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Isolated teeth and bones of rhinoceroses are sometimes difficult to identify at specific level because our poor knowledge of the morphological variability of fossil and, often, of extant species. During the past century, several studies have been devoted to highlight morphological and morphometric differences among the fossil Pleistocene species, but only a few of them considered the variability of the extant taxa, in particular Dicerorhinus sumatrensis , to support and interpret their results. Dicerorhinus sumatrensis is the closest extant relative of the Eurasian Stephanorhinus , as recently evidenced by molecular analyses, and it provides useful data to better understand the variability in extinct Pleistocene Eurasian taxa. Taking into account the observations on the extant Sumatran rhinoceros, the validity of some morphological traits, often used for specific attribution in Early Pleistocene fossil rhinoceroses, is revised. Sexual dimorphism and ontogenetic stage drive the development of some important features in Eurasian rhinoceroses and a detailed comparison among crania lead to new considerations on taxonomy, and, therefore, on paleobiogeography and biochronology of the different species. [ABSTRACT FROM AUTHOR]

Sexual dimorphism has been poorly evaluated or investigated in Pleistocene Eurasian Stephanorhinus species, leaving a gap in our knowledge about their morphometric variability. Among the representatives of this genus, S. etruscus is the most abundant species, with several remains collected from Western European localities, allowing us to investigate the presence of sexual dimorphism in the limb bones of this taxon. We considered measurements taken on 45 postcranial variables and three different statistical metrics to identify patterns of bimodality in the dataset. This work represents the first application of sex-combined statistical analysis to a dataset composed of individuals from various European localities. The morphometrical analyses revealed that a relatively weak sexual dimorphism is present in all the considered bones. Larger forelimbs and hindlimbs are interpreted as belonging to adult males of S. etruscus, similarly to what was observed in the modern Sumatran rhino, where males are a little bit larger than females. The recognition of a weak sexual dimorphism in the postcranial bones of S. etruscus increases our understanding of the paleoecology of this extinct taxon. However, only a better study of the morphological and morphometrical variability of the crania of fossil rhinoceroses could deeply contribute to the investigation of the social habits and behavior of these taxa. [ABSTRACT FROM AUTHOR]

The present study deals with a new bovid assemblage from Kataphyto, Northeastern Greece, providing additional information concerning this species dispersal and stature in Greece. It consists of 155 teeth and bone remains attributed to the extinct species Bos primigenius Bojanus, 1827. The 14C dating analysis of the fossils resulted in a Late Pleistocene age, with a median at 23,800 calBP, similar to the age obtained using the Useries geochronology. A brief outline of the evolutionary history, morphology and ecology of the Pleistocene aurochs is reported, with a comprehensive update of the palaeontological and archaeological records of the species in Greece reviewed here. Morphometric data have been investigated in order to provide an evaluation of the species body size variations throught time in Greece. The results show that the populations of aurochs in Greece were small in size during the Middle Pleistocene, increased in size during the end of the Late Pleistocene, followed by a decrease during the Holocene. Moreover, aurochs remains from Greece, compared with those obtained from the Italian Peninsula, contribute to knowledge about body size variation during the Pleistocene and Holocene. This study revealed a different trend compared to other European sites and depict a more complex and articulated framework on aurochs body size variation than previously thought. [ABSTRACT FROM AUTHOR]

Pliocene and earliest Pleistocene Northern Eurasian rhinocerotines are poorly documented and understudied in comparison to Pleistocene and Miocene ones. However, they represent a key-group of species for understanding the phylogeny and historical biogeography of their Pleistocene relatives. In the present paper, we revise the abundant material from the late Pliocene locality of Kvabebi, Georgia from a systematic, phylogenetic and palaeobiogeographical perspective. The specimens from Kvabebi are documented by two partially preserved skulls, one mandible and several postcranial remains. Morphological and morphometric comparison with the type material assigned to Pliocene and earliest Pleistocene Northern Eurasian Rhinocerotinae reveal that the specimens from Kvabebi have close affinities with the poorly known Dicerorhinus miguelcrusafonti Guérin & Santafé-Llopis, 1978, described from Layna (Spain). The latter species, represented by scanty remains from the Iberian Peninsula, is usually excluded from morphological and morphometrical comparisons and no findings were reported after the 1990s. Pliocene rhinocerotine species have monotonous dental and postcranial morphologies and only a few features allow us to discern the different species. The material from Layna and Kvabebi is somewhat smaller than that of other Pliocene taxa, except for the largest representatives of Stephanorhinus etruscus (Falconer, 1868). Accordingly, the earliest specimens assigned to S. etruscus on morphometric grounds should be revised in the light of the new data here presented. A cladistic analysis performed on 280 characters and 30 species suggests that the emblematic early Pliocene European species, 'Dihoplus' megarhinus (de Christol, 1834), is sister taxon to the Layna and Kvabebi rhinoceroses. Accordingly, both species are here assigned to a new genus named Pliorhinus gen. nov. Although distinct, this clade has close affinities with the paraphyletic genus Stephanorhinus, therefore suggesting the co-occurrence of at least two distinct rhinocerotine lineages raised in the late Miocene interval in Northern Eurasia. [ABSTRACT FROM AUTHOR]

A new species of Rhinocerotidae (Perissodactyla), 'Ceratotherium' advenientis sp. nov., from the late Miocene (8.1–7.2 Ma) locality of Cava Gentile, Calabria (Southern Italy), is described. 'Ceratotherium' advenientis displays morphological characters close to Rhinocerotina, in particular to dicerotines, and can be distinguished from the late Miocene elasmotheres, teleoceratines and aceratheres recorded in Eurasia and Africa. The new taxon clearly differs from the European latest Miocene species Dihoplus schleiermacheri, Dihoplus pikermiensis, 'Dihoplus' megarhinus and Ceratotherium neumayri, and from the African species Ceratotherium douariense, Ceratotherium? primaevum and Paradiceros mukirii. 'Ceratotherium' advenientis also differs from the Chinese dicerotine Diceros gansuensis and from the extant African species. The new taxon is characterized by peculiar features, in particular in the morphology and dimension of the neurocranial portion, and by having a nuchal crest wider than in the extant African rhinoceroses, C. neumayri, C. douariense, and European latest Miocene species. A cladistic analysis places 'Ceratotherium' advenientis in a polytomy with the extant Diceros bicornis, C. neumayri and a small clade composed by C. simum and C. antiquitatis. The African affinities of the new taxon support the Calabrian-Peloritan arc as a northern extension of the African continental shelf during the late Miocene. [ABSTRACT FROM AUTHOR]

The Plio-Pleistocene fossil rhinoceroses of Eurasia are relatively well known, although their taxonomy and phylogeny are still debated. The fossil rhinoceros material collected at the Bethlehem site during the first half of the 20th century is revised and re-described. Based on morphological comparison of the Bethlehem material with the latest Miocene, Pliocene and Early Pleistocene Eurasian and African species, the specimens are assigned as Rhinocerotini indet., Dihoplus sp. and 'Dihoplus' bethlehemsis sp. nov. The comparison revealed that the studied material displays more features in common with the genus Dihoplus than with the genus Stephanorhinus. Tooth microwear analysis revealed that the diet of Bethlehem rhinoceros was probably composed of leaves of trees, bushes, or part of dicots. The new species could be one of the latest representatives of the Miocene genus Dihoplus that survived in Eurasia until the end of the Pliocene. Thus again the site of Bethlehem being unique in its biogeographically setting, provides novel understanding of the 'Levantine Corridor' dispersal route. [ABSTRACT FROM AUTHOR]

The Rhinocerotidae material from the early Pleistocene Tetoiu Formation of Colțești (southwestern Romania) is here described for the first time. The rhinoceros is documented by calcaneus and second, third, and fourth metatarsals, probably belonging to the same individual. The morphology and the dimensions of these specimens enable us to record the presence of Stephanorhinus jeanvireti , a relatively rare rhinoceros usually reported from late Pliocene European localities and recently considered a junior synonym of S. elatus. Nevertheless, the taxon Rhinoceros elatus is here regarded as a nomen dubium and the name S. jeanvireti is retained in order to maintain nomenclatural stability, being it based on much more diagnostic material. In Romania, S. jeanvireti has been listed within a few late Pliocene faunal assemblages, but the rhinoceros remains are fragmentary and isolated bones. The Colțești find is among the best documented records of S. jeanvireti in Romania. The biochronological distribution of S. jeanvireti is mainly confined within the MNQ16 (early and early middle Villafranchian), and its first occurrence is doubtfully reported in late MNQ15 faunas. The record of Colțești (MNQ17/MNQ18) represents, instead, the last occurrence of this taxon in Europe. [ABSTRACT FROM AUTHOR]

Most of the research on fossil mammals from Umbria (central Italy) has been carried out in the southwestern branch of the Tiber basin, due to its paleontological richness. This portion of the basin extends from Perugia to Terni and corresponds to a well-defined half-graben filled with fluvial-lacustrine deposits. The paleontological sample presented here was discovered in a sand and gravel quarry at Podere San Lorenzo, East of the town of Deruta. The stratigraphic succession exposed in the quarry is no longer visible, but we describe here a new outcrop (Palazzone), which is not far from Podere San Lorenzo and shows comparable facies associations. The two successions were deposited in a fluvial environment characterized by an average reduction of the hydrodynamic energy from the bottom upwards. They are referred to the Early Pleistocene Santa Maria di Ciciliano Subsyntheme (Madonna dei Bagni Lithofacies). Large mammal remains are attributed to Mammuthus cf. meridionalis (Nesti, 1825), Stephanorhinus etruscus (Falconer, 1859), Equus stenonis Cocchi, 1867, Leptobos cf. etruscus (Falconer, 1868), 'Pseudodama' nestii (Azzaroli, 1947), and Sus strozzii Forsyth Major, 1881. Some hyena coprolites are also reported. The assemblage is typical of the early Late Villafranchian Land Mammal Age and can be referred to the Olivola/Tasso Faunal Units (about 2.0-1.8 Ma). This is in agreement with the alleged age of some other assemblages found in the southwestern branch of the Tiber basin (e.g., Torre Picchio, Villa San Faustino, Colle Sant'Andrea, Pantalla). [ABSTRACT FROM AUTHOR]

Several taxa belonging to the so called megafauna became extinct during the late Quaternary in Eurasia. The extinction chronology of the narrow-nosed rhinoceros, Stephanorhinus hemitoechus, in Europe is still uncertain and only estimated around 45 ka. A systematic revision of several rhinoceros findings reveals that this species occurred in Italy at least untill 41 ka BP, at the onset of the Heinrich Event 4. Climatic fluctuations during MIS 3 and habitat fragmentation probably created an increase of small S. hemitoechus populations in southern Europe which had a tendency to become extinct. [ABSTRACT FROM AUTHOR]

We undertake a comparative mammalian zoogeographic analysis with the aim of revealing the extent to which the Dmanisi Early Pleistocene large mammal assemblage resembles, at the genus level, African, Arabian, and Eurasian localities of similar age. The inclusion of Old World Pliocene and Pleistocene mammalian faunas provides us with insights into the provincial origins of specific mammalian taxa and permits us to assess the relative affiliation of the Dmanisi mammalian faunas to other faunas in the Old World. Our analysis also allows us to consider hypotheses about the timing and direction of zoogeographic connections between western Eurasia and Africa during the Early Pleistocene. We utilize multiple zoogeographic analytical tools as a cross-comparison of Dmanisi with 42 other Eurasian and African mammalian-bearing localities between 2.7 and 0.7 Ma. Overall, we find that Dmanisi compares most closely with a subgroup of Greek, Italian, and Spanish localities that are slightly younger than Dmanisi itself. This could suggest a progressive dispersal from East to West of the large mammal communities during the late Early Pleistocene and the first occurrence at Dmanisi, and then later in Western Europe, of some taxa such as Stephanorhinus ex gr. etruscus-hundsheimensis , Equus altidens, Bison georgicus, Soergelia minor, Megantereon whitei , Canis borjgali , Canis (Xenocyon) lycaonoides. Dmanisi's habitats included drier areas, probably of open wooded savannah and grassland and by mountainous to semiarid rocky terrain. There is evidence that Dmanisi records short intervals of increased aridity in the middle part of the succession contemporaneous with the occurrence of Homo. [ABSTRACT FROM AUTHOR]

Hippopotamuses were present in Western Europe during the Pleistocene with at least two continental species: Hippopotamus antiquus and H. amphibius. The former was the large European hippopotamus, which appeared in the fossil record around 2 Ma and successfully spread across Europe before its last appearance datum (LAD) in Portugal and Greece, dated to 0.4 Ma. The extant hippopotamus, H. amphibius , first occurred in Africa around 1.5 Ma, and was well-established in Europe during the Late Pleistocene; however, its first occurrence datum (FOD) in Europe is still highly debated. Most of the H. amphibius remains collected from Middle Pleistocene localities are either badly preserved and fragmented or lacking clear stratigraphical data, in turn leading to doubtful specific attributions. A well-preservedmandible herein described, collected from Fosso Malafede (Latium, central Italy), allows to re-assess the occurrence of H. amphibius and to critically revise its earliest records. The morphological and morphometric characters of the studied remain, such as the general robustness of the studied mandible, the convex ventral profile of the mandibular corpus, the length of the teeth series, and the shape of the sagittal cross section of the mandibular symphysis, allow us to ascribe it to Hippopotamus amphibius. Fosso Malafede has been chronologically assigned to MIS 7 (∼0.25 Ma) and represents one of the best preserved H. amphibius specimen of the Middle Pleistocene of the European mainland. Accordingly, we discuss biochronological and paleobiogeographic implications, suggesting a critical revision of several remains from the Middle Pleistocene of southern Europe. • A mandible collected from a Middle Pleistocene site in Italy adds new morphological and morphometric data on European Hippopotamus. • 3D GMM is here applied to investigate hippopotamid mandibles. • The virtual restoration of the specimen represents an important step towards a better valorisation of the fossil material. • During MIS 7 Hippopotamus amphibius was spread in the Italian Peninsula. [ABSTRACT FROM AUTHOR]

Equus stenonis is one of the most iconic fossil species of the Early Pleistocene. Although it is known from the 19th century, its taxonomic and evolutionary position has been extensively debated among equid specialists, including multiple interpretations. The current work aims to review the European E. stenonis record, by morphological, morphometrical and statistical analyses combined with body mass and net primary production estimates. Furthermore, we investigate the evolution of E. stenonis cranial anatomy in comparison with the Plio-Pleistocene species from North America, Eurasia and Africa, and extant zebras and asses. Our review of European E. stenonis samples has closely considered the E. stenonis subspecies Equus stenonis vireti , Equus stenonis guthi , Equus stenonis pueblensis , Equus stenonis olivolanus and Equus stenonis stenonis which we recognize as being ecomorphotypes of Equus stenonis , with the exception of Equus stenonis senezensis and Equus stenonis mygdoniensis. These two last subspecies warrant the species ranks of Equus senezensis and Equus altidens , respectively. This has led us to propose a new revised biochronology of European Early Pleistocene Equus. Our evolutionary interpretations based on cranial morphology reveals that the evolutionary lineage of extant zebras can be recognized in deep time from the North American species Equus simplicidens, through the European E. stenonis and the African Equus koobiforensis. European stenonine horses likewise represent the original source for the African Pleistocene fossil species and extant Equus grevyi. • This revision provides new insights on the Early Pleistocene species Equus stenonis and on its relatives forms. • The Equus stenonis subspecies shown a broad intraspecific variability shown in cranial and postcranial elements. • E. stenonis vireti , E. stenonis guthi , E. stenonis pueblensis , E. stenonis olivolanus and E. stenonis stenonis are different ecomorphotypes of E. stenonis. • Equus stenonis senezensis and Equus stenonis mygdoniensis are related to species rank of Equus senezensis and Equus altidens. • The evolutionary lineage of the extant largest zebras can be recognized through the North American E. simplicidens , the European E. stenonis and the African E. koobiforensis. [ABSTRACT FROM AUTHOR]