Abstract
The taxonomy of Eurasian rhinoceroses has long been complicated by extensive homoplasy, incomplete fossil records, and shifting diagnostic criteria, particularly affecting the genera Dicerorhinus, Dihoplus, Pliorhinus, Rhinoceros, and Stephanorhinus. Rhinocerotid material from the Late Pliocene locality of Kvabebi (Georgia), previously referred to Pliorhinus miguelcrusafonti, is re-evaluated here using an expanded morphological dataset and phylogenetic analyses. The results consistently recover the Kvabebi rhinoceros as a distinct dicerorhinine lineage positioned close to the divergence between Coelodonta and Stephanorhinus. On the basis of its unique combination of cranial, dental, and postcranial characters, and its stable phylogenetic placement outside both Coelodonta and Stephanorhinus, the Kvabebi material is provisionally assigned to Scythicorhinus vekuai (Tsiskarishvili, 1987) gen. nov. et comb. nov. Although alternative placements within the early Coelodonta–Stephanorhinus radiation cannot be statistically rejected, a close relationship with Stephanorhinus miguelcrusafonti is strongly unsupported. Two fragmentary maxillae from Berehove (Crimea, Ukraine) are also examined and are tentatively referred to Scythicorhinus vekuai, pending the discovery of more diagnostic material. Discrete-character and morphometric analyses indicate that both the Kvabebi and Berehove rhinoceroses fall outside the observed morphological variation of Coelodonta and Stephanorhinus, supporting their distinction at the generic level, albeit provisionally given the limited and partially worn material. The results further support rejection of Pliorhinus as a valid genus. Overall, this study refines the systematic framework of late Neogene rhinocerotids and underscores the importance of expanded comparative datasets for resolving evolutionary relationships within this highly homoplastic group.
Acknowledgements
The authors extend their sincere thanks to S. Dominici, L. Bellucci, and A. Savorelli for providing access to rhinoceros specimens housed in the Paleontology Section of the Museum of Natural History, University of Florence (Italy), enabling direct examination of the material. A.B. also thanks superintendent A. Betori (Soprintendenza Archeologia, Belle Arti e Paesaggio per le Province di Frosinone e Latina) for his kind support. P.P.A.M. was supported in the present study by the PAULMAZZARICATEN26—Mazza P. Fondo Ateneo 2026 MUR (Italian Ministry of Universities and Research) grant, as well as by the Institutional Recognition (Art. 4, para. 2 of Call 2025) awarded by the Ministry of Foreign Affairs and International Cooperation (MAECI) to project Arc-3089. We further wish to thank three anonymous reviewers for their insightful comments and helpful suggestions that significantly improved the manuscript.
Funding
This research was partly supported by the PAULMAZZARICATEN24 – Mazza P. Fondo Ateneo 2024 grant from MIUR (Italian Ministry of Education, Universities, and Research), which funded P.P.A.M.’s contribution. Additional support was provided by the National Academy of Sciences of Ukraine (0125U000927, 0124U000572) for O.K. and A.D. as well as by subsidy 2025; (501); MPK 2599280000 ZP; discipline 73 awarded to K.S. and U.R.S. Computational analyses were partly conducted using resources provided by the Wrocław Center for Networking and Supercomputing (grant No. 307 and 442).
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Kovalchuk, O., Borrani, A., Mackiewicz, P. et al. Scythicorhinus vekuai gen. nov. et comb. nov. (Mammalia, Rhinocerotidae) from the Pliocene of Georgia and its implications for the early evolution of Coelodonta and Stephanorhinus.
Sci Rep (2026). https://doi.org/10.1038/s41598-026-57113-0
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DOI: https://doi.org/10.1038/s41598-026-57113-0
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