European Pleistocene Research Articles

Refining the environmental and climatic background of the Middle Pleistocene human cranium from Gruta da Aroeira (Torres Novas, Portugal)

The recent publication of a human cranium, dating to ca. 436–390 ka, from Gruta da Aroeira provides an important input to the debate about human evolution during the Middle Pleistocene in Europe and the origin of the Neanderthals. This cranium, chronologically located to Marine Isotope Stage 11c (MIS 11c), appears in association with Acheulean lithic tools and several faunal remains, among which cervids and equids are predominant. The small vertebrate assemblage associated with the Aroeira-3 cranium suggests an environment and a climate related to an open-woodland landscape and relatively mild and humid conditions, probably in connection with the end of MIS 11c. Moreover, comparison with other Middle Pleistocene Iberian sites with human remains suggests that these hominins lived in open forest areas during mild climatic substages.

The MIS 13 interglacial at Ceprano, Italy, in the context of Middle Pleistocene vegetation changes in southern Europe

Climatic and environmental changes of the Middle Pleistocene in Europe provide the context for an important phase in the evolution and dispersal of early hominins. Pollen records from terrestrial and marine sediment sequences reveal patterns not usually visible in palaeoenvironmental reconstructions from archaeological sites alone and show that hominin evolution took place against a background of marked environmental change as forests expanded and contracted in concert with global and regional climatic shifts. It is rare to find archaeological material in stratigraphically long and continuous palaeoenvironmental sequences, yet this is what is needed to enable particular phases of hominin evolution to be securely associated with climatic and environmental changes. At the well-known Middle Pleistocene archaeological site of Ceprano in Italy we have been able to produce a pollen record from the sedimentary unit directly below the stratum of the hominin remains. The new pollen data from Ceprano are presented here in full for the first time and provide a detailed picture of interglacial vegetation development during Marine Isotope Stage (MIS) 13. This evidence contributes significantly to our knowledge of the spatial heterogeneity of ecosystem responses to climatic change during this period and helps to further constrain the age of the hominin remains. New sites combining both archaeology and long, continuous high quality palaeoecological records would help clarify the relationship between hominin presence and palaeoclimatic and palaeoenvironmental conditions. In the meantime, better communication between the research communities can also go a long way to improving our understanding of the links between the two.

Unravelling an essential archive for the European Pleistocene. The human occupation in the Manzanares valley (Madrid, Spain) throughout nearly 800,000 years

Since 1862, one of the largest accumulations of Palaeolithic sites of Europe has been found in the Manzanares valley (Madrid, Spain). Virtually all the fluvial deposits preserved from the Lower Pleistocene to the end of the Late Pleistocene contain remains of human occupation, that date to the Lower Palaeolithic ("core and flake assemblages" and Acheulean), Ancient Middle Palaeolithic and Middle Palaeolithic, as well as Upper Palaeolithic (Solutrean and perhaps Aurignacian). A good deal of these assemblages lacked a timeframe sufficiently delimited to be integrated within the European Pleistocene general framework. In this paper we provide, for first time, a geochronological context to several tens of sites. Together with those sites for which this information was previously available, an actual analysis of information will be possible, and will allow research on: the variability on the behaviour of human groups throughout 800 ka in the same geographic framework; human groups with flake and core industry during the first half of the Middle Pleistocene; the Acheulean technocomplex between MIS 13/11 and MIS 7/6; the coexistence between the Acheulean and the Middle Palaeolithic towards the end of the Middle Pleistocene; the peopling dynamics of Neanderthal groups in the open air; the end of the Middle Palaeolithic and therefore late Neanderthal survival; the human occupation in the interior of the Iberian Peninsula during the Late Pleniglacial; and territory management during the Solutrean period.

New material of Sus strozzii (Suidae, Mammalia) from the Early Pleistocene of Italy and a phylogenetic analysis of suines

