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Postcranial Osteology of the Basally Branching Hadrosauroid Dinosaur Tanius Sinensis from the Upper Cretaceous Wangshi Group of Shandong, China

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Postcranial Osteology of the Basally Branching Hadrosauroid Dinosaur Tanius Sinensis from the Upper Cretaceous Wangshi Group of Shandong, China Journal of Vertebrate Paleontology ISSN: (Print) (Online) Journal homepage: https://www.tandfonline.com/loi/ujvp20 Postcranial osteology of the basally branching hadrosauroid dinosaur Tanius sinensis from the Upper Cretaceous Wangshi Group of Shandong, China Niclas H. Borinder, Stephen F. Poropat, Nicolás E. Campione, Tomas Wigren & Benjamin P. Kear To cite this article: Niclas H. Borinder, Stephen F. Poropat, Nicolás E. Campione, Tomas Wigren & Benjamin P. Kear (2021): Postcranial osteology of the basally branching hadrosauroid dinosaur Taniussinensis from the Upper Cretaceous Wangshi Group of Shandong, China, Journal of Vertebrate Paleontology, DOI: 10.1080/02724634.2021.1914642 To link to this article: https://doi.org/10.1080/02724634.2021.1914642 © 2021 Niclas H. Borinder, Stephen F. View supplementary material Poropat, Nicolás E. Campione, Tomas Wigren, and Benjamin P. Kear Published online: 03 Jun 2021. Submit your article to this journal Article views: 557 View related articles View Crossmark data Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=ujvp20 Journal of Vertebrate Paleontology e1914642 (20 pages) Published with license by the Society of Vertebrate Paleontology DOI: 10.1080/02724634.2021.1914642 ARTICLE POSTCRANIAL OSTEOLOGY OF THE BASALLY BRANCHING HADROSAUROID DINOSAUR TANIUS SINENSIS FROM THE UPPER CRETACEOUS WANGSHI GROUP OF SHANDONG, CHINA NICLAS H. BORINDER,*,1,8 STEPHEN F. POROPAT, 2,3,4 NICOLÁS E. CAMPIONE, 5,6,7 TOMAS WIGREN,8 and BENJAMIN P. KEAR 8* 1Geological Survey of Sweden, Fångvallsgatan 4, SE-752 37 Uppsala, Sweden, [email protected]; 2Faculty of Science and Technology, Swinburne University of Technology, John St, Hawthorn, Victoria 3122, Australia; 3Australian Age of Dinosaurs Museum of Natural History, Lot 1 Dinosaur Drive, PO Box 408, Winton, Queensland 4735, Australia; 4Melbourne Museum, Museums Victoria, 11 Nicholson St, Carlton, Victoria 3053, Australia, [email protected], [email protected]; 5Palaeoscience Research Centre, University of New England, Armidale, New South Wales, 2350, Australia; 6Palaeobiology Programme, Department of Earth Sciences, Uppsala University, