Stephanorhinus Research Articles

New early Pliocene Rhinocerotidae findings from Tuscany (Italy) and the Pliocene rhinocerotine record in Italy

An almost complete mandible from the Lower Pliocene deposits of Arcille (Tuscany) is described and investigated for the first time. The specimen is assigned to Pliorhinus megarhinus, which is documented in Italy from the latest Miocene to the mid-Pliocene. The studied specimen is larger than Stephanorhinus etruscus and S. jeanvireti, and morphologically fits P. megarhinus from the Early Pliocene of Montpellier (France), Vera Basin (Spain) and Val di Pugna Fangonero (Italy). This specimen represents the first stratigraphically well-documented Pliocene record of this species in Italy, the previous discoveries dating back to the 19th century or the beginning of the 20th century, hence lacking stratigraphic control. According to the obtained results, it is possible to assess the occurrence of P. megarhinus during the earliest Pliocene in Italy as well as to confirm the long-lasting persistence of this species, whose fossil record covers a timespan between 6.7 Ma and 3.5 Ma. During that time, P. megarhinus did seemingly undergo a trend of body mass reduction.

The postcranial variability of Quaternary European rhinoceroses: the case study of Stephanorhinus hundsheimensis from the Middle Pleistocene site of Contrada Monticelli (Apulia, southern Italy)

ABSTRACT Five rhinoceros species, mainly distinguishable through cranial, mandibular and dental morphology, were recognized during the Quaternary across Europe. The postcranial morphology of these taxa is quite variable, with strong superposition among species, especially between Stephanorhinus etruscus and Stephanorhinus hundsheimensis. Here, the complete sample including 25 postcranial bones from the early Middle Pleistocene site of Contrada Monticelli (Apulia, southern Italy) is described and compared through morphological and biometric analyses. A few bones, in addition to cranial and mandibular remains from this site, have been previously referred to Stephanorhinus hundsheimensis. We explored the morphological variability of Stephanorhinus hundsheimensis from Contrada Monticelli against the diagnostic characters described in the literature. The size variation of the Contrada Monticelli sample is also investigated, through a comparison with other Quaternary European rhinoceros samples. Our results indicate a wide morphological variability with several distinct characters and a strong morphological affinity between Stephanorhinus etruscus and S. hundsheimensis. Our results suggest that Stephanorhinus hundsheimensis from Contrada Monticelli is a species with a high degree of intraspecific variability, possibly a hybrid population or a population including two morphotypes of the same species.

Sus strozzii (Suidae, Mammalia) from the historical locality of Quercia (Early Pleistocene, Italy)

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.

Extinct Eurasian rhinoceros Coelodonta and Stephanorhinus dental pathologies and tooth change modus

Dental pathologies are described for four extinct rhinoceroses Coelodonta antiquitatis (Blumenbach, 1799) and Stephanorhinus hemitoechus (Falconer, 1868) of the Late Pleistocene Weichselian Glacial (MIS 3-5 d), Stephanorhinus kirchbergensis Jäger, 1839 of the Late Pleistocene Eemian Interglacial (MIS 5e), and Stephanorhinus hundsheimensis (Toula, 1902) from the Middle Pleistocene Cromerian Mosbach Interglacial (MIS 11) which material is mainly from the Rhine Valley megafauna migratory channel of Central Europe. All three extinct and even extant rhinoceros share similar ontogenetic P4 pathologic misplacement pathologies. Those caused in some cases further dental abnormalities impacting the lower jaw dentition, even in rare cases including complete deformation of the lower jaw. In the browsing Stephanorhinus (S. kirchbergensis, S. hundsheimensis) the P4 is more commonly misplaced; in the Mid-Pleistocene Mosbach S. hundsheimensis are three examples with similar misplacements on the right and left maxillaries, which have a similar genetic pool background of a local palaeopopulation. The grazer rhinoceros C. antiquitatis has a further pathology type with loss or even genetically pre-assessed non-development of the first upper or lower PM premolars or last M molars. The strong wear, damage, and loss of the anterior premolars in elderly individuals as ontogenetically based pathology, seem to result from intensive grazer and opal phytoliths abrasion. This loss finally resulted in irregularities of the permanent dentition and influenced wear stages and irregular abrasion angles of the tooth rows, affecting the upper and lower rows each others.

The woolly rhinoceros (Coelodonta antiquitatis) from Ondorkhaan, eastern Mongolia

The Mammoth Faunas, the famous cold‐adapted mammal assemblages, were distributed widely across northern Eurasia and North America during the Late Pleistocene. The now extinct woolly rhinoceros, Coelodonta antiquitatis, was a major component. Abundant fossil remains of this species with radiocarbon dates have been reported through almost all of northern Eurasia, but the fossil rhinoceroses of Mongolia are poorly known. Here, we describe a rhinocerotid skeleton from Ondorkhaan, eastern Mongolia, and compare it with four Late Pleistocene rhinoceros species of northern Eurasia (Elasmotherium sibiricum, Stephanorhinus hemitoechus, Stephanorhinus kirchbergensis and C. antiquitatis), resulting in its identification as a woolly rhinoceros (C. antiquitatis). Accelerator mass spectrometry radiocarbon dates obtained from two samples (ODK01 and ODK02) of the skeleton essentially agree, 42 160–40 040 cal. a BP and 42 105–39 955 cal. a BP, and the two samples had δ13C values of −19.5‰ and −20.2‰ SMMKW, respectively. This find suggests that the Mammoth Faunas were distributed in eastern Mongolia c. 45–40 ka during the period of climatic amelioration between Heinrich events 5 (H5: 46 000 cal. a BP) and 4 (H4: 39 000 cal. a BP).

New Pliocene Rhinocerotidae findings from the Iberian Peninsula and the revision of the Spanish Pliocene records

A fragmentary mandible from the early Pliocene of the Vera Basin is here investigated to clarify its taxonomic position. The studied specimen is here assigned to Pliorhinus megarhinus, documented in Europe from the latest Miocene to the mid-Pliocene. The morphometrical comparison reveals that this taxon is larger than Stephanorhinus etruscus and S. jeanvireti but overlaps with Dihoplus schleiermacheri. P. megarhinus and D. schleiermacheri differ in some characters of the lower molars, such as the length of the metalophid and the shape of the trigonid in occlusal view, thus allowing new considerations on the Pliocene Spanish rhinocerotine record. According to the results here obtained, three species are at least documented in Span during the Pliocene: P. megarhinus, P. miguelcrusafonti, and S. etruscus. The presence of S. jeanvireti has to be verified and confirmed by further evidence. The presence of Dihoplus schleiermacheri as a relict taxon in the Pliocene of Spain is here discarded.

New taxonomical, biochronological and palaeoenvironmental data from the Middle Pleistocene site of Cúllar de Baza 1 (Granada, Spain)

The intra-montane Guadix-Baza Basin is one of the few continental basins in Europe that hosts a well-dated set of fossiliferous sites spanning from the latest Miocene to the late Middle Pleistocene. The Cúllar de Baza 1 (CB-1) represents a key site to investigate the effects of the Early-Middle Pleistocene Transition, considered a fundamental transformation in the Earth’s climate state. Our review and update of the large mammal assemblage, and particularly equids, is of paramount relevance to understand the systematic affinities and the evolution of the Early and Middle Pleistocene European horses. We confirm the occurrence of two different taxa, the medium sized Equus altidens and the larger E. suessenbornensis. Moreover, we illustrate that CB-1 is essential for the biochronological studies of the latest Early Pleistocene/Middle Pleistocene transition (Epivillafranchian/Galerian ELMA); in particular with regard to the Last Occurrences of the Etruscan rhino Stephanorhinus etruscus and the large deer Megaloceros savini and the First Occurrence of the water-rat Arvicola mosbachensis in the Iberian peninsula. Finally, a development of a mosaic environment characterised the CB-1 site contrasting with the conditions reported for other Iberian late Early and Middle Pleistocene localities.

The Villafranchian perissodactyls of Italy: knowledge of the fossil record and future research perspectives

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.

The Koskobilo (Olazti, Navarre, Northern Iberian Peninsula) paleontological collection: New insights for the Middle and Late Pleistocene in Western Pyrenees

The destroyed site(s) of Koskobilo (Olazti, Navarre, Northern Iberian Peninsula) have yielded unique archaeo-paleontological evidence in the Western Pyrenees region. The quarry uncovered a karstic site with faunal remains in 1940, and fossils were recovered both in situ and from the quarry dump. Ten years later, while the quarry was still working, a new visit to the dump yielded a large lithic assemblage and additional fossil remains with a different taphonomic pattern, which has been interpreted as the remains coming from a different site or zone within the same karst system. Here we re-study the paleontological evidence and provide new dating on a speleothem covering a Stephanorhinus hemitoechus tooth, which has yielded a minimum date of c. 220 ka for part of the assemblage. In total, the fossil assemblage comprises 38 mammal and six avian taxa and three fish remains. The faunal evidence indicates that in 1940 a mix of taxa from both the Middle and Upper Pleistocene were recovered, and it is difficult to assign most of them to a concrete period. However, based on biochronological criteria some of the identified taxa (e.g., Ursus thibetanus, Ursus cf. deningeri, Cuon cf. priscus, Macaca sylvanus, cf. Megaceroides) could be roughly contemporaneous with the dated rhino tooth, which would provide a new window to the Middle Pleistocene of the region, with deposits from MIS 7d and/or older. Despite the difficulties in studying this collection, recovered without stratigraphic context and in a salvage operation, Koskobilo has yielded an important paleontological assemblage which helps to understand the paleoecology of the Middle Pleistocene human occupations in the Western Pyrenees.

Rhinoceros (Stephanorhinus hemitoechus) exploitation in Level F at the Caune de l'Arago (Tautavel, Pyrénéés‐Orientales, France) during MIS 12

AbstractCaune de l'Arago is a Middle Pleistocene site in Southern France, where Acheulean artefacts and hominin fossils were excavated. Rhinoceros (Stephanorhinus hemitoechus) remains from Level F (MIS 12) were studied from a zooarchaeological and taphonomic perspective to investigate the potential human exploitation of this large taxon. The well‐represented butchering marks, as well as scarce carnivore marks, indicate primary access to rhinoceros carcasses by hominins. The juvenile‐dominated mortality profile further suggests aggressive scavenging or occasional opportunistic hunting. Furthermore, differential skeletal representation shows that humans selectively transported the nutrient‐rich body sections. In addition, thorough processing of the carcasses for consumption inside the cave is highlighted by the frequent cut marks, intensive fragmentation, and regular spatial distribution. The analyses of the Arago Level F rhinoceros confirm the rhinoceros exploitation by humans during the Lower Palaeolithic; however, the rhinoceros mortality profile differs from that of large mammals such as equids, argali, and all other game species of this archaeological layer. Caune de l'Arago is the earliest hominin site where the systematic exploitation of rhinoceros has been documented.

Biogeography and chronology of the Eurasian extinct rhinoceros Stephanorhinus etruscus (Mammalia, Rhinocerotidae)

Stephanorhinus etruscus is one of the most abundantly recorded and better known Eurasian Early Pleistocene rhinoceroses. Nevertheless, the first and last appearances of this species, as well as its paleogeographic distribution, are controversial and debated in literature. S. etruscus is documented since the latest Pliocene in Spain (Las Higueruelas), Italy (Montopoli and Castelnuovo di Barardenga), France (Perrier–Les Étouaires) and Romania (Iarăș–Cariera Veche). During the Early Pleistocene, S. etruscus occurred in several Spanish, French and Italian localities, as well as in The Netherlands (e.g., Tegelen), Germany (e.g., Thiede), Greece (e.g., Aivaliki) and Israel (e.g., Ubeidiya). The last appearance of S. etruscus in Eurasia is debatable. Etruscan rhino populations survived till the Jaramillo subchrone (around 1.1 Ma) in France (Bois-de-Riquet), Romania (Betfia XII) and Hungary (Osztramos 2 and 8), and close to the early–middle Pleistocene transition in Spain (Cueva Victoria, Huéscar 1, Atapuerca TD4, TD6 and TD8), and Italy (Monte delle Piche).

NEW DATA ON THE LOWER PLEISTOCENE (GELASIAN) LIGNITE BEDS OF CASTEL SAN PIETRO (RIETI, CENTRAL ITALY)

A multiproxy study of the lignite-bearing lacustrine sequence cropping out at Castel San Pietro (CSP) (Rieti, central Italy) provides new insights on the paleoenvironmental evolution of an extensional basin, located on the Tyrrhenian side of the Apennine range. The CSP fossiliferous levels reflect deposition in a stable lacustrine environment. Mammal remains collected from this locality during the past centuries include Stephanorhinus etruscus, Anancus arvernensis and the beaver Castor fiber. The co-existence of the Etruscan rhinoceros and the mastodont suggests a middle Villafranchian age (Montopoli or Coste San Giacomo Faunal Unit) for the lignite beds of CSP. New paleoflora findings are in agreement with an attribution to the Piacenzian or Gelasian age for these deposits whilst the presence of the ostracod Qinghaicypris cf. Q. riojensis suggests a time interval from Zanclean to Gelasian (between ~4.5 and ~1.8 Ma). Accordingly, the lignite beds of CSP most likely were deposited during the Gelasian (from ca. 2.5 to 1.8 Ma; from Montopoli to Coste San Giacomo FUs or possibly to Olivola FU) in forested area and warm and humid conditions.

LATE PLEISTOCENE LAST OCCURRENCES OF THE NARROW-NOSED RHINOCEROS STEPHANORHINUS HEMITOECHUS (MAMMALIA, PERISSODACTYLA) IN ITALY

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.

New taphonomic data of the 1 Myr hominin butchery at Untermassfeld (Thuringia, Germany)

Abstract Untermassfeld has provided a butchered bone assemblage and a Mode 1 lithic record demonstrating hominin presence at a site which was classically viewed as paleontological. This archaeological record was found during fieldwork and surveying in fluvial river banks and low-to-medium energy channel erosion sediments. Paleomagnetism and biostratigraphy has yielded an age for this hominin occupation close to the onset of the Jaramillo subchron (ca. 1.07 Myr). In this paper we present new taphonomic data of the cut-marked and hammer-related bone material, corresponding mainly to large-sized herd species (i.e., Bison ), but also from cervids ( Eucladoceros giulii and Cervus s.l. nestii vallonnetensis ) and megafaunal species (i.e., Stephanorhinus hundsheimensis and Hippopotamus amphibius antiquus ). Distribution of skeletal elements and bone surface features suggests that this bone assemblage consists of animal remains with different taphonomic origins. The assemblage formed during at most two or three sediment-laden floods, transporting both lithic tools and remains of animals that died of natural causes or were killed by predators. The new taphonomic results presented here are important to analyze hominin colonization and adaptation to European mid-latitude environments at 1 Myr, in competition for meat resources with large carnivores (i.e., Pachycrocuta brevirostris and Panthera onca gombaszoegensis ).

Stephanorhinus hundsheimensis (Mammalia, Rhinocerotidae) from the late early Pleistocene deposits of the Denizli Basin (Anatolia, Turkey)

Pleistocene rhinoceroses are poorly documented in Turkey where they have been reported only from the late early Pleistocene (1.3–1.1Ma) travertine deposits of the Denizli Basin. In this work, new rhinoceros remains collected from this basin are assigned to a relatively large-sized Stephanorhinus hundsheimensis on the basis of their morphology and morphometry. The first Turkish record of this species is approximately coeval with the first appearance of S. hundsheimensis in Europe, chronologically referred to the late early Pleistocene, ca. 1.2Ma. During that time, S. etruscus still survived in Iberian Peninsula, central Italy and Dacian Basin. The presence of two successive evolutionary morphs for S. hundsheimensis during the Pleistocene is not confirmed.

In situ high spatial resolution 87Sr/86Sr ratio determination of two Middle Pleistocene (c.a. 580 ka) Stephanorhinus hundsheimensis teeth by LA–MC–ICP–MS

Bone and tooth tissues are important biological archives to study eating habits and provenance of ancient humans and animals. By taking advantage of the high spatial resolution offered by the Laser Ablation Multi Collector Inductively Coupled Plasma Mass Spectrometry (LA–MC–ICPMS) technique, we investigated the 87Sr/86Sr intra-tooth variability of two Stephanorhinus hundsheimensis tooth fragments from the Middle Pleistocene site of Isernia La Pineta. We detected significant Sr isotopes variability within the rhinos tooth enamel (enamel average ±2σ: sample RH–IS30 0.70951±0.00014; sample RH–IS26 0.70976±0.00015) with values higher than the “local” 87Sr/86Sr ratio (dentine average ±2σ: sample RH–IS30 0.70918±0.00013; sample RH–IS26 0.70934±0.00009). This is likely linked to a different water and food intake with a Sr isotopic signature similar to volcanic soils nearby (Roccamonfina: from 0.7093 to 0.7100; Colli Albani: >0.7100) and supports the idea that the Stephanorhinus hundsheimensis species moved around seasonally. The improvement of non-destructive, accurate and precise analytical methods to decrypt the information hidden within bone and tooth hard tissues of archeological material is crucial to unravel critical questions about evolution, migration and ecology of human and animals. We have successfully took upon this challenge using three matrix-matched reference materials, with variable Sr concentration (c.a. 100–1000ppm), to correct unresolved interferences arising from LA analyses.

Villafranchian-Galerian mammal faunas transition in South-Western Europe. The case of the late Early Pleistocene mammal fauna of the Frantoio locality, Arda River (Castell’Arquato, Piacenza, Northern Italy)

We investigated the fossil mammal assemblage from the late Early Pleistocene (early Galerian) site of the Arda River. Seven taxa were identified: one carnivore (Ursus dolinensis), five artiodactyls (Sus strozzii, Pseudodama farnetesis, Praemegaceros sp., Bison sp., and Hippopotamus sp.) and one perissodactyl (Stephanorhinus hundsheimensis). The Arda River faunal assemblage includes taxa already present in the Villafranchian, such as Pseudodama and Sus strozzii. The bear U. dolinensis and the bison, a more evolved form than Bison (Eobison), are taxa typical of the beginning of the Galerian mammal age. Paleontological, sedimentological and paleobotanical data point towards the presence of woodlands settled on an alluvial plain with patches of grassland dated around 1.0 Ma. The continental sediments of the Arda River represent one of the first steps of the forming of the alluvial Po valley.