Sus strozzii is the only species of Suinae occurring in the first part of Early Pleistocene in Europe and western Asia. It is characterized by a large body size and several morphological similarities with extinct and extant pigs from Island South East Asia (ISEA), such as the Javan Warty Pig S. verrucosus. In this study, a new mandible of S. strozzii from the Early Pleistocene of Pantalla (central Italy) is described and the diagnosis of the species is emended. Based on 52 craniomandibular and dental morphological characters, the first phylogenetic analysis of both living and fossil Eurasian and African Suinae is carried out to reconstruct their relationships. Our results are in agreement with the literature based on molecular and/or morphological analyses. The late Miocene Microstonyx is the first taxon to branch off the monophyletic Suinae clade. The separation between the African (Kolpochoerus/Hylochoerus and Metridiochoerus/Phacochoerus) and the Eurasian (Sus) clades occurred early, probably even in the late Miocene. The inclusion of Potamochoerus in the latter group is questionable, being probably due to the retaining of similar plesiomorphic characters in this African pig and in Sus. The Wild Boar S. scrofa and the Early–Middle Pleistocene S. lydekkeri are the species of Sus that display the most numerous plesiomorphic character states. They are followed by the monophyletic group of suines that show verrucosic lower canines, which includes the Pliocene S. arvernensis, S. strozzii, and the extinct and extant verrucosic pigs from ISEA.

Early bison remains from Mygdonia Basin (Northern Greece)

The continuous enrichment of the European Pleistocene mammal fossil record leads inevitably to a comprehensive forthcoming review of the biogeographic and evolutionary history of bisons on the continent. New and revised data from the post-Olduvai (<1.8 Ma)-pre-Jaramillo (>1.0 Ma) time span of the Mygdonia basin (N. Greece), exposed here, indicate the presence of a small bison referred to as Bison cf. degiulii (Masini, Palombo & Rozzi, 2013) due to its similarities with the contemporaneous type material from the Italian Peninsula. A series of minor morphometrical changes detected across chronologically successive Mygdonia bison populations is interpreted as local adaptations to harsher environmental conditions. A re-evaluation of several morphological and biometrical features of the European early bison populations in comparison with later and extant taxa leads to the reconfirmation of some characters as taxonomically and phylogenetically important, while questioning others. The results allow recognizing B. degiulii as most likely member of the true bison clade Bison (Bison) Hamilton Smith, 1827 and reconsidering the current B. (Eobison) Flerov, 1972 concept. The data from Mygdonia basin and Northern Greece confirm the short co-occurrence of Bison Hamilton Smith, 1827 and Leptobos Rütimeyer, 1877-1878 in the Balkans and suggest the appearance of at least two bison lineages in Southern Europe between 1.7-1.5 Ma.

The fossil fallow deer Dama geiselana (Cervidae, Mammalia, upgrade to species level) in the context of migration and local extinctions of fallow deer in the Late and Middle Pleistocene in Europe

The fossil fallow deer Dama dama geiselana Pfeiffer from Neumark-Nord (Saxony-Anhalt, Germany) is upgraded to species level and discussed within the current taxonomy of recent and fossil fallow deer. Typical antler and skeletal characteristics are discussed in comparison with the recent Dama dama, Dama mesopotamica and the fossil species Dama clactoniana, Dama nestii and Dama rhenana. The phylogenetic relationship of fallow deer can be traced back to the Late Pliocene and distinguished morphologically from Cervus elaphus, Cervus nippon and Axis axis. Late and Middle Pleistocene finds from Germany are presented and discussed in the context of the finds from the Mediterranean region and Great Britain. The differences in antler morphology and bone dimensions in West German and North-East German fallow deer from the Late Pleistocene support the hypothesis of different immigration channels, on the one hand from the eastern Mediterranean along the Danube and on the other from the west along the Rhone and Rhine. In the Middle Pleistocene, Dama mesopotamica is considered as the typical fallow deer in the eastern Mediterranean, while in the west, Dama clactoniana is widespread. The hypothesis of immigration from the eastern Mediterranean is supported by the fossil record in Germany with the fallow deer from Edesheim. Conversely, Dama geiselana probably influenced East Mediterranean populations. Special tooth characteristics of Dama geiselana occur with lower frequency in Dama mesopotamica. In the Bronze Age, the fallow deer from Kastanas (Macedonia) shares antler characteristics, the high frequency of specific features of the scapula, and the astragalus with Dama geiselana. A relict population of Dama geiselana probably reached the Eastern Mediterranean at the beginning of the last cold stage.

Taxonomical revision of fossil Canis in Middle Pleistocene sites of Zhoukoudian, Beijing, China and a review of fossil records of Canis mosbachensis variabilis in China

Canis mosbachensis is a widely distributed species from the Early to Middle Pleistocene in Europe. In Eastern Asia, Canis variabilis was supposed to be a local subspecies of C. mosbachensis, but such hypothesis has never been tested in details. Here we restudied the Canis materials from type locality, Loc.1 of ZKD (Zhoukoudian) of Middle Pleistocene, as well as other localities that produced important materials of this taxon (e.g. Loc.13 of ZKD). To better evaluate the variability of the Pleistocene Canis, a series of morphotypes are described. Based on morphotype and morphometric, the present study reveals there is only one species present at Loc.1 and Loc.13 of ZKD, which is quite similar to European C. mosbachensis and should be revised as a subspecies of C. mosbachensis, namely C. mosbachensis variabilis. C. m. variabilis is more derived and less hypercarnivorous than the Early Pleistocene Canis chihliensis (as well as European Canis etruscus and Canis arnensis) in craniodental characters, whereas it is less derived and less hypercarnivorous than Canis lupus. A review of fossil records of C. m. variabilis in China was made based on these different characters. Canids from Gongwangling, Lantian of Shaanxi Province (middle or late Early Pleistocene) are the earliest known representatives of C. mosbachensis in China, which are still more primitive than C. m. variabilis from the Middle Pleistocene sites of ZKD. Reliable records of C. mosbachensis are only known from the Early Pleistocene to the late Middle Pleistocene in Northern China. C. m. variabilis is not direct ancestor to C. lupus but a close relative of the latter, while it is far from the Early Pleistocene C. chihliensis. The latter remains more primitive characters that are more similar to those of European Canis etruscus and Canis arnensis.

High-resolution paleoenvironmental context for human occupations during the Middle Pleistocene in Europe (MIS 11, Germany)

High-resolution paleoecological proxies, such as stable isotopes or tooth microwear in large mammals, are often used for their potential to deliver information about the paleodietary traits of individuals and populations at the time of death. Such proxies are of interest in high resolution sites because they provide accurate data regarding the diet of large herbivores as well as the habitats that were available at the time of formation of the site, and by inference can detect seasonality in the formation of the assemblages. The integration of two techniques, tooth mesowear and microwear, applied to Middle Pleistocene assemblages of large herbivores from Steinheim and Heppenloch did not indicate seasonality at any of the two sites, most likely due to low resolution and time averaging of the dietary signal. However, the combination of the two proxies was highly informative for reconstructing the paleodiets of the large herbivores. The two paleodietary proxies provided consistent results that permitted us to propose a reconstruction of the paleodiets and habitats available at each site.

Chronological and Isotopic data support a revision for the timing of cave bear extinction in Mediterranean Europe

The Cave Bear, Ursus spelaeus (sensu lato), was one of many megafaunal species that became extinct during the Late Pleistocene in Europe. With new data we revisit the debate about the extinction and paleoecology of this species by presenting new chronometric, isotopic and taphonomic evidence from two Palaeolithic cave bear sites in northeastern Italy: Paina Cave and Trene Cave. Two direct radiocarbon dates on well-preserved collagen have yielded ages around 24,200–23,500 cal yr BP, which make them the latest known representatives of the species in Europe. The carbon (δ13C) and nitrogen (δ15N) isotopic values of bone collagen exhibit values similar to those of older cave bears from Swabian Jura and France, suggesting that the feedings preferences of cave bears remained unchanged until the disappearance of this species in Europe. Several bear remains preserved traces of human modification such as cut marks, which enables a reconstruction of the main steps of fur recovery and the butchering process. The broad range of plant types available and the favorable location of Berici Hills may have played an important role in the range expansion of cave bears and their interaction with the Paleolithic hunters settled the same area.

The diverse dietary profiles of MIS 3 cave bears from the Romanian Carpathians: insights from stable isotope (δ13C and δ15N) analysis

AbstractLate Pleistocene European cave bears (Ursus spelaeus) have been considered to be largely vegetarian, although stable isotope data (δ13C and δ15N values) from the Romanian Carpathians has suggested considerable dietary variation. Here we evaluate previous and additional adult cave bear isotopic data from four Marine Isotope Stage 3 (MIS 3) sites in the Carpathians. Peştera Urşilor (N = 35), Peştera Cioclovina (N = 32), Peştera Muierilor (N = 8), and Peştera cu Oase (N = 72) provide both a dichotomy between samples suggesting vegetarian diets (from Cioclovina and Muierilor) and more omnivorous diets (from Urşilor and Oase), and considerable isotopic variation within samples from each site. While an inference of a strictly vegetarian diet may apply to groups that lived in ecosystems which restricted the available animal protein for these large ursids, the within and between sample isotopic variation among the Carpathian cave bears indicates considerable flexibility in their sources of protein and hence in their dietary regimes. In addition, developmental assessment of Cioclovina isotopic profiles (neonates, juveniles, sub‐adults and adults) provides patterns of transfer of stable isotope signatures throughout immature life for both δ13C and δ15N (increase and decrease, respectively), whereas those from Urşilor show little developmental shift.

Hominin and carnivore interactions during the Early Pleistocene in Western Europe

The first peopling of Europe has been widely discussed for the last decades. The many findings recorded in recent years have confirmed that Europe was occupied by humans during the Early Pleistocene for over a million years. However, several issues are still questioned in the current debate about the first peopling of Europe, including the continuity or discontinuity of this event. In this regard, a revision of the available zooarchaeological evidence for the Early Pleistocene in Europe is proposed in this article, discussing the influence on hominin behaviour of meat-resource acquisition. The climatic fluctuations which characterized this period, as well as the diversity of ecosystems found in the Mediterranean area and in the whole continent, make meat consumption a key resource concerning the adaptive possibilities of local hominins. Thus, the persistence of hominin settlement in Europe during the Early Pleistocene may have depended on the social cohesion of the groups and their capacity to provide a regular supply of meat resources.

Evolutionary History of Saber-Toothed Cats Based on Ancient Mitogenomics

Saber-toothed cats (Machairodontinae) are among the most widely recognized representatives of the now largely extinct Pleistocene megafauna. However, many aspects of their ecology, evolution, and extinction remain uncertain. Although ancient-DNA studies have led to huge advances in our knowledge of these aspects of many other megafauna species (e.g., mammoths and cave bears), relatively few ancient-DNA studies have focused on saber-toothed cats [1-3], and they have been restricted to short fragments of mitochondrial DNA. Here we investigate the evolutionary history of two lineages of saber-toothed cats (Smilodon and Homotherium) in relation to living carnivores and find that the Machairodontinae form a well-supported clade that is distinct from all living felids. We present partial mitochondrial genomes from one S.populator sample and three Homotherium sp. samples, including the only Late Pleistocene Homotherium sample from Eurasia [4]. We confirm the identification of the unique Late Pleistocene European fossil through ancient-DNA analyses, thus strengthening the evidence that Homotherium occurred in Europe over 200,000 years later than previously believed. This in turn forces are-evaluation of its demography and extinction dynamics. Within the Machairodontinae, we find a deep divergence between Smilodon and Homotherium (∼18 million years) but limited diversity between the American and European Homotherium specimens. The genetic data support the hypothesis thatall Late Pleistocene (or post-Villafrancian) Homotherium should be considered a single species, H.latidens, which was previously proposed based on morphological data [5, 6].

Revisiting Ursus etruscus (Carnivora, Mammalia) from the Early Pleistocene of Greece with description of new material

The Villafranchian ursids of Greece are scanty, represented by a few isolated teeth and bones. During our last field campaigns in the Mygdonia Basin (Macedonia, Greece) we discovered an almost complete cranium, as well as some cranial, dental and postcranial remains, which are presented in this article. The new material originates from the locality Tsiotra Vryssi (TSR), dated to the late Villafranchian (1.8–1.2 Ma). The specimens are described and compared with ursids from various European localities; their morphological characters and dimensions place it with Ursus etruscus Cuvier, 1823, while the dental features are most similar to the specimens from Pietrafitta (Italy). The TSR cranium was CT-scanned in order to create a 3D virtual model and study its internal anatomy. Its observed endocranial traits exhibit primitive conditions, confirming the basal position of U. etruscus among the Pleistocene European ursids. Most of these features are shared with U. arctos and clearly differ from U. deningeri and U. spelaeus. Finally, we provide a revision of the known Greek material of U. etruscus and the biostratigraphic distribution of the Miocene–Pleistocene ursids of Greece. The earliest secure appearance of Ursus etruscus in Greece is found in the locality Dafnero 1 (Western Macedonia), dated to the end of the middle Villafranchian at ∼2.0 Ma, whereas its probable last occurrence is known from the locality Apollonia 1 of the Mygdonia Basin, dated to the latest Villafranchian at ∼1.2 Ma.

Small mammal fauna in Europe during the second half of the Middle Pleistocene

Evolutionary changes in European small mammals during the second half of the Middle Pleistocene, from the Likhvin (Holsteinian, Hoxnian) Interglacial (MIS 11) to the beginning of the Mikulino (Eemian) Interglacial (MIS 5e), that is between 424 ka BP and 130 ka BP were traced. Trends in evolutionary change were documented, and East European and West European faunas were compared. An integrated analysis of available theriological, geological, and geochronological data for the second half of the Middle Pleistocene in Europe has shown marked changes in the small mammal fauna throughout the period under consideration and provided information on the climate and environments at different time intervals. Changes traceable in the Arvicolinae phyletic lines made a correlation between the West European and East European mammal localities possible. The biostratigraphic scheme of the second half of the Middle Pleistocene has been developed and maps of small mammal localities compiled.

Evidence of Neanderthals in the Balkans: The infant radius from Kozarnika Cave (Bulgaria)

Excavations conducted by a Bulgarian-French team at Kozarnika Cave (Balkans, Bulgaria) during several seasons yielded a long Paleolithic archaeological sequence and led to the discovery of important faunal, lithic, and human samples. This paper aims to describe the unpublished radius shaft of an infant who died approximately before the sixth month postnatal that was recovered from layer 10b, which contained East Balkan Levallois Mousterian with bifacial leaf points. The layer was dated between 130 and 200 ka (large mammals biochronology) and between 128 ± 13 ka and 183 ± 14 ka (OSL), i.e. OIS6. Here we show that, given the scarcity of Middle Pleistocene infant remains in general, and Middle Paleolithic human remains from this part of Eastern Europe in particular, the study of the Kozarnika specimen is of special interest. We discuss its place in the Middle Pleistocene European hominine record and substantiate the hypothesis of early Neanderthal presence in the eastern Balkans.

New luminescence-based geochronology framing the last two glacial cycles at the southern limit of European Pleistocene loess in Stalać (Serbia)

Abstract A new geochronology was established for the Stalać loess-paleosol sequence (LPS) in Serbia. The section is located in the interior of the Central Balkan region, south of the typical loess distribution, in a zone of paleoclimatic shifts between continental and Mediterranean climate regimes. The sampled sequence contains four well-developed paleosol and loess layers, a crypto tephra and one visible tephra layer. Optically stimulated luminescence measurements showed a strong dependency of preheat temperature on equivalent dose for one fine-grained quartz sample, which makes it unsuitable for dating. A firm chronology framing the last two glacial cycles was established using fine-grained polyminerals and the post-infrared infrared stimulated luminescence (pIR50IR290) protocol instead. The characteristics of dated paleosols indicate similar climatic conditions during the last interstadial and interglacial phases, which were different from the penultimate interglacial period. The tephra within the L2 loess, probably related to tephra layers also found in other sections in Southeastern Europe, was sandwich-dated. The results indicate an age between 118 ka and 141 ka. Furthermore, a weak pedogenic layer dated to between 126 ka and 148 ka gives a first numerical age to this soil formation in Southeastern Europe.

Non‐traditional isotope perspectives in vertebrate palaeobiology

AbstractThe recent development of multi‐collector inductively coupled plasma mass spectrometry (MC‐ICP‐MS) notably in the disciplines of earth sciences, now allows the precise measurement of isotope ratios, even at low concentration. Non‐traditional isotope systems, such as alkaline earth (Ca, Mg) and transition (Cu, Fe, Zn) metals are now being measured in a variety of biological tissues, including bone and teeth. Although our understanding of the environmental and biological mechanisms behind the fractionation of such elements is still in its infancy, some of these isotopes are suspected to fractionate along the food chain, as has been reported in the literature for calcium, magnesium and zinc. Other geochemical methods, such as concentration analyses, permit a prior assessment of diagenesis in the fossils and such an approach indicates that in some circumstances, not only enamel but also dentine or bone can preserve its original biogenic composition. The aims here are to: review the current knowledge surrounding these various isotopic tools; address their potential preservation in biological apatite; and provide the palaeobiologist with a guide to the different toolkits available, including a discussion of their potential applications in vertebrate palaeobiology with a case study involving two mammal assemblages from the Pleistocene of Europe.

Disentangling relationships among the members of the Silene saxífraga alliance (Caryophyllaceae): Phylogenetic structure is geographically rather than taxonomically segregated

AbstractOut of ca. 700 species belonging to Silene, 15 to 35 were classified in S. sect. Saxifragoideae. This section has its highest diversity on the Balkan Peninsula, which was the most important European Pleistocene refugium. The most widespread of the species is S. saxifraga ranging from the Pyrenees to the Carpathians. Together with ten morphologically similar taxa it is often treated as the S. saxifraga group, but the relationships among the taxa are unclear and blurred due to the occurrence of transitional forms. In this study, we used sequences of the nuclear ribosomal internal transcribed spacer (ITS) and the plastid rps16 intron as well as amplified fragment length polymorphisms (AFLPs) to elucidate phylogenetic relationships among the taxa belonging to—or hypothesized to be closely related to—S. sect. Saxifragoideae, covering the section’s entire geographical range. Phylogenetic analyses of 112 populations belonging to 33 taxa clarified previous classifications and showed that S. cephallenia, S. congesta, S. linoides, S. multicaulis, S. schwarzenbergeri and S. waldsteinii are clearly differentiated from all other taxa, which in turn form the newly circumscribed S. saxifraga group. Phylogenetic relationships within the latter are unresolved, and are governed by geographic proximity rather than by taxonomic identity. This is likely a result of recent rapid radiation and range expansion, as well as of convergent morphological evolution triggered by similar environmental selection pressures. The southern parts of the Balkan Peninsula are the centre of genetic diversity of the group, underlining the area’s importance as a sanctuary of diversification of European biota.

Early Pleistocene hominin deciduous teeth from the Homo antecessor Gran Dolina-TD6 bearing level (Sierra de Atapuerca, Spain).

During the last 13years, the late Early Pleistocene Gran Dolina-TD6-2 level (Sierra de Atapuerca, northern Spain) has yielded an additional sample of 26 dental specimens attributed to Homo antecessor. In this report, we present a descriptive and comparative study of the six deciduous teeth. We provide external and internal morphological descriptions following classical terminology, as well as the mesiodistal and buccolingual measurements of the teeth. The internal morphology was described by means of micro-CT technique. The TD6 deciduous teeth preserve primitive features regarding the Homo clade, such as the presence of styles in lower and upper canines and developed anterior and posterior foveae in the dm2 . However, other features related to the complexity of the crown morphology (e.g., cingulum) are not present in this sample. Furthermore, the great reduction of the talonid of the dm1 s is also noteworthy. Despite the limited comparative evidence, the presence of a remarkably well-developed tuberculum molare in the dm1 and dm1 s from TD6 can be also considered a derived feature in the genus Homo. The TD6 hominins exhibit dental dimensions similar to those of other Pleistocene hominins. The dm1 s are buccolingually elongated and the buccolingual diameter of ATD6-93 is the largest recorded so far in the Homo fossil record. This study expands the list of plesiomorphic features of H. antecessor, and provides some information on the evolutionary status of this species. However, the identification of some advanced traits evinces a step towards the derived morphology of European Pleistocene teeth. The study of the deciduous dentition confirms the mosaic pattern of H. antecessor morphology revealed in previous studies of this hominin sample.

Morphometric-palaeoecological discrimination between Bison populations of the western Palaearctic

The present study outlines a comprehensive chrono-spatial framework of the palaeoecological adaptations among several western Palaearctic Bison populations. Possibly arrived during the middle early Pleistocene (late Villafranchian), bisons were present with various morphotypes for more than 1.5 million years in the European mammalian palaeocommunities. Based on dental wear and postcranial morphological traits, bisons can be attributed to distinct size classes and shape trends, associated with distinct feeding and locomotor adaptations across regional provinces and time intervals. During the progressively cooler and more arid climate conditions towards the late Pleistocene, a body mass increase and a significant distal limb elongation are recorded. After the last glacial climatic deterioration, the restoration of a more temperate climate and the expansion of woodlands triggered the general size decrease and limb shortening towards the beginning of the Holocene, both reflected in extant Holarctic bisons. The phenotypic diversity, imprinted in further limb structure alterations, was driven by climatic instability and landscape vegetational heterogeneity. The morphological variability among early Pleistocene forms can be attributed to evolutionary processes or most likely multiple Bison invasions at that time. The vast majority of Pleistocene European bisons primarily inhabited relatively open and dry environments, suited for grazing though they were able to exploid a variety of open and fairly wooded habitats, consuming a broad vegetational spectrum. In contrast, the increasing arid character of the southeastern part of Epivillafranchian Europe resulted in periodically harsh substrate and a more grazing specialized feeding mode.