The field vole, an abundant and widespread microtine rodent, is a complex comprised of three cryptic species: the short-tailed field vole (Microtus agrestis) which is present over much of Eurasia, the Mediterranean field vole (Microtus lavernedii) in southern Europe, and the Portuguese field vole (Microtus rozianus) in western Spain and Portugal. Previous research has shown high genomic differentiation of these three lineages. However, the details of the process underlying their divergence remain unknown. We analyse 70 mitogenomes and 16 nuclear genomes of modern specimens, and 83 mitogenomes and 12 nuclear genomes of ancient specimens spanning the last 75 thousand years (ka). We estimate the divergence of Portuguese from short-tailed and Mediterranean field voles to be ca. 220 ka ago and of the latter two species to be ca. 110 ka ago, earlier than previous estimates involving only modern sequences. The divergence times we obtain match those between major mitochondrial lineages of cold-adapted and steppe rodents in Europe. We find signatures of gene flow within and between field vole lineages, with some analyses suggesting a hybrid origin of the Mediterranean lineage. Ancient specimens from the Italian Peninsula reveal a previously unrecognised lineage that show evidence of genetic exchange with other populations. The pattern of genetic variation in the field vole species complex demonstrates the impact of stadial-interstadial cycles in generating recurrent episodes of allopatry and connectivity of populations, a situation which could only be revealed by our dense genomic sampling over time.
& Lstrok;abajowa Cave, located in the Krakow-Cz & eogon;stochowa Upland (southern Poland), preserves a complex sedimentary sequence that documents environmental and depositional dynamics from the early Late Pleistocene to the Holocene. The cave's infill reflects a non-continuous, multi-phase sedimentary history shaped by both natural and potentially anthropogenic processes. Through an integrated multidisciplinary approach, combining micromorphology, radiocarbon and OSL dating, paleontology, anthracology, and ancient DNA studies on fossil voles, this study reconstructs the chronological, paleoenvironmental, and paleoclimatic framework of the site. The strati-graphic sequence reveals alternating phases of slow accumulation, erosion, and redeposition, with weak anthropogenic inputs in the lower units. Faunal assemblages dominated by small mammals, birds, and molluscs reflect significant climatic oscillations between Marine Isotope Stage (MIS) 5 and MIS 2/1, with shifts from mainly temperate to cold-adapted taxa. The presence of charcoal may also indicate episodic fire activity, probably of natural origin. The stratigraphic complexity of & Lstrok;abajowa Cave provides a valuable record for reconstructing the interactions between sedimentary dynamics, climate variability, and faunal changes during the Late Pleistocene in Central Europe.
The prevailing paleobiogeographic hypothesis suggests that many steppe and tundra-steppe taxa currently found in Central Asia expanded into Europe during Pleistocene glacial periods, when open habitats dominated. However, previous studies have shown that one such species, the European narrow-headed vole (Stenocranius anglicus), diverged from its Asiatic counterparts over 200 thousand years ago, implying its prolonged isolation and survival in European refugia through the Eemian interglacial period. To test whether this was an exception or part of a broader pattern, we analysed the mitochondrial genomes from 33 Late Pleistocene and Holocene small hamster (Cricetinae) remains from Central and Western Europe, the Balkans, and Anatolia, all previously identified morphologically as grey dwarf hamster (Nothocricetulus migratorius). Contrary to expectations, 16 Late Pleistocene Central European samples were assigned to the hairy-footed hamster (Cricetiscus sungorus), a species currently restricted to northern Kazakhstan and southern Russia, whereas 17 samples from the Balkans and Anatolia belonged to the grey dwarf hamster. In both cases, the Late Pleistocene samples formed clades sister to modern populations; however, with relatively recent divergence times. This suggests population continuity or repeated expansion of hamsters from Asiatic sources rather than long-term isolation in Europe, as previously observed in narrow-headed voles. Our findings indicate that steppe species, despite occupying similar ecological niches, respond to past climate change in a species-specific manner. Therefore, broad generalisations regarding their evolutionary history may be misleading.
