr ago), Lycaon falconeri from Rebielice 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.

Ancient DNA of narrow-headed vole reveal common features of the Late Pleistocene population dynamics in cold-adapted small mammals.

The narrow-headed vole, collared lemming and common vole were the most abundant small mammal species across the Eurasian Late Pleistocene steppe-tundra environment. Previous ancient DNA studies of the collared lemming and common vole have revealed dynamic population histories shaped by climatic fluctuations. To investigate the extent to which species with similar adaptations share common evolutionary histories, we generated a dataset comprised the mitochondrial genomes of 139 ancient and 6 modern narrow-headed voles from several sites across Europe and northwestern Asia covering approximately the last 100 thousand years (kyr). We inferred Bayesian time-aware phylogenies using 11 radiocarbon-dated samples to calibrate the molecular clock. Divergence of the main mtDNA lineages across the three species occurred during marine isotope stages (MIS) 7 and MIS 5, suggesting a common response of species adapted to open habitat during interglacials. We identified several time-structured mtDNA lineages in European narrow-headed vole, suggesting lineage turnover. The timing of some of these turnovers was synchronous across the three species, allowing us to identify the main drivers of the Late Pleistocene dynamics of steppe- and cold-adapted species.

A 41,500 year-old decorated ivory pendant from Stajnia Cave (Poland).

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 Tien Shan vole (Microtus ilaeus; Rodentia: Cricetidae) as a new species in the Late Pleistocene of Europe.

Grey voles (subgenus Microtus) represent a complex of at least seven closely related and partly cryptic species. The range of these species extends from the Atlantic to the Altai Mountains, but most of them occur east of the Black Sea. Using ancient DNA analyses of the Late Pleistocene specimens, we identified a new mtDNA lineage of grey voles in Europe. Phylogenetic analysis of mitochondrial DNA cytochrome b sequences from 23 voles from three caves, namely, Emine-Bair-Khosar (Crimea, Ukraine), Cave 16 (Bulgaria), and Bacho Kiro (Bulgaria), showed that 14 specimens form a previously unrecognized lineage, sister to the Tien Shan vole. The average sequence divergence of this lineage and the extant Tien Shan vole was 4.8%, which is similar to the divergence of grey vole forms, which are considered distinct species or being on the verge of speciation; M. arvalis and M. obscurus or M. mystacinus and M. rossiaemeridionalis. We estimated the time to the most recent common ancestor of the grey voles to be 0.66 Ma, which is over twice the recent estimates, while the divergence of the extant Tien Shan vole and the new lineage to be 0.29 Ma. Our discovery suggests that grey voles may have been more diversified in the past and that their ranges may have differed substantially from current ones. It also underlines the utility of ancient DNA to decipher the evolutionary history of voles.

New perspectives on Neanderthal dispersal and turnover from Stajnia Cave (Poland).

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.

Highly divergent lineage of narrow-headed vole from the Late Pleistocene Europe.

During the Late Pleistocene, narrow-headed voles (Lasiopodomys gregalis) inhabited Eurasia's vast territories, frequently becoming the dominant small mammal species among steppe-tundra communities. We investigated the relationship between this species' European and Asiatic populations by sequencing the mtDNA genomes of two extant specimens from Russia and 10 individuals from five Central European sites, dated to the post-LGM period. Phylogenetic analyses based on a large portion of mtDNA genomes highly supported the positioning of L. gregalis within Arvicolinae. The phylogeny based on mtDNA cytochrome b sequences revealed a deep divergence of European narrow-headed voles from Asiatic ones and their sister position against the extant L. gregalis and L. raddei. The divergence of the European lineage was estimated to a minimum 230 thousand years ago. This suggest, contrary to the current biogeographic hypotheses, that during the interglacial periods narrow-headed vole did not retreat from Europe but survived the unfavourable conditions within the refugial areas. Based on this result, we propose to establish a cryptic species status for the Late Pleistocene European narrow-headed vole and to name this taxon Lasiopodomys anglicus.

Human-mediated dispersal of cats in the Neolithic Central Europe.

Archeological and genetic evidence suggest that all domestic cats derived from the Near Eastern wildcat (Felis silvestris lybica) and were first domesticated in the Near East around 10,000 years ago. The spread of the domesticated form in Europe occurred much later, primarily mediated by Greek and Phoenician traders and afterward by Romans who introduced cats to Western and Central Europe around 2000 years ago. We investigated mtDNA of Holocene Felis remains and provide evidence of an unexpectedly early presence of cats bearing the Near Eastern wildcat mtDNA haplotypes in Central Europe, being ahead of Roman period by over 2000 years. The appearance of the Near Eastern wildcats in Central Europe coincides with the peak of Neolithic settlement density, moreover most of those cats belonged to the same mtDNA lineages as those domesticated in the Near East. Thus, although we cannot fully exclude that the Near Eastern wildcats appeared in Central Europe as a result of introgression with European wildcat, our findings support the hypothesis that the Near Eastern wildcats spread across Europe together with the first farmers, perhaps as commensal animals. We also found that cats dated to the Neolithic period belonged to different mtDNA lineages than those brought to Central Europe in Roman times, this supports the hypothesis that the gene pool of contemporary European domestic cats might have been established from two different source populations that contributed in different periods.

Retreat and extinction of the Late Pleistocene cave bear (Ursus spelaeus sensu lato).

The cave bear (Ursus spelaeus sensu lato) is a typical representative of Pleistocene megafauna which became extinct at the end of the Last Glacial. Detailed knowledge of cave bear extinction could explain this spectacular ecological transformation. The paper provides a report on the youngest remains of the cave bear dated to 20,930 ± 140 14C years before present (BP). Ancient DNA analyses proved its affiliation to the Ursus ingressus haplotype. Using this record and 205 other dates, we determined, following eight approaches, the extinction time of this mammal at 26,100-24,300 cal. years BP. The time is only slightly earlier, i.e. 27,000-26,100 cal. years BP, when young dates without associated collagen data are excluded. The demise of cave bear falls within the coldest phase of the last glacial period, Greenland Stadial 3. This finding and the significant decrease in the cave bear records with cooling indicate that the drastic climatic changes were responsible for its extinction. Climate deterioration lowered vegetation productivity, on which the cave bear strongly depended as a strict herbivore. The distribution of the last cave bear records in Europe suggests that this animal was vanishing by fragmentation into subpopulations occupying small habitats. One of them was the Kraków-Czestochowa Upland in Poland, where we discovered the latest record of the cave bear and also two other, younger than 25,000 14C years BP. The relatively long survival of this bear in karst regions may result from suitable microclimate and continuous access to water provided by deep aquifers, indicating a refugial role of such regions in the Pleistocene for many species.

Synchronous genetic turnovers across Western Eurasia in Late Pleistocene collared lemmings.

Recent palaeogenetic studies indicate a highly dynamic history in collared lemmings (Dicrostonyx spp.), with several demographical changes linked to climatic fluctuations that took place during the last glaciation. At the western range margin of D. torquatus, these changes were characterized by a series of local extinctions and recolonizations. However, it is unclear whether this pattern represents a local phenomenon, possibly driven by ecological edge effects, or a global phenomenon that took place across large geographical scales. To address this, we explored the palaeogenetic history of the collared lemming using a next-generation sequencing approach for pooled mitochondrial DNA amplicons. Sequences were obtained from over 300 fossil remains sampled across Eurasia and two sites in North America. We identified five mitochondrial lineages of D. torquatus that succeeded each other through time across Europe and western Russia, indicating a history of repeated population extinctions and recolonizations, most likely from eastern Russia, during the last 50 000 years. The observation of repeated extinctions across such a vast geographical range indicates large-scale changes in the steppe-tundra environment in western Eurasia during the last glaciation. All Holocene samples, from across the species' entire range, belonged to only one of the five mitochondrial lineages. Thus, extant D. torquatus populations only harbour a small fraction of the total genetic diversity that existed across different stages of the Late Pleistocene. In North American samples, haplotypes belonging to both D. groenlandicus and D. richardsoni were recovered from a Late Pleistocene site in south-western Canada. This suggests that D. groenlandicus had a more southern and D. richardsoni a more northern glacial distribution than previously thought. This study provides significant insights into the population dynamics of a small mammal at a large geographical scale and reveals a rather complex demographical history, which could have had bottom-up effects in the Late Pleistocene steppe-tundra ecosystem.

On the origin of the Norwegian lemming.

The Pleistocene glacial cycles resulted in significant changes in species distributions, and it has been discussed whether this caused increased rates of population divergence and speciation. One species that is likely to have evolved during the Pleistocene is the Norwegian lemming (Lemmus lemmus). However, the origin of this species, both in terms of when and from what ancestral taxon it evolved, has been difficult to ascertain. Here, we use ancient DNA recovered from lemming remains from a series of Late Pleistocene and Holocene sites to explore the species' evolutionary history. The results revealed considerable genetic differentiation between glacial and contemporary samples. Moreover, the analyses provided strong support for a divergence time prior to the Last Glacial Maximum (LGM), therefore likely ruling out a postglacial colonization of Scandinavia. Consequently, it appears that the Norwegian lemming evolved from a small population that survived the LGM in an ice-free Scandinavian refugium.

