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Sus arvernensis is a Pliocene species that occupies a key position in the evolution of suids (Suidae, Artiodactyla, Mammalia) in Eurasia, and besides, it is considered important for biochronological correlations and paleoecological inferences. However, our knowledge on S. arvernensis is largely based on fossil remains from southwestern Europe. Here, we present a revision of the Hungarian fossil record of S. arvernensis. Up to now, the species was known from only two localities of Hungary, Gödöllő (central Hungary) and Süttő (northwest Hungary), and the latter occurrence has even been questioned. After the comparison with other relevant samples of S. arvernensis, of the Early Pleistocene S. strozzii, and of the extant wild boar S. scrofa (motivated by previous attributions and the chronology of the localities), the presence of S. arvernensis from Gödöllő and Süttő is confirmed, and more material of the species is described from Beremend (southern Hungary) and Kisláng (western Hungary). Collectively, the results of the revision carried out herein reveal a relatively widespread distribution of S. arvernensis in Hungary, hence providing an important link from the eastern to western European fossil record of the species. The specimens from Gödöllő and Süttő are slightly larger than the other material of S. arvernensis from France and Italy included in the biometric comparison, although the paucity of the material precludes to evaluate whether these differences are significant and to relate them to a chronological and/or geographical context. The occurrence of S. arvernensis in the Hungarian localities considered in this work is a biochronological indication of an age older than at least 2.6 Ma, since the species is not recorded after the Pliocene–Pleistocene transition. This in agreement with the age estimates available so far for some of the localities or provides new insights. At Süttő, in particular, the identification of S. arvernensis reinforces the view that travertine deposition started already in the Pliocene. [ABSTRACT FROM AUTHOR]

Located at the eastern corner of Mediterranean Europe, Greece occupies a critical position for mammal dispersals to/from Europe, Asia, and Africa and constitutes a potential passageway towards Western Europe. During recent decades, numerous fieldwork campaigns in several Pliocene–Pleistocene sites have greatly enriched the fossil record and provided valuable taxonomic and biostratigraphic data. However, a fully developed reference biochronological unit scheme for the Greek record that could contribute to correlations at a continental scale is still pending. In this article, we provide the updated Late Pliocene to Middle Pleistocene large mammal succession, and we introduce the Faunal Units (FUs) of Greece. We define eight FUs, the Milia, Dafnero, Gerakarou, Tsiotra Vryssi, Krimni, Apollonia, Marathousa, and Apidima FUs (from the oldest to the youngest), which are determined by a set of first and last local occurrences. The results form the basis for discussion of already set turnovers, dispersals, and extinction/immigration events and showcase the importance of the local record for the investigation of the European terrestrial ecosystems. By developing the first detailed biochronological scheme for the Pliocene–Pleistocene of Southeastern Europe, this study comprises the basis for an expanded Balkan faunal unit scale and a reference framework for future investigations. [ABSTRACT FROM AUTHOR]

Located at the eastern corner of Mediterranean Europe, Greece occupies a critical position for mammal dispersals to/from Europe, Asia, and Africa and constitutes a potential passageway towards Western Europe. During recent decades, numerous fieldwork campaigns in several Pliocene–Pleistocene sites have greatly enriched the fossil record and provided valuable taxonomic and biostratigraphic data. However, a fully developed reference biochronological unit scheme for the Greek record that could contribute to correlations at a continental scale is still pending. In this article, we provide the updated Late Pliocene to Middle Pleistocene large mammal succession, and we introduce the Faunal Units (FUs) of Greece. We define eight FUs, the Milia, Dafnero, Gerakarou, Tsiotra Vryssi, Krimni, Apollonia, Marathousa, and Apidima FUs (from the oldest to the youngest), which are determined by a set of first and last local occurrences. The results form the basis for discussion of already set turnovers, dispersals, and extinction/immigration events and showcase the importance of the local record for the investigation of the European terrestrial ecosystems. By developing the first detailed biochronological scheme for the Pliocene–Pleistocene of Southeastern Europe, this study comprises the basis for an expanded Balkan faunal unit scale and a reference framework for future investigations.

The Grăunceanu site in the Olteţ River Valley has yielded a rich carnivoran assemblage including at least 10, possibly 11 species: Vulpes alopecoides, Nyctereutes megamastoides, Ursus etruscus, Meles thorali, Lutraeximia sp., Pliocrocuta perrieri, Lynx issiodorensis, Puma pardoides, Megantereon cultridens, Homotherium latidens and possibly Pachycrocuta brevirostris. The faunal assemblage is compared with approximately coeval sites Europe and western Asia. This comparative analysis shows that the Grăunceanu assemblage shows the greatest similarities with sites to the west such as Senèze and, particularly, Saint-Vallier, rather than more easternly ones such as Liventsovka and Dmanisi. The relative abundance of the taxa at Grăunceanu was compared to that of Saint-Vallier and broad similarities were found, except for the absence of some cursorially adapted taxa present at Saint-Vallier but absent from Grăunceanu. The absence at Grăunceanu of taxa with a reconstructed cursorial hunting strategy, such as Chasmaporthetes and Acinonyx, may suggest less open habitat at Grăunceanu than indicated in previous paleoenvironmental reconstructions. [ABSTRACT FROM AUTHOR]

