Early Pleistocene rhinoceroses (Mammalia, Perissodactyla, Rhinocerotidae) from Northern Greece: biochronological and paleobiogeographic implications

Rhinoceroses were well-distributed across Southern Europe during the Early Pleistocene with abundant remains, but they are rather scarce in the Greek record and even less studied from a systematic point of view. In this article, we study a total of 124 specimens from eight fossil-bearing Lower Pleistocene localities of Northern Greece, spanning from ~ 2.3 to ~ 1.2 Ma. The localities of Tsiotra Vryssi and Krimni in the Mygdonia Basin provided the best material in terms of both number and preservation of specimens, whereas in the rest of the sites from Mygdonia, Drama and Neapolis-Grevena Basins less than ten specimens for each site are known. The present study documents the first occurrence in Greece of Stephanorhinus cf. S. hundsheimensis in the localities of Riza-1, Krimni-1, 3 and Tsiotra Vryssi, which suggests that the species may have entered earlier than the rest of the Europe. Additionally, the last occurrence of Stephanorhinus jeanvireti in Greece is recorded in Dafnero-3. The presence of Stephanorhinus etruscus is confirmed in several localities including Volax, Tsiotra Vryssi, Libakos and Apollonia-1, and indicates a chronological range from ~ 2.3 to ~ 1.2 Ma. At the beginning of this time frame both S. jeanvireti and S. etruscus co-occur in Greece, whereas the presence of both S. etruscus and Stephanorhinus cf. S. hundsheimensis at Tsiotra Vryssi testifies their first local co-occurrence.