The DNA analysis - and later, the ancient DNA analysis - has profoundly reshaped taxonomy, shifting it from a framework based primarily on morphology to one grounded in molecular evidence. Here it is demonstrated in a bat genus Plecotus, considered until 1960, a monotypic clade, in its current view comprising at least 20 separate extant cryptic species. A rich Quaternary fossil record of the genus used to be attributed to a fossil species Plecotus abeli Wettstein-Westersheim, 1923 described from a glacial assemblage in Austrian Alps, traditionally reported as an ancestor of extant Plecotus auritus (Linnaeus, 1758). Yet, two hypotheses are under investigation: (i) the separate species status of P. abeli is not substantiated, (ii) P. abeli is identical with extant P. auritus. Additionally, with a discovery of Plecotus macrobullaris alpinus Kiefer & Veith, 2002, a third hypothesis arose merging its identity with P. abeli (iii). We tested those hypotheses with the aid of detailed phenotype comparisons of extant clades and 790 fossil specimens, including the type series of P. abeli, as well as aDNA analysis of mid-European Late Pleistocene and Holocene samples of the genus, for which 42 bone and tooth samples from 20 sites across Central Europe were used. We successfully generated 35 mitogenomes, 29 belonging to Plecotus. They all, including the P. abeli paratype, were identified as P. auritus of the Eastern European clade. Thus, both our morphological and genetic data support that P. abeli is identical with P. auritus, rejecting hypothesis (iii), while (i) and (ii) are robustly supported. In consequence, vast majority of the Early Pleistocene and all the Middle and Late Pleistocene record of the genus (identified mostly as P. abeli) seems to belong to P. auritus, indicating at the same time a broad span of ecological plasticity and adaptive capacity of that species.
Southern Europe has long been recognized as a critical refuge for faunal and human populations during the Pleistocene, offering stable climatic conditions within restricted geographic areas. This study examines local climatic variations over a chronological range of 200 ka (MIS 7-3) across three key regions: Northeastern Spain, Southeastern France, and Northeastern Italy. Using micromammals as paleoenvironmental markers and applying a bioclimatic model, we reconstructed five climatic parameters (Mean Annual Temperature, Mean Temperature of the Hottest Month, Mean Temperature of the Coldest Month, Mean Annual Precipitation, and Drought Length) to identify regional differences in climate dynamics. Results indicate that Northeastern Spain experienced higher temperatures and precipitation levels, reflecting a relatively stable Mediterranean influence that created consistent refugial conditions. In contrast, Southeastern France and Northeastern Italy showed more pronounced climatic fluctuations, influenced by the Alps and a more continental climate, resulting in harsher conditions and diverse microhabitats. These findings emphasize the variability within Southern European refuges. This study underscores the complexity of refugial dynamics and their implications for species survival during the Pleistocene.
The Late Pleistocene and Holocene climatic fluctuations profoundly influenced the demographic patterns of many species. Small mammals, particularly rodents, are well-suited for such studies due to their abundance and high environmental sensitivity. A suitable subject is the tundra vole, Alexandromys oeconomus, with changing past and present distributions across the Holarctic. Using ancient DNA, we reconstructed its phylogeography and identified 12 main lineages, including extinct lineages, which highlight greater historical variability of this species. Our analyses revealed eastward and westward expansions, extinctions, and lineage replacements driven by climate changes. Originating in Central/Western Asia, the tundra vole expanded around 110 ka (thousand years ago) into Europe, diversifying into multiple lineages. Two migrations from Central Asia to north-eastern Asia occurred at roughly 70 and 16 ka, and preceded the colonization of North America by 11.5 ka. Europe also experienced intense population turnovers, with remigrations into Western Asia. Fennoscandia was colonized three times after 15 ka by two distinct routes. These migrations were tied to climate changes, with population size increasing during warming periods and declining during cooling periods, especially during the Last Glacial Maximum. This research provides new findings on how climate and environmental shifts shaped the evolution and distribution of A. oeconomus, highlighting the resilience and adaptability of small mammals.
The Pirro Nord quarry (Apricena, Foggia, southern Italy, 41°48’6’’N, 15°23’5’ E) is a well-known paleontological and archaeological locality. In the past, the high number of fossil remains allowed to describe a highly diverse assemblage of mammals, birds, amphibians and reptile species related to Late Villafranchian and Early Biharian large and small mammal ages.The small mammal material (insectivores, rodents, and bats), excavated between 2010 and 2022, was studied to describe the small mammals and to reconstruct the environment that surrounded the site using the Habitat Weighting and the Bioclimatic model methods.The rodent assemblage is composed mainly by Allophaiomys cf. ruffoi, Apodemus sp., and Hystrix refossa, while Eliomys cf. intermedius is reported for the first time in the Pirro Nord fauna. Among insectivores, Talpa gr. minor-caeca is reported together with Asoriculus sp., Erinaceus praeglacialis, and Sorex sp. Bats are represented by Rhinolophus ferrumequinum, Rhinolophus gr. euryale-mehelyi, Myotis gr. myotis-blythii, Myotis sp., and Miniopterus schreibersii.The landscape reconstruction attested the presence of a rich and diverse environment with open areas and forest patches surrounding the Pirro Nord 13 site. The climate was slightly colder than nowadays and with higher precipitations. The presence of different ecosystems nearby the sites constituted suitable conditions for the first human dispersion in Europe, as attested in Barranco León D, Atapuerca TE7-14, and Fuente Nueva 3 in the Iberian Peninsula.The study of the small mammal assemblage contributes to define the chronological boundaries of the human occupation of Pirro Nord 13, adding important information about the context of the first peopling of the Mediterranean region between 1.6 and 1.3 million years ago.
We present the results of the first dating study of site P13 at Pirro Nord, Italy, which documents some of the earliest evidence for hominin presence in western Europe. Our multi-technique dating approach is based on a combination of palaeomagnetism, electron spin resonance (ESR), thermally-transferred optically stimulated luminescence (TT-OSL) and combined U-series/ESR applied to both fossil material and host sediment. It provides ages ranging from 627 +/- 59 to 1006 +/- 126 ka and clustering around 0.8 Ma. One additional fossil tooth collected from the nearby P10 site also returns an age within this range. Ordinarily, this outcome would naturally lend itself to the straightforward conclusion that Pirro Nord has a late Early Pleistocene age of similar to 0.8 Ma. However, this interpretation is complicated by the fact that these numerical dating results are in contradiction with the biochronological evidence, which suggests a much older age on the order of 1.3-1.7 Ma. Consequently, we explore the various potential sources of bias that could have influenced the numerical dating methods and the biochronological inferences. In particular, the critical evaluation of the palaeomagnetic data available for various sites belonging to the younger Colle Curti Faunal Unit (FU) indicates that there is non-negligible age uncertainty on the allegedly minimum age of similar to 1.1 Ma traditionally assigned to the Pirro FU. Moreover, while the combined U-series/ESR dataset could accommodate an older age for the fossil remains if uranium uptake in the dental tissues occurred relatively rapidly before the closure of the system (CSUS model), the ages obtained from the two semi-independent quartz dating methods (ESR and TT-OSL) both appear to indicate that the sediment was last exposed to sunlight about 0.8 Ma. This disparity opens up the possibility that the sediment and fossil assemblage may not be coeval. In other words, it is possible that the fossil remains may have been reworked into younger deposits that entered the karst about 0.8 Ma. Though feasible from a karst sedimentary dynamics perspective, this hypothesis is not consistent with previous taphonomic studies that indicate an absence of evidence for fossil reworking after entering the karst. At the very least, our dating results indicate that site formation processes at Pirro Nord site P13 are more complex than previously considered.
The Pirro Nord quarry (Apricena, Foggia, southern Italy, 41°48′6″N, 15°23′5″ E) is a well-known palaeontological and archaeological locality comprising an abundant and diverse Early Pleistocene vertebrate assemblage of mammals, birds, amphibians, and reptiles related to the Late Villafranchian and Early Biharian mammal ages. In this study, we describe the small mammal assemblages (insectivores, rodents, and bats) excavated between 2010 and 2022 to reconstruct the palaeoenvironment surrounding the site using Habitat Weighting and the Bioclimatic model methods. The rodent assemblage comprises mostly Allophaiomys cf. ruffoi, Apodemus sp., and Hystrix refossa, whereas Eliomys cf. intermedius was reported for the first time. Insectivores includes Talpa gr. minor-caeca; Asoriculus sp.; Erinaceus praeglacialis; and Sorex sp. Bats are represented by Rhinolophus ferrumequinum, Rhinolophus gr. euryale-mehelyi, Myotis gr. myotis-blythii, Myotis sp., and Miniopterus schreibersii. Landscape reconstruction attested to the presence of a rich and diverse environment with open areas and forest patches surrounding the Pirro Nord 13 site. The climate was slightly colder than that of current times in the same area, with higher precipitation. The presence of different ecosystems near the sites constituted suitable conditions for the first human dispersion in Europe, as already revealed by evidence from Barranco León D, Atapuerca TE7–14, and Fuente Nueva 3 elsewhere on the Iberian Peninsula. The study of small mammal assemblages improves our understanding of the chronological boundaries of the human occupation of Pirro Nord 13, adding important information in the context of the first peopling of the Mediterranean region between 1.6 and 1.3 million years ago.
Lithic use-wear analysis, through defining site function and allowing reconstructing of patterns of human occupation, can contribute to our understanding of archaeological palimpsests. The Ciota Ciara cave represents an excellent case study for this methodology. Multidisciplinary research so far conducted on the materials recovered from the atrial sector of the cave distinguishes three archaeological units from a Middle Palaeolithic occupation of the site: stratigraphic units (SUs) 13, 14, and 15. Each unit is interpreted as referring to a period of numerous, superimposed episodes of human occupation, the characteristics of which we try to reconstruct and present in this work through use-wear studies. The functional analysis of lithic industries from the upper units (13 and 14) has already been published previously; here, we report corresponding new data from the lowest level, SU 15. By comparing the use-wear results from the three units and integrating the findings with data from the geoarchaeological, palaeontological, zooarchaeological, and technological studies, we attempt to reconstruct the different phases of human occupation represented in the site through time, contributing to current interpretations regarding settlement dynamics and human behaviour in the Middle Palaeolithic of north-western Italy.
New excavations in the lower part of the sequence dated between 670 and 695 ka by 40Ar/39Ar and ESR-U-Th at Notarchirico revealed layers with lithic and bone remains attesting several phases of human occupations. Some of these occupations are located at the top of residual pebble/cobble lags along former water channels, while others are more disturbed. All the layers yield faunal and lithic remains. Here, we aim to discuss the interpretative limits of traces of hominin occupations in such Early Palaeolithic sites through a multidisciplinary approach focusing on depositional and post-depositional processes in sedimentary units applied on the micro/macro-mammal remains, artefacts (surfaces, micro-wear traces), and spatial distribution of the archaeological material. These data are then compared with those from M. Piperno’s previous excavations in the upper part of the sequence (610–670 ka). As is often the case in open-air deposits and wetland environments, the majority of the bone surface modifications are related to natural abrasion caused by trampling and water flow. By contrast, the lithic material provides more relevant results both on taphonomic processes before and after the hominin occupations and function of the site. Despite the strong impact of post-depositional processes on archaeological material, evidence of hominin activities can nonetheless be inferred, shedding light on early hominin occupations of western Europe at the MIS 17/MIS 16 transition. The discussion takes penecontemporaneous open-air sites into consideration.
Confirming ephemeral human occupation is a crucial issue in cave archaeology. The project 'Tracing human presence in caves of Polish Jura' focuses on the application of molecular methods to decode the history of past human activities in cave sediments in the Krakow-Czestochowa Upland. The results will be compared with archaeological and palaeoecological proxies.
The recent developments in palaeoecological reconstruction methods, collagen extraction of small bone samples and ancient DNA analyses led us to test new approaches to enhance the chronological resolution of past climate reconstructions inferred from small mammal assemblages. Grotta della Ferrovia (Fabriano, Ancona, Italy, 40 degrees 25'36''N, 13 degrees 0'11''E, 215 m a.s.l.) is a small cave that opens on the southern bank of the Esino river, around 45 km west of Ancona. Discovered in 1966, it is famous among scholars because it is the first locality where the fossil birch mouse was recorded in Italy and because of the remarkable archaeological lithic evidence assemblage found in the cave related to the Late Epigravettian. Across the small mammal sequence, distributed into seven layers (from GDF7 to GDF2), two main phases can be recognized: in the first phase, from GDF7 to GDF5, Microtus arvalis is dominant over Microtus agrestis, in a poorly diversified assemblage. The sequence gradually changes in GDF4 and a second phase can be observed in GDF3 and GDF2, where Apodemus gr. sylvaticus-flavicollis dominates, but combined with a higher number of taxa. From GDF7 to GDF3, few individuals of Sicista cf. subtilis and Alexandromys oeconomus are recorded. Environmental and climatic reconstructions calculated with the Habitat Weighting and the Bioclimatic methods show a dominance of open meadows (GDF7 to GDF4) that were gradually replaced by closed or semi-closed forests (GDF3 and GDF2) in a context of gradually increasing temperature, shifting from a cold phase to conditions similar to the current ones in the area. Direct radiocarbon dates on 18 rodent bones and aDNA analyses on three Microtus arvalis teeth helped us to better understand the significance of the oscillation inferred from the small mammal sequences. A robust series of nine radiocarbon measurements from GDF7 to GDF5 shows that this part of the sequence accumulated between similar to 24,600e19,60 0 cal BP (95.4% probability), which includes the final phases of the Last Glacial Maximum (LGM). The ancient DNA from Microtus arvalis confirmed the chronological framework and the presence of italic populations in this area during the LGM, similar to the slightly more ancient Grotta del Sambuco on the western side of the central Peninsula. The small mammal radiocarbon dates from GDF4 and above are less consistent with their stratigraphic position, probably related to postdepositional disturbance of the sediments, but suggest that the last part of the sequence accumulated during the Late Glacial.
Although multiple Mesolithic cave sites have been recognised in Europe, the use of such sites by Early Holocene hunter gatherers was extremely scarce north of the Carpathians. Single Mesolithic artefacts have been found thus far only in six cave sites in Poland. The rich Early Mesolithic assemblage found in Bramka Rockshelter in southern Poland seems to be quite extraordinary in such a context. The site was excavated over 50 years ago, but the results have never been published or analysed beyond a short mention of child burial found in the Mesolithic context. The paper presents new radiocarbon dates obtained for the Mesolithic occupation and the child burial, showing the Early Holocene chronology of the Komornician assemblage and the Late Bronze Age/Early Iron Age chronology of the burial. The assemblage's techno-typological analysis allowed us to identify six human occupation phases at the site, with the most intensive phase connected to the Mesolithic. Comparative analyses of other Mesolithic assemblages from the region, together with an extremely high percentage of debitage found in the Bramka Rockshelter assemblage, allow us to discuss the possibility of identifying the site as a knapping workshop located in the vicinity of Jurassic flint outcrops.
The Holocene has always been considered a crucial epoch where the major cultural steps of humankind took place. Understanding past Holocene climatic variability, shifts in vegetation, and faunal communities are among crucial challenges in predicting the upcoming changes of the natural environment. In Central Europe, the Kraków-Częstochowa Upland is one of the best study areas for detecting faunal, environmental, and climate shifts. For that reason, we choose to study the small vertebrate and mollusc communities of three archaeological sites with Holocene horizons: Sąspowska Zachodnia Cave, Ciasna Cave, and Małe Rockshelter. We analysed the faunal community shifts, and we compared our results with the major Holocene sites in the area with the correspondence and cluster analyses. The environment and climate were reconstructed through Habitat Weighting, Quantified Ecology, malacological spectra and Bioclimatic methods. Environment and climate values showed a temporary replacement of the Late Glacial tundra and grasslands with boreal forest mixed with open wetlands during the Preboreal and then stable landscapes dominated by temperate forests. Models based on rodent data show that temperature and precipitation generally increased, starting from the Boreal, to reach values near, or sometimes overwhelm, the current climate values of the area during the Atlantic and Subboreal. The climate values obtained from herpetofaunal analyses show very slight changes between the Preboreal and the Atlantic (Ciasna Cave layers 3 and 4), except for the winter temperatures, whereas results obtained for Sąspowska Zachodnia Cave are probably biased by a very low number of recovered remains. Among rodents, a major negative oscillation detected in layers related to the edge of Early Holocene was related to the Preboreal Oscillation or long-lasting effect of the Younger Dryas.
The study of faunal communities of the past, and how they have changed to become what they are now, is fundamental to the understanding of the changes in ecosystems that we are witnessing and that are coming in the future. In this paper, we present the small mammal assemblage from Grotta del Sambuco (Massa Marittima, Grosseto), belonging to two archaeologic stratigraphic Units, 5 and 6, both dated to Marine Isotope Stage 2. The assemblage from Unit 6 can be defined as “non-analogue”, due to the occurrence of the Narrow-headed Vole Lasiopodomys anglicus and the Southern Birch Mouse Sicista subtilis. Non-analogue communities (i.e., with no equivalent in present time) are frequent during Late Pleistocene in the Italian Peninsula. Starting with Grotta del Sambuco, we describe some of the different factors leading to the origins and disappearance of past non-analogue small mammal assemblages in Italy. Climatic changes and the consequent shifts in geographic distributions of the species play a major role in shaping these communities, eventually leading to extirpation and extinction of cold- or continental-adapted species.
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.
In this paper, first lower molars identified as Microtus arvalis from nine archaeological and palaeontological sites of the Italian Peninsula were analysed from a morphometric point of view, focusing on the development of the anterior part of the tooth. We used the data obtained to reveal possible patterns of evolution during the Late Pleistocene, as well as similarities and differences among the Italian samples and between Italian and European populations from the same timeframe. Factors such as the relative isolation provided by the Italian Peninsula, its milder climatic setting, intra-specific population dynamics, and the south-north displacement of Italian groups are taken into account to explain the environmental and historic causes of the peculiarities displayed by Italian samples of Microtus arvalis.
The passage between the Late Pleistocene and the Early Holocene is considered to have developed during a period of abrupt climate changes that drove large and small mammals to adapt to new environments. In this context, the southwestern Italian Peninsula is a key area to study mammal adaptation to past environmental changes. One of the key localities in this region is the Grotta del Romito, an archaeological site that bears one of the most important Late Palaeolithic and Mesolithic sequences in Europe. To understand the faunal dynamics in the region during this period, we studied the large and small mammal sequences coming from the deposit investigated in the “Rock shelter” area using environment and climate reconstruction methods such as Habitat Weighting, Bioclimatic model, and Quantified Ecology. In the lower part of the sequence, the large mammals are dominated by Capra ibex, Cervus elaphus, and Capreolus capreolus, while the upper portion is dominated by C. elaphus, Sus scrofa, and C. capreolus. This change is also visible among small mammals, where a high biodiversity is registered in the lower units, while Glis glis is dominant in the upper sequence. The environmental reconstructions show a trend from a starting condition where grasslands and rocky landscapes were widespread to another where forests were the dominant biome in the area. The climate results show an increase in temperatures, even though there are some incongruities among the models used. We correlated the Grotta del Romito – “Rock shelter” sequence to the major oscillations in climatic and environmental conditions detected in other proxies during the passage between the Late Pleistocene and the Holocene; the results allowed us to detect a first warming oscillation (STR 7-7D to STR 6A-6B), which can be related to the Bølling-Allerød Interstadial. A relatively mild and quick change in favour of open environments, together with a drop in diversity species and index, was tentatively correlated with the passage between the Younger Dryas and the Preboreal (STR 4B-4C and 4-4A). The sequence ends with STR 3, related to the Early Holocene, where forested habitats were dominant. The disappearance of environments suitable for Capra ibex may have led to the progressive extirpation of this species from the southwestern Italian Peninsula. This ungulate probably moved to more suitable ecological niches (high mountains or steep slopes) from the Bølling-Allerød Interstadial. The last individuals of this ungulate species probably disappeared from this region during the Early Holocene.