The Grăunceanu site in the Olteţ River Valley has yielded a rich carnivoran assemblage including at least 10, possibly 11 species: Vulpes alopecoides, Nyctereutes megamastoides, Ursus etruscus, Meles thorali, Lutraeximia sp., Pliocrocuta perrieri, Lynx issiodorensis, Puma pardoides, Megantereon cultridens, Homotherium latidens and possibly Pachycrocuta brevirostris. The faunal assemblage is compared with approximately coeval sites Europe and western Asia. This comparative analysis shows that the Grăunceanu assemblage shows the greatest similarities with sites to the west such as Senèze and, particularly, Saint-Vallier, rather than more easternly ones such as Liventsovka and Dmanisi. The relative abundance of the taxa at Grăunceanu was compared to that of Saint-Vallier and broad similarities were found, except for the absence of some cursorially adapted taxa present at Saint-Vallier but absent from Grăunceanu. The absence at Grăunceanu of taxa with a reconstructed cursorial hunting strategy, such as Chasmaporthetes and Acinonyx, may suggest less open habitat at Grăunceanu than indicated in previous paleoenvironmental reconstructions.

Karnezeika is an Early Pleistocene locality in the Peloponnese, South Greece. More than 3000 specimens have been unearthed and studied, revealing the presence of a large mammal assemblage, which corresponds to the middle Villafranchian biochronological unit. The bovid fauna of the site has already been presented, as well as the scarce remains of a large cercopithecid, which characterizes Karnezeika as a primate-bearing fossil site. Herein, we describe the Carnivora of this locality. The carnivore guild of Karnezeika was found to consist of seven different taxa including Vulpes alopecoides , Felis sp., Megantereon cultridens , Ursus etruscus , Pachycrocuta brevirostris , Baranogale helbingi , and a Mustelidae indet. Interestingly, the presence of the giant, short-faced hyena P. brevirostris among the assemblage shows that this species was present in SE Europe before the Olduvai subchron, similarly to Western Europe. Furthermore, based on the new and relatively well-preserved specimens belonging to the rare mustelid B. helbingi , we discuss some possible taxonomic interpretations of this species. Finally, the majority of the carnivora taxa present in Karnezeika indicate a mixed habitat which agrees with previous palaeoenvironmental results concerning the site. [ABSTRACT FROM AUTHOR]

More than 3000 fossil vertebrate specimens have been unearthed from the new Lower Pleistocene locality of Karnezeika, Peloponnese, Greece. The vast majority of the material (~90%) represents bovids, and their systematic study is provided herein. The bovid assemblage consists of four taxa. A significant portion of the collected bovid material belongs to a small sized gazelle, attributed to Gazella bouvrainae, and to the large Rupicaprini Gallogoral meneghinii. A relatively small sized form of Gazellospira torticornis has been recorded as well, and in addition the scarce dental material belonging to an unspecified Caprini form is mentioned as Caprini gen. et sp. indet. This combination of taxa is typical for the Middle Villafranchian of Greece (2.6–1.8 Ma). [ABSTRACT FROM AUTHOR]

Some remains of suids were recovered during the second half of the 1800s from Quercia (Tuscany, Italy), at a close distance to, but from a lower stratigraphic position than the better-known mammal fauna of Olivola. Although they were collected a long time ago, Quercia suids are described for the first time in this work. This sample represents one of the earliest occurrences (middle Villafranchian) of Sus strozzii , a large-sized suid present in Europe during the Early Pleistocene, but only abundant ∼2.0–1.8 Ma. A biometric comparison with selected samples of Pliocene to early Middle Pleistocene European suids is carried out, showing some dimensional changes in S. strozzii as well as differences between S. strozzii and other species. Quercia has been somehow eclipsed by other historical samples from Tuscany, namely the extensive collection of the Upper Valdarno and the diverse fauna of Olivola, but it is a different and important palaeontological locality. Apart from S. strozzii , the local fauna of Quercia-Vaccareccia includes Anancus arvernensis , Canis sp., Stephanorhinus etruscus , Leptobos etruscus , ' Pseudodama ' sp., and Castor sp., representing one of the few mammal assemblages referable to the Faunal Unit of Coste San Giacomo (late middle Villafranchian, MNQ 17b), corresponding to ∼2.2–2.1 Ma. [ABSTRACT FROM AUTHOR]

The dispersal of primitive elephantines and monodactyl equids in Eurasia has long been regarded as representative of a substantial turnover in mammal faunas, denoting the spread of open environments linked to the onset of cold and dry conditions in the Northern Hemisphere. During the 1980s, this event was named the "Elephant-Equus event" and it was correlated with the Gauss-Matuyama reversal, today corresponding to the Pliocene-Pleistocene transition and the beginning of the Quaternary, dated at ~2.6 Ma. Therefore, the Elephant-Equus event became a concept of prominent biochronological and paleoecological significance, especially in western Europe. Yet, uncertainties surrounding the taxonomy and chronology of early "elephant" and "Equus", as well as conceptual differences in adopting (or understanding) the Elephant-Equus event as an intercontinental dispersal event or as a stratigraphic datum, engendered ambiguity and debate. Here, we provide a succinct review of the Elephant-Equus event, considering separately the available evidence on the "elephant" and the "Equus". Elephantines dispersed out of Africa during the Pliocene (Piacenzian). Their earliest calibrated occurrences from eastern Europe date at ~3.2 Ma and they are usually referred to Mammuthus rumanus, although the allocation of several samples to this species is tentative. Available dating constraints for other localities do not resolve whether the dispersal of Mammuthus was synchronous across Eurasia, but this possibility cannot be ruled out. The spread of Mammuthus was part of an intercontinental faunal exchange between Africa and Eurasia that occurred during the Piacenzian, but in this scenario, Mammuthus is quite unique in being the only genus of African origin dispersing to western Eurasia. The arrival of monodactyl equids from North America coincides with the Pliocene-Pleistocene transition, with several occurrences dated or calibrated at ~2.6 Ma and no compelling evidence prior to this age. In Europe, early monodactyl equids are often aligned to Equus livenzovensis, but the material from the type locality of this species is chronologically time-averaged and taxonomically heterogeneous, and western European samples are seldom abundant or informative. Regardless, this does not diminish the biochronological significance of the "Equus event". Indeed, while the term "Elephant-Equus event" should no longer be used, as the appearance of elephantines in the European fossil record markedly precedes that of monodactyl equids, we endorse the use of the "Equus event" as a valid alternative to refer to the intercontinental dispersal event that characterizes the middle Villafranchian faunal turnover, epitomized by but not limited to monodactyl equids. [ABSTRACT FROM AUTHOR]

A new Lower Pleistocene vertebrate site, Krimni-3 (KMN) from Mygdonia Basin, Greece is originally presented herein, providing a preliminary analysis of its taphonomic context and faunal content. The KMN fossil mammal fauna includes the following: Erinaeus sp., Canis sp., Hyaenidae indet., Ursus sp., Equus altidens, Equus sp. (large-sized), Stephanorhinus cf. hundsheimensis, Sus sp., Palaeotragus inexpectatus, Cervidae indet. (medium-sized), Praemegaceros sp., Bison cf. degiulii, Soergelia brigittae, and Hemitragus sp. Additionally, an avian femur, fully described and compared, is attributed to the giant ostrich Pachystruthio dmanisensis. This is the first record of a giant ostrich in Greece and southeastern Europe in general, and by documenting the southernmost occurrence of the species, it expands significantly its known geographical distribution. Large-sized ungulates predominate in the assemblage, while a large number of postcranial remains appear articulated. Bone surfaces are mostly fresh or only slightly weathered, yet post-depositionally cracked. Minor to moderate rounding and polishing indicate the action of water to a certain degree, while several bone breakages and marks suggest carnivore activity (including carnivore gnawing on the ostrich femur). The comparison of the KMN fauna with those already known from Mygdonia Basin and Europe suggests a late Villafranchian age, intermediate between the faunas from Tsiotra Vryssi and Apollonia-1, within the 1.8–1.5 Ma time frame. This is a crucial interval of time for mammal dispersal and turnovers, and KMN opens another window into the Early Pleistocene ecosystems of western Eurasia. [ABSTRACT FROM AUTHOR]

The fossil record of lynxes provides clear evidence of a large range across the North Hemisphere during the Pliocene and Pleistocene. However, their origin, systematics and evolutionary relationships are still fraught with difficulties and controversy. Here we report a complete hemimandible of a medium-sized felid from the Early Pleistocene (MN17, middle Villafranchian, 2.05 Ma) site of La Puebla de Valverde (Teruel, Spain). Based on comparative and multivariate analyses of the lower dentition of 458 individuals of medium-sized Lynx, Caracal and Leptailurus, we confidently ascribe the remains to Lynx aff. issiodorensis. Although the dental proportions are somewhat different from those of the Eurasian L. issiodorensis (smaller canines and more elongated p4/m1), Lynx aff. issiodorensis shows affinities with the Issoire lynx from the contemporaneous site of Saint Vallier (France), sharing a similar morphology of the mandible, reduced canines, and long m1. We further test the hypothesis that examines the presence of the African/Asian Caracal in the European Plio/Pleistocene for C. depereti and C. issiodorensis, and discard the attribution of L. issiodorensis into Caracal. This mandible extends the record of the genus and contributes to update our understanding of the Lynx lineage and its variability within the European fossil record. [ABSTRACT FROM AUTHOR]

The climatic instability of the Early Pleistocene induced regional paleoenvironmental changes, which impacted faunal composition and dynamics. It coincides with the Homo dispersal out of Africa. The role played by the Eastern Balkans in this dispersal is at the heart of current scientific debates. The rich fossil assemblage of Gerakarou-1 (2–1.8 Ma; northern Greece) can provide important insight into the environmental conditions of that time and area. In this study, we provide insights into the dietary habits of the herbivore guild of Gerakarou-1 at both short and long-term scales based on two complementary dental proxies, mesowear and microwear textures applied on 70 specimens and we infer the local paleoecology and paleoenvironment. As diet varies between and among populations within the same species, we conduct a population-level study, including a reference dataset of 144 extant individuals belonging to six populations of ruminants with well-known ecology. As we aim to contribute to the collective effort of the community to improve the mesowear methodology, we propose and discuss here the benefits of an alternative, quantitative Geometric Morphometric approach. We show that the occlusal relief, as calculated in the classical mesowear approach, is a variable of little relevance and that the quantitative approach allows better discrimination between dietary categories. By proposing for the first time a multi-scale and multi-proxies study of the entire herbivore guild at Gerakarou-1, we nuance and clarify the paleoecology and niche-partitioning of the ungulates from this site and improve past paleoenvironmental interpretations. • Two complementary dental proxies offer a better overview of past ungulate ecology. • Gerakarou-1 ungulates are mixed-feeders to grazers. • Gerakarou-1 was a predominantly open landscape with little woody vegetation. • The occlusal relief used in the classical mesowear approach is of little relevance. • The Geometric Morphometrics mesowear is more discriminating than the classical one. [ABSTRACT FROM AUTHOR]

The present work provides a revision of the complete fossil collection of Equus stehlini from the Italian Early Pleistocene (late Villafranchian) of the Upper Valdarno Basin (Tuscany, Italy). The species was originally described in the second half of the last century, even if recent studies questioned its distinct species status, considering it a subspecies of Equus senezensis (i.e., E. senezensis stehlini). Over the succeeding decades the debate about the taxonomic status of E. stehlini extended to its possible evolutionary origin from two different Equus species from the European Early Pleistocene, Equus stenonis or E. senezensis. The revision undertaken here of E. stehlini, by means of morphological, morphometric and statistical comparisons, provides new insights into the taxonomic status and the evolutionary history of this enigmatic species. It is concluded that E. stehlini is a distinct species derived from the medium-sized E. senezensis. This work further highlights new insights and perspectives in the evolutionary framework and in the paleoecological understanding of the medium to small-sized Equus species from the early to middle Early Pleistocene of Europe. [ABSTRACT FROM AUTHOR]

The present work provides a revision of the complete fossil collection of Equus stehlini from the Italian Early Pleistocene (late Villafranchian) of the Upper Valdarno Basin (Tuscany, Italy). The species was originally described in the second half of the last century, even if recent studies questioned its distinct species status, considering it a subspecies of Equus senezensis (i.e., E. senezensis stehlini). Over the succeeding decades the debate about the taxonomic status of E. stehlini extended to its possible evolutionary origin from two different Equus species from the European Early Pleistocene, Equus stenonis or E. senezensis. The revision undertaken here of E. stehlini, by means of morphological, morphometric and statistical comparisons, provides new insights into the taxonomic status and the evolutionary history of this enigmatic species. It is concluded that E. stehlini is a distinct species derived from the medium-sized E. senezensis. This work further highlights new insights and perspectives in the evolutionary framework and in the paleoecological understanding of the medium to small-sized Equus species from the early to middle Early Pleistocene of Europe.

The Early Pleistocene site of Pietrafitta (central Italy) produced a rich vertebrate assemblage from the Late Villafranchian Land Mammal Age (late MN18). Geological and paleobotanical data from Pietrafitta indicate a lacustrine environment, surrounded by a humid deciduous broadleaved forest with a temperate climate. The vertebrate assemblage consists of at least 40 taxa including actinopterygians, amphibians, reptiles, birds, and mammals. Here, we concentrate on the ichthyofaunal and herpetofaunal remains. The ichthyofauna includes Barbus Cuvier and Cloquet, 1816, Scardinius Bonaparte, 1837, aff. Squalius Cuvier, 1817, and Tinca Cuvier, 1817. The two anuran genera are the large-sized alytid frog Latonia Meyer, 1843 and the "green frog" Pelophylax Fitzinger, 1843. Three snake precloacal vertebrae were recognized, one attributed to Colubrines indet., another one to Natrix sp. Laurenti, 1768, and the largest and most complete vertebra is referred to the genus Vipera s.l. Laurenti, 1768 (cf. gr. "Oriental vipers"). The chelonian fossils, including some complete carapaces and plastrals, are attributed to the European pond turtle (Emys gr. Orbicularis Linnaeus, 1758) and Hermann's tortoise (Testudo hermanni Gmelin, 1789). [ABSTRACT FROM AUTHOR]

The evolution of large mammal faunas during the Pleistocene of Europe has been widely investigated using taxonomical and/or ecological-functional categories, with special emphasis on the implications for reconstructing hominin dispersal and ecological role. Here, an ecospace modelling approach is for the first time applied to Pleistocene carnivorans of Europe. Examining ecospace occupancy and disparity through time effectively conveys major aspects of faunal dynamics, while offering new insights into species relationships from a trait-space-explicit perspective. Several replacements occurred within-family and concerned species of similar functional role, but between-family competition was also important, especially for hyaenids. When hominins first dispersed into Europe during the Early Pleistocene, carnivorans of middle size that are deemed either highly social active hunters or primarily scavengers were both lacking, a substantial ecospace vacuum that could have been filled by the newcomers. Hominins then became widespread since the Middle Pleistocene, when a decrease of ecospace density is documented. This suggests that adaptability, mainly behavioural for hominins, ecomorphological for other species, might have been key for survival, especially in the more unstable conditions encountered since the latest Early Pleistocene. The uncertainty in categorising the ecology of extinct species is a limitation shared with previous studies, but reducing this multidimensional problem to one ecospace facilitates addressing the main pattern and future assessment. • Ecospace modelling is for the first time applied to Pleistocene carnivorans of Europe. • New insights into species relationships are provided from a trait-space-explicit perspective. • Within-family replacements of species of similar functional role were common. • Hominins may have initially occupied the vacant space of middle-sized and highly social carnivorans. • A Middle Pleistocene drop in ecospace density indicates the key role of adaptability for survival. [ABSTRACT FROM AUTHOR]

The Villafranchian fossil record of Perissodactyla is of a great interest because the appearance of the genus Equus and disappearance of the genus Tapirus mark important faunal turnovers. Here, we provide new data on Italian Villafranchian Tapiridae, Rhinocerotidae, and Equidae with updates from the last comprehensive review; most relevant are those of the Rhinocerotidae and Equidae. At present only two Villafranchian rhinoceros' species are documented in Italy: Stephanorhinus jeanvireti and Stephanorhinus etruscus. The two species can be distinguished by several morphological features and by the dimensions of their bones. Stephanorhinus etruscus occurred throughout the Villafranchian; despite the numerous records, its remains are not enough to investigate morphometric trends. The Equidae include seven species including new occurrences of Hipparionine horses and of the genus Equus. The statistical analyses performed on the Equus species show two different evolutionary trends, namely the stenonine lineage " Equus livenzovensis - Equus stenonis " and the small-to-medium-sized lineage " Equus senenzensis - Equus stehlini ", suggesting a different evolution of these species. Moreover, the analyses show close relationships between the Italian Equus stenonis samples and three European Equus stenonis subspecies (E. stenonis vireti, E. stenonis guthi , and E. stenonis pueblensis). We begin to reconsider the validity of E. stenonis subspecies identification in order to provide new perspectives on the taxonomy of this species. [ABSTRACT FROM AUTHOR]

After more than sixty years since its discovery, the fossils from Monte Peglia (late early Biharian, Umbria) are reconsidered in their entirety. The small mammals of Monte Peglia upper and lower levels had been studied in the past, whereas the remains of large mammals of Monte Peglia lower level, stored in several Institutions of central Italy, are described here for the first time. The following taxa have been described: Hystrix refossa, Homotherium latidens, Panthera cf. P. gombaszoegensis, Felis cf. F. lunensis, Canis mosbachensis, Vulpes alopecoides, Meles meles, Pannonictis cf. P. nestii, Mustela palerminea, Ursus cf. U. etruscus; Macaca sylvanus florentinus, Equus altidens, Stephanorhinus cf. S. hundsheimensis, Sus sp., Capreolus sp., Axis eurygonos, Hemitragus cf. H. orientalis, Bison degiulii. Moreover, the list of small mammals of the lower level has been updated with the addition of three new small vertebrate taxa: Rana sp., Myotis sp. (large size), cf. Miniopterus sp. The study of the remains of large mammals of the lower layer indicate the survival of a number of taxa of latest Villafranchian age. If we accept the biochronological correlation of Monte Peglia with the Colle Curti local fauna, its age should be ~1.072 Ma. In this case, it is possible to pinpoint the accumulation of the lower level to the MIS 35/33, as the small mammals confirm the presence of a mixed environment with forested and open spaces and warm temperate climate. The accumulation of the upper level, characterized by taxa typical of open spaces and steppes and a cooler climate, probably occurred during MIS 34/32.

Palynological investigations were carried out on the basal sediments of the Aulla-Olivola basin, precisely on the Aulla lacustrine deposit, which underlies by about 200 m, both stratigraphically and geometricall", the Olivola village, known for its famous Villafranchian fauna. This research has revealed; • a very rich palynoflora and a long, complex, vegetational succession. dominated by arboreal formations, with high abundance of exotic taxa, including also tropical-subtropical elements; • a climatic Sequence characterized by oscillations reflecting seasonality, especially in pluvial regime. A most evident climatic change occurs in the middle of the sequence. Moreover, the palynological sequence made it possible: • to assign the examined sequence to the Pliocene, because of the abundance of "Tertiary" elements (Taxodium-type, Sciadopitys, Sequoia-type, Nyssa, Palmae, etc.); • to-ascertain the rather archaic character of the Aulla pollenflora because of the significant occurrence Of pollen assemblages mainly confined to pre-Pliocene and early Pliocene (Symplocos, Sapotaceae, Clethraceae/Cyrillaceae-type, Magnolia-type, Sapindaceae- type, Sterculiaceae, cfr. Castanopsis, Tricolporopollenites sp. of archaic-type, etc.); • to compare the Aulla pollenflora with Other Pliocene pollen microfloras from Italy and Other areas and to evidence affinities and differences. From these comparisons it appears clearly that the Aulla lacustrine episode represents the most ancient continental Pliocene deposit discovered in Italy by means of pollen analyses so far. In fact, the palynological and climatic features indicate that the pollen sequence of Aulla is certainly older than the establishement of glacial/interglacial climatic cycles in the Northern Hemisphere (—2,5 Ma). Also the Villafranca d'Asti deposit (Early Villafranchian in age) is older than 2,5 Ma on the base of the paleomagnetic data. But the pollen floras of Villafranca d'Asti (Fornace R.D.B.) do not include the pre- Pliocene and early Pliocene elements, which characterize the Aulla pollen records. All these considerations lead to Suggest an older chronostratigraphic collocation of the Aulla sequence, most likely in a pre-Vilafranchian interval, i.e., in the Ruscinian stage interval. The large floristic affinities with the pollen zones of the Brunssumian of the Netherlands, entirely included in the Ruscinian stage, and also the pollen affinities with the continental middle-Pliocene western-alpine deposits support the chronostratigraphical assignation of the Aulla deposit to a (probably late) Ruscinian phase. Finally, the pollen record of Aulla represents a particular uniform floristic biozone, which is named here "Macrian" phase, easily distinguished from the contiguous vegetational biozones. The results reported above allow to extend the chronostratigraphic scheme of the paleofloristic evolution in Northern-Central Italy during the Pliocene.

After more than sixty years since its discovery, the fossils from Monte Peglia (late early Biharian, Umbria) are reconsidered in their entirety. The small mammals of Monte Peglia upper and lower levels had been studied in the past, whereas the remains of large mammals of Monte Peglia lower level, stored in several Institutions of central Italy, are described here for the first time. The following taxa have been described: Hystrix refossa, Homotherium latidens, Panthera cf. P. gombaszoegensis, Felis cf. F. lunensis, Canis mosbachensis, Vulpes alopecoides, Meles meles, Pannonictis cf. P. nestii, Mustela palerminea, Ursus cf. U. etruscus; Macaca sylvanus florentinus, Equus altidens, Stephanorhinus cf. S. hundsheimensis, Sus sp., Capreolus sp., Axis eurygonos, Hemitragus cf. H. orientalis, Bison degiulii. Moreover, the list of small mammals of the lower level has been updated with the addition of three new small vertebrate taxa: Rana sp., Myotis sp. (large size), cf. Miniopterus sp. The study of the remains of large mammals of the lower layer indicate the survival of a number of taxa of latest Villafranchian age. If we accept the biochronological correlation of Monte Peglia with the Colle Curti local fauna, its age should be ~1.072 Ma. In this case, it is possible to pinpoint the accumulation of the lower level to the MIS 35/33, as the small mammals confirm the presence of a mixed environment with forested and open spaces and warm temperate climate. The accumulation of the upper level, characterized by taxa typical of open spaces and steppes and a cooler climate, probably occurred during MIS 34/32. [ABSTRACT FROM AUTHOR]

The Early Pleistocene of Eurasia is marked by significant climatic, environmental, and faunal shifts and is the time during which Homo first appears in the Eurasian fossil record. To better characterize the environments that were available to these hominins, accurate data regarding the faunal composition of eastern European sites are necessary, as this is the region through which Homo is hypothesized to have dispersed into Europe. Here we present updated and revised taxonomy and biochronology for sites in the Olteţ River Valley (ORV) of central Romania. The site of Grăunceanu is one of the most fossiliferous deposits from this time. Excavations and descriptions of the ORV sites took place during the 1960–1980s, but since that time many taxonomic revisions have been published. Here, we present a reassessment and update of the fauna from the ORV sites. We report several new taxa (e.g., Pachystruthio , Smutsia) and expand the known biogeographic range for other taxa (e.g., Puma pardoides). Our biochronological assessment of this updated taxonomy is consistent with previous reports, though with some refinement. We suggest Grăunceanu is Late Villafranchian (~2.2–1.9 Ma) and compositionally similar to the sites of Saint-Vallier (France) and Vatera (Greece). Similarly, the La Pietriş assemblage was likely deposited >1.7 Ma. The younger site of Fântâna lui Mitilan was deposited after 1.8 Ma, and perhaps as recently as 1.1 Ma. Thus, the ORV faunal assemblages provide an excellent record of nearly the entire span of the Late Villafranchian for both central Romania and eastern Europe. [ABSTRACT FROM AUTHOR]

Herein we describe a fragment of a mandible with a deciduous fourth premolar (dp4) from the Early Pleistocene locality of Dunaalmás, representing the first confirmed report of Sus strozzii from Hungary. The comparison of dp4 measurements supports a statistically significant distinction between S. strozzii and Sus scrofa. The two species overlap in time during the late Early Pleistocene of Europe (Epivillafranchian), but suid remains of this time-span are seldom classified at a species level. The correct taxonomic identification of the Epivillafranchian suids, which are often associated with evidences of hominin presence, is of great palaeoenvironmental value because S. scrofa and S. strozzii possess different ecological requirements. [ABSTRACT FROM AUTHOR]

The extinct bovid Leptobos is one of the most characteristic elements of Eurasian faunal assemblages during most of the Villafranchian Land Mammal Age (i.e., from the late Pliocene to most of the early Pleistocene). Several species of this genus have been established since the end of XIX Century, but their taxonomic status and phylogenetic relationships remain unclear due to the fact that most of them are described on the basis of scanty material. European species are divided into two groups or lineages. The first includes L. stenometopon, L. merlai, and the poorly known L. furtivus, the second L. etruscus and L. vallisarni. While the last two species are well documented in the Italian early Pleistocene fossil record, very little is known on the L. stenometopon-merlai-furtivus group and especially on L. merlai, whose richest sample is from the French locality of Saint Vallier. Here, we describe new material of L. merlai from the early late Villafranchian of Umbria (central Italy), including a nearly complete female cranium and a male neurocranium with horn cores. These remains constitute the best-preserved and complete sample of L. merlai in the Italian Peninsula and bring new consistency to the fossil record of this species. In addition, they serve to confirm the extension of the spatial distribution of L. merlai to the south and of the chronological distribution of this taxon from the middle to the early late Villafranchian. Finally, we offer critical remarks on some not well defined Leptobos species. [ABSTRACT FROM AUTHOR]

Abstract 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. [ABSTRACT FROM AUTHOR]

Background and scope: The late Villafranchian large mammal age (~2.0–1.2 Ma) of the Early Pleistocene is a crucial interval of time for mammal/hominin migrations and faunal turnovers in western Eurasia. However, an accurate chronological framework for the Balkans and adjacent territories is still missing, preventing pan-European biogeographic correlations and schemes. In this article, we report the first detailed chronological scheme for the late Villafranchian of southeastern Europe through a comprehensive and multidisciplinary dating approach (biochronology, magnetostratigraphy, and cosmogenic radionuclides) of the recently discovered Lower Pleistocene vertebrate site Tsiotra Vryssi (TSR) in the Mygdonia Basin, Greece. Results: The minimum burial ages (1.88 ± 0.16 Ma, 2.10 ± 0.18 Ma, and 1.98 ± 0.18 Ma) provided by the method of cosmogenic radionuclides indicate that the normal magnetic polarity identified below the fossiliferous layer correlates to the Olduvai subchron (1.95–1.78 Ma; C2n). Therefore, an age younger than 1.78 Ma is indicated for the fossiliferous layer, which was deposited during reverse polarity chron C1r. These results are in agreement with the biochronological data, which further point to an upper age limit at ~1.5 Ma. Overall, an age between 1.78 and ~1.5 Ma (i.e., within the first part of the late Villafranchian) is proposed for the TSR fauna. Conclusions: Our results not only provide age constraints for the local mammal faunal succession, thus allowing for a better understanding of faunal changes within the same sedimentary basin, but also contribute to improving correlations on a broader scale, leading to more accurate biogeographic, palaeoecological, and taphonomic interpretations.

This paper aims to contribute to the stratigraphic and geochronological evaluation of the primate bearing Dafnero fossil site of Northern Greece by means of lithostratigraphic, paleomagnetic and paleontological analyses. The 60 m thick fossiliferous deposits of fluviatile origin are recognized as representing a typical braided-river sequence unconformably overlying molassic sediments. Rock magnetic investigations indicate the presence of both medium and low coercivity minerals. Paleomagnetic sampling of the Dafnero sediments yielded a stable magnetic remanence, and the characteristic remanent magnetization directions pass reversal test with dual polarity. Based on calibration from mammal fossils, the normal polarity magnetozone N1 located in the upper third of the studied section could correlate with chron C2n (the Olduvai subchron), suggesting that the fossil horizon is within C2r with an extrapolated age of 2.4–2.3 Ma and rather closer to the upper age limit. The results allow the re-calibration of several middle Villafranchian assemblages of S. Balkans and the correlation of the corresponding mammal fauna with the environmental shifts of Praetiglian, as it is recorded in climatostratigraphic data from the Black Sea.

Abstract A study of the terrestrial gastropods of the Pliocene-Quaternary succession of the Beni Saf sea cliff, NW Algeria, at the Playa Port locality, is presented herein. The sedimentary succession is subdivided into four lithostratigraphic units, on the basis of their lithologic and biogenic contents: the three first units (A, B and C) are of marine origin; the last unit (D) is of continental origin and includes three beds yielding terrestrial gastropod fossils, which form laterally traceable horizons. Herein, we describe the section's lithology, present stratigraphical considerations regarding its age and thoroughly described its terrestrial gastropod fossil fauna. In total, 13 species are reported here from Beni Saf: 2 caenogastropods (family Pomatiidae) and 11 stylommatophorans (family Achatinidae and superfamily Helicoidea). The sedimentological data indicate that the depositional setting at Beni Saf was a dune system flanked by wadi floodplains deposits (snail levels); the ecological preferences of the gastropods largely agrees with this scenario. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

In the present study we provide a paleontological and chronostratigraphic review of the Villafranchian fossiliferous sites of Latium, revising the biochronologic attribution based on their framing within the geodynamic and paleogeographic evolutionary picture for this region. Aimed at this scope, we reconstruct the sedimentary and structural history of the Early Pleistocene marine basins through the review and the regional correlation of published stratigraphic sections and borehole data. Moreover, we combine the chronostratigraphic constraints provided in this study to the near-coast deposits of Gelasian-Santernian age (2.58–1.5 Ma) with the results of a recent geomorphologic study of this area, allowing us to reconstruct a suite of terraced paleo-surfaces correlated with marine isotopic stages 21 through 5. By doing so, we provide further age constraints to the sedimentary and tectonic processes acting on the Tyrrhenian Sea margin in Quaternary times, highlighting a possible different paleogeographic evolution of the southern coastal area where the Middle-Villafranchian type-section of Coste San Giacomo site is located, with respect to the northern sector. [ABSTRACT FROM AUTHOR]

Climatic oscillations at the Gelasian/Calabrian transition modified terrestrial palaeoenvironmental settings in the European region. A gradual drop in global temperatures beginning about 2.7 Ma led to drier conditions and to a reduction in, and subsequent disappearance of, sub-tropical vegetation in the central Mediterranean area by ca 1.2 Ma. Large ungulates are sensitive to vegetation changes and faced with harsher environmental settings may shift their feeding strategies to exploit available food resources in different ecosystems. In fossil assemblages such dietary adaptations are reflected by tooth morphology (a phylogenetic signal) and tooth wear degree (a direct signal of the species' diet). In this paper, we investigate how large herbivores responded to palaeoenvironmental changes that occurred at the passage between the Gelasian and Calabrian ages in the Italian Peninsula, analysing the dental wear patterns and hypsodonty of the early Pleistocene fossil ungulates assemblage of Olivola (Aulla, Central Italy). We found that while ungulate feeding behaviours during the Gelasian spanned from browsers to grazers, in the locality of Olivola this group of herbivores display a narrower range of diet types with many taxa adopting a mixed feeding behaviour. Cervids in particular, whose fossils are often associated only with wooded environments, as a response to the reduction of covered sub-tropical vegetation, shifted from a strict browsing diet in mostly closed habitats to a more abrasive one taking advantage of the spread of open landscapes. We also provide new data on the feeding behaviour of the rare fossil caprines, Procamptoceras and Gallogoral meneghinii , suggesting a grass-rich mixed diet for the former and a certain degree of dietary plasticity for the latter. Our research thus contributes to the better understanding of how ungulates adapted in the past to exploit different resource types during pivotal climatic changes and how environments changed in Central Italy on the onset of colder and more arid conditions. [ABSTRACT FROM AUTHOR]

The early Pleistocene is represented by a succession of glacial–interglacial cycles characterized by a general tendency towards global cooling, with increasing aridity and seasonality. The large deer Eucladoceros is found in abundance in Europe during this period of faunal dispersions. The dietary plasticity of Eucladoceros and how it can mirror early Pleistocene climatic variations will be explored here using Dental Microwear Texture Analysis. The wide range of dental microwear textures for Eucladoceros reflects a low selectivity and high plasticity in its diet. It is an appropriate proxy to track vegetal resource availability. Oscillations were identified between a browsing and a grazing signal. This study proposes that a browsing signal is associated with a fossil assemblage deposited during an interglacial event characterized by warmer temperatures and deciduous vegetation. A grazing signal more likely indicates a glacial event with cooler temperatures and a developed herbaceous, bushy layer. [ABSTRACT FROM AUTHOR]

Among living canids, the genus Nyctereutes Temminck, 1838 was the first to appear in the Western European fossil record. In the Italian Peninsula, scanty remains from a few Plio-Pleistocene localities of central Italy, referable to the Triversa Faunal Unit (FU) and the Montopoli FU, were historically attributed to Nyctereutes megamastoides (Pomel, 1842). Here we describe the partially unpublished Nyctereutes remains from two localities of the Lower Valdarno (Tuscany), S. Giusto (Florence; early Villafranchian) and Montopoli (Pisa; middle Villafranchian), and compare them with both extant and fossil species of this genus. The material described herein most closely resembles the remains from Spain, France and other European sites previously attributed to N. megamastoides in proportions and morphological features, hence confirming the presence of this relatively widespread species in Italy from the Pliocene to the Early Pleistocene. Besides the scanty material, the Lower Valdarno record of N. megamastoides represents one of the earliest of the species in Europe and allows to report, for the first time, the occurrence of a derived form of Nyctereutes in the Late Pliocene of Italy. This early record confirms that the evolutionary pattern of Asia, where the advanced N. sinensis is associated with the primitive N. tingi, was different in the Pliocene of Europe.

Among living canids, the genus Nyctereutes Temminck, 1838 was the first to appear in the Western European fossil record. In the Italian Peninsula, scanty remains from a few Plio-Pleistocene localities of central Italy, referable to the Triversa Faunal Unit (FU) and the Montopoli FU, were historically attributed to Nyctereutes megamastoides (Pomel, 1842). Here we describe the partially unpublished Nyctereutes remains from two localities of the Lower Valdarno (Tuscany), S. Giusto (Florence; early Villafranchian) and Montopoli (Pisa; middle Villafranchian), and compare them with both extant and fossil species of this genus. The material described herein most closely resembles the remains from Spain, France and other European sites previously attributed to N. megamastoides in proportions and morphological features, hence confirming the presence of this relatively widespread species in Italy from the Pliocene to the Early Pleistocene. Besides the scanty material, the Lower Valdarno record of N. megamastoides represents one of the earliest of the species in Europe and allows to report, for the first time, the occurrence of a derived form of Nyctereutes in the Late Pliocene of Italy. This early record confirms that the evolutionary pattern of Asia, where the advanced N. sinensis is associated with the primitive N. tingi, was different in the Pliocene of Europe. [ABSTRACT FROM AUTHOR]

Various species and ecomorphotypes of the genus Equus are recorded in several southern European Early Pleistocene local faunal assemblages (LFAs), though their taxonomy, phylogenetic relationships, and chronological distribution are still a matter of debate. This article aims to increase knowledge on the European pre-Olduvai stenonoid horses by describing and discussing the equid sample from the middle Villafranchian (Gelasian) Italian site of Coste San Giacomo (CSG; Anagni). Although horse remains from CSG are scanty, the morphological traits, dimensions, and proportions of teeth and some limb bones suggest some affinities with middle-sized European stenonoid horses, in particular with Equus senezensis, as supported by statistical analysis. This opens a new window on the possible phylogenetic relationships of the middle-sized, slender middle Villafranchian horses, although some questions about their actual taxonomic rank cannot be firmly answered based on available data. The complex relationship among environment, structure of the accompanying mammalian fauna, and the presence in the Early Pleistocene LFAs of only one equid, large or middle-sized, or even of more horse species with different size is briefly discussed. [ABSTRACT FROM AUTHOR]

A recent find of a Middle Villafranchian (c. 2.35–2.10 Ma) Canis cf. C. etruscus in the trawlings from the Oosterschelde concerns the oldest dog known from the Netherlands and is the first appearance record of this canid in the North Sea Basin. It shows that the tribe Canini was dispersed beyond south central Europe up to the northwestern edge of the continent. The find confirms the lack of synchroneity and usefulness of the so-called ‘Wolf Event’. [ABSTRACT FROM PUBLISHER]