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Cranial Anatomy of Csakvaromys bredai (Rodentia, Sciuridae, Xerinae) and Implications for Ground Squirrel Evolution and Systematics

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Abstract

We provide the first detailed description of the cranial anatomy of an early marmotine ground squirrel Csakvaromys (= Spermophilinus) bredai, based on a well-preserved facial skull and associated specimens from the early late Miocene (11.1–9.7 Ma) Grytsiv fauna in western Ukraine. Notable cranial features of C. bredai include a shallow, broad, and elongated facial skull with a straight dorsal profile, massive rostrum, deeply concave lateral lip of the infraorbital foramen, shortened infraorbital canal, large lacrimal foramen, an elongated anteromedial edge of the zygomatic arch extending onto the premaxilla, a broad and flat interorbital region with closed supraorbital foramina, and non-elevated supraorbital margins. Phylogenetic analysis of a 136 craniodental character matrix reconstructs Csakvaromys, together with Palaeosciurus, Protospermophilus, and Sciurotamias, within the subtribal clade Sciurotamiina. We provide the first reliable morphological synapomorphies for Xerinae, and suggest a revised definition for a monophyletic Sciurotamiina, which is supported by a large lacrimal foramen ventrally contributed by the maxilla, the facial tuberosity with a longitudinal crest for the superficial masseter, rudimentary ectopterygoid crest, and the presence of an expansive attachment area for the anterior fibers of the anterior deep masseter muscle. The craniofacial anatomy reveals that C. bredai was a frugivore with a predominantly hard-nut diet and arboreal or semi-arboreal habit.

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Data Availability

All data associated with this manuscript are presented in the supporting files and deposited on MorphoSpace. All fossil materials are housed at NMNHU-P.

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Acknowledgements

We are grateful to Olga V. Makarova (ZIN), Frank Zachos, Alexander Bibl, and Ursula B. Göhlich (NMW), Lyudmila S. Shevchenko (NMNHU-Z), Oleksandr M. Kovalchuk and Vadym O. Yanenko (NMNHU-P) for access to specimens in their care. We also thank Boris Kryštufek (Slovenian Museum of Natural History, Ljubljana, Slovenia) for providing the photographic images of the skull of Sciurotamias forresti. Finally, we thank the editor, John R. Wible, and reviewers, Isaac Casanovas-Vilar (Institut Català de Paleontologia Miquel Crusafont, Barcelona, Spain) and William W. Korth (Rochester Institute of Vertebrate Paleontology, New York, USA) for their constructive comments, which helped improve the manuscript. Financial support for this study was provided by Act 211 of the Government of the Russian Federation, contract No. 02.A03.21.0006, by a Grant to the Department of Science, Roma Tre University (MIUR-Italy, Dipartimenti di Eccellenza, Art.1 C, 314-327, L.232/2016), and partly by institutional support RVO67985831 of the Institute of Geology of the Czech Academy of Sciences.

Funding

Maxim V. Sinitsa was supported by Act 211 of the Government of the Russian Federation, [contract No. 02.A03.21.0006] and by a Grant to the Department of Science, Roma Tre University (MIUR-Italy, Dipartimenti di Eccellenza, Art.1 C, 314–327, L.232/2016); Stanislav Čermák was supported by institutional support RVO67985831 of the Institute of Geology of the Czech Academy of Sciences.

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Authors and Affiliations

  1. Institute of Natural Sciences and Mathematics, Ural Federal University, Mira 19, 620002, Ekaterinburg, Russia

    Maxim V. Sinitsa

  2. Dipartimento Di Scienze, Università Roma Tre, Largo S. L. Murialdo 1, 00146, Rome, Italy

    Maxim V. Sinitsa

  3. Department of Paleontology, National Museum of Natural History, National Academy of Sciences of Ukraine, Bohdan Khmelnytsky 15, 01030, Kiev, Ukraine

    Maxim V. Sinitsa

  4. Institute of Geology of the Czech Academy of Sciences, Rozvojová 269, Prague, 6, Czech Republic

    Stanislav Čermák

  5. Saint Petersburg State University, Universitetskaya Emb. 7/9, 199034, Saint Petersburg, Russia

    Lyudmila Yu. Kryuchkova

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  3. Lyudmila Yu. Kryuchkova
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Contributions

MVS conceived the study, conducted the phylogenetic analysis, virtual reconstructions of the specimens, prepared illustrations, and wrote the main part of the manuscript. SC prepared and wrote nomenclatural parts of the manuscript. LYK carried out the micro-CT scans and wrote the related parts of the manuscript. All authors discussed the results, edited the manuscript, and approved the final version for publication.

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Correspondence to Maxim V. Sinitsa.

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Sinitsa, M.V., Čermák, S. & Kryuchkova, L.Y. Cranial Anatomy of Csakvaromys bredai (Rodentia, Sciuridae, Xerinae) and Implications for Ground Squirrel Evolution and Systematics. J Mammal Evol 29, 149–189 (2022). https://doi.org/10.1007/s10914-021-09561-w

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