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Postcranial anatomy of Besanosaurus leptorhynchus (Reptilia: Ichthyosauria) from the Middle Triassic Besano Formation of Monte San Giorgio (Italy/Switzerland), with implications for reconstructing the swimming styles of Triassic ichthyosaurs.

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Peer-reviewed

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Abstract

UNLABELLED: Besanosaurus leptorhynchus Dal Sasso & Pinna, 1996 was originally described on the basis of a single complete fossil specimen excavated near Besano (Italy). However, a recent taxonomic revision and re-examination of the cranial osteology allowed for the assignment of five additional specimens to the taxon. Here, we analyse, describe and discuss the postcranial anatomy of Besanosaurus leptorhynchus in detail. The size of the specimens examined herein ranged from slightly more than one meter to eight meters. Overall, several diagnostic character states for this taxon are proposed, demonstrating a mosaic of plesiomorphic and derived features. This is best exemplified by the limbs, which show very rounded elements in the forelimbs, and pedal phalanges with retained rudimentary shafts. We suggest that the widely spaced phalanges in the forefins of Besanosaurus leptorhynchus were embedded in a fibrocartilage-rich connective tissue, like in modern cetaceans. We also review the similarities of Besanosaurus with Pessopteryx and Pessosaurus, allowing us to conclude that Besanosaurus is not a junior synonym of either of the two taxa. Lastly, to test the swimming capabilities of Besanosaurus leptorhynchus, we expanded on a previously published study focussing on reconstructing the swimming styles of ichthyosaurs. Besanosaurus leptorhynchus was found to possess a peculiar locomotory mode, somewhat intermediate between anguilliform swimmers, such as Cymbospondylus and Utatsusaurus, and some shastasaur-grade (e.g., Guizhouichthyosaurus) and early-diverging euichthyosaurian (e.g., Californosaurus) ichthyosaurs. Based on our results, we furthermore suggest that mixosaurids acquired their characteristic body profile (dorsal fin and forefins that are distinctly enlarged compared to the hindfins) independently and convergently to the one that later appeared in Parvipelvia. Moreover, the different swimming styles inferred for Cymbospondylus, Mixosauridae, and Besanosaurus strengthen the earlier hypothesis of niche partitioning among these three distinct ichthyosaur taxa from the Besano Formation. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1186/s13358-024-00330-9.

Description

Acknowledgements: We thank the volunteers of the former “Gruppo paleontologico di Besano”, who unearthed the holotype of Besanosaurus leptorhynchus, and many other exceptional fossils. We also thank D. Affer, F. Fogliazza and L. Magnoni (MSNM), C. Egli (PIMUZ), and B. Riss (PIMUZ) for preparation of the specimens; M. Auditore for anatomical drawings; A. Ciaffi for the artwork; L. Forzenigo, C. Bonelli, and G. Terribile (Fondazione IRCCS “Cà Granda” Ospedale Maggiore Policlinico di Milano) for CT scanning the holotype of Besanosaurus leptorhynchus; the “Ministero della Cultura” and the “Soprintendenza Archeologia, Belle Arti e Paesaggio delle province di Como, Lecco, Monza-Brianza, Pavia, Sondrio e Varese” for permissions. For 3D-scanning the holotype of Besanosaurus leptorhynchus, we thank M. Merella and F. Nobile (Università di Pisa). For access to key specimens in museum collections, we thank C. Klug (PIMUZ), B. P. Kear (PMU), E. E. Maxwell (SMNS) and R. B. Hauff (Urweltmuseum Hauff, Holzmaden). We also thank M. Auditore, M. Balini, A. Crasti, B. P. Kear, S. Maganuco, and G. Teruzzi, for helpful discussions and the Willi Hennig Society for making TNT 1.5 freely available. Finally, we thank editor Daniel Marty and three anonymous reviewers for their feedback, which helped us to improve our manuscript.

Journal Title

Swiss J Palaeontol

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Journal ISSN

1664-2376
1664-2384

Volume Title

143

Publisher

Springer Science and Business Media LLC

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Except where otherwised noted, this item's license is described as http://creativecommons.org/licenses/by/4.0/
Sponsorship
National Natural Science Foundation of China (42202006)
Bekker Programme of the Polish National Agency for Academic Exchange (BPN/BEK/2022/1/00194)
Swiss National Science Foundation (31003A_179401)