Late cenozoic history of the genus Micromys (mammalia, rodentia) in Central Europe.

Molecular phylogeography suggests that Micromys minutus, the sole extant species of the genus, colonized its extensive range quite recently, during the Late Pleistocene-Holocene period. Rich Pliocene and Pleistocene fossil records both from Europe and China suggest rather continuous and gradual in situ phenotype rearrangements from the Pliocene to the Recent periods. To elucidate the discrepancy we reexamined a considerable part of the European fossil record of the genus (14 sites from MN15 to Q3, 0.4-4.2 Ma, including the type series of M. preaminutus from MN15 Csarnóta 2), analyzed them with the aid of detailed morphometric comparisons, and concluded that: (a) The European Pliocene form, M. praeminutus, differs significantly from the extant species; (b) it exhibits a broad phenotypic variation covering the presumptive diagnostic characters of MN16 M. caesaris; (c) despite having smaller dimensions, the Early and Middle Pleistocene forms (MN17-Q3, 2.6-0.4 Ma) seem to be closer to M. praeminutus than to the extant species; (d) the extinction of M. praeminutus during Q3 and the re-occupation of its niche by the recent expansion of M. minutus from E-European-C Asiatic sources (suggested by phylogeographic hypotheses) cannot be excluded. Discussing interpretations of the phylogenetic past of the genus we emphasize the distinct history of the West Palearctic clade (Late Miocene-Early Pleistocene) terminating with M. praeminutus and the East Asiatic clade (chalceus, tedfordi, minutus), and the possible identity of the Western clade with the Late Miocene genus Parapodemus.

The first Neanderthal tooth found North of the Carpathian Mountains.

An upper second permanent molar from a human was found alongside numerous tools of the Micoquian tradition and was excavated in Stajnia Cave, which is located over 100 km North of the Carpathian Mountains in southern Poland. The age of these finds has been established within a time-span of late Saalian to early Weichselian, most likely to OIS 5c or 5a, according to the palaeontological, geological, archaeological and absolute dating of the layer from which they were obtained. An examination of the morphology of the human molar indicates that this tooth exhibits many traits frequently occurring in Neanderthal upper molars. Although the occurrence of derived Neanderthal traits in the Stajnia molar cannot be firmly established because of degradation of its cusps, the presence of the above-mentioned features allows the assertion that this tooth belonged to a Neanderthal. The age of the Stajnia tooth and the archaeological context of this find also indicate that this molar is of Neanderthal origin.

Molecular phylogeny of the speciose vole genus Microtus (Arvicolinae, Rodentia) inferred from mitochondrial DNA sequences.

Voles of the genus Microtus represent one of the most speciose mammalian genera in the Holarctic. We established a molecular phylogeny for Microtus to resolve contentious issues of systematic relationships and evolutionary history in this genus. A total of 81 specimens representing ten Microtus species endemic to Europe as well as eight Eurasian, six Asian and one Holarctic species were sequenced for the entire cytochrome b gene (1140 bp). A further 25 sequences were retrieved from GenBank, providing data on an additional 23, mainly Nearctic, Microtus species. Phylogenetic analysis of these 48 species generated four well-supported monophyletic lineages. The genus Chionomys, snow voles, formed a distinct and well-supported lineage separate from the genus Microtus. The subgenus Microtus formed the strongest supported lineage with two sublineages displaying a close relationship between the arvalis species group (common voles) and the socialis species group (social voles). Monophyly of the Palearctic pitymyid voles, subgenus Terricola, was supported, and this subgenus was also subdivided into two monophyletic species groups. Together, these groupings clarify long-standing taxonomic uncertainties in Microtus. In addition, the "Asian" and the Nearctic lineages reported previously were identified although the latter group was not supported. However, relationships among the main Microtus branches were not resolved, suggesting a rapid and potentially simultaneous radiation of a widespread ancestor early in the history of the genus. This and subsequent radiations discernible in the cytochrome b phylogeny, show the considerable potential of Microtus for analysis of historical and ecological determinants of speciation in small mammals. It is evident that speciation is an ongoing process in the genus and that the molecular data provides a vital insight into current species limits as well as cladogenic events of the past.