The influence of climate on species distribution over time and space during the late Quaternary

Understanding the effect of climate on the composition of communities and its change over time and space is one of the major aims in ecology and paleoecology. Herein, we tackled on this issue by studying late Quaternary large mammal paleocommunities of Eurasia. The late Quaternary was a period of strong environmental instability, especially characterized by the occurrence of the last glacial maximum (LGM). We used community phylogenetics and joint species distribution models in order to understand the factors determining paleocommunity composition in the late Quaternary. Our results support the existence of strong climatic selection operating on the LGM fauna, both through the disappearance of warm-adapted species such as Elephas antiquus, Hippopothamus amphibious, and Stephanorhinus hemitoechus, and by setting the stage for the existence of a community characterized by cold-adapted large mammals. Patterns of abundance in the fossil record, co-occurrence between species pairs, and the extent of climatic forcing on faunal composition, differ between paleocommunities, but not between extinct and extant species, which is consistent with the idea that climate change, rather than the presence of humans, exerted a major effect on the survival of the late Quaternary megafauna.

Middle Pleistocene large‐mammal faunas from North Iberia: palaeobiogeographical and palaeoecological implications

The rich Quaternary fossil record from the Cantabrian region (NW Iberia) is virtually restricted to the Late Pleistocene, with Middle Pleistocene findings very rare. This work presents the study of two Middle Pleistocene palaeontological sites in the Cantabrian region: Llantrales quarry and Mestas de Con mine, both discovered and first described in the 1950s but never studied in detail thereafter. The material was here re‐analysed and re‐evaluated. Taxonomic attributions have been changed in most cases and, consequently, chronological assignments of the fossils have also been changed. This first detailed study of the assemblage from Llantrales quarry revealed the occurrence of the large‐sized deer Praemegaceros solilhacus, which has never been reported previously in North Iberia. Cervus cf. elaphus and Stephanorhinus cf. hundsheimensis were also identified. The age of these fossils was originally regarded as Pontian (late Miocene), but new palaeontological results indicate a much younger age, most likely from around 0.8 to 0.5 Ma. A taxonomic re‐evaluation of the fossils from Mestas de Con provided different results with respect to the 1950s works. The faunal association is composed of a large‐sized deer (probably corresponding to Praemegaceros), Cervus elaphus, Capreolus cf. capreolus, Bison sp. (small sized), Equus sp. (large sized), Stephanorhinus hemitoechus, Ursus sp. and Homotherim latidens. This faunal association was evaluated within a western European context and proves to be highly analogous to other contemporary assemblages there. The chronology was initially regarded as late Villafranchian (in a broader sense). New results, documenting the co‐occurrence of Stephanorhinus hemitoechus and Homotherium latidens, indicate a more precise and younger age, most likely between 0.6 and 0.4 Ma. These two fossil assemblages provide new and relevant information on the Middle Pleistocene faunas from a geographical region where this information, compared with other western European areas, is rare and poorly known.

Two new vertebrate localities from the Early Pleistocene of Mygdonia Basin (Macedonia, Greece): Preliminary results

Two new vertebrate fossiliferous localities from the Pleistocene of the Mygdonia Basin (Macedonia, Greece), Tsiotra Vryssi (TSR) and Platanochori-1 (PLN), are presented in this article. TSR belongs to the Gerakarou Formation and its preliminary faunal list includes the corvid Corvus pliocaenus, the hyaenid Pachycrocuta brevirostris, the canid Canis etruscus, the rhinoceros Stephanorhinus sp., two species of Equus, the bovid Bison sp., the cervid Metacervocerus rhenanus and the giraffid Palaeotragus sp. PLN belongs to the Platanochori Formation (overlying Gerakarou Formation) and its fauna includes the following taxa: Stephanorhinus hundsheimensis, Equus apolloniensis, Bison sp. and Pontoceros ambiguus. The preliminary comparison of these faunal assemblages with the already known localities from the Mygdonia Basin and the wider region, combined with the geological data, suggest a Late Villafranchian age for TSR, chronologically intermediate between the localities Gerakarou-1 and Apollonia-1, while the fauna from PLN is indicative of a Latest Villafranchian age, similar to that of Apollonia-1. These new localities will enhance our understanding of the crucial time period of the earliest dispersal of hominins into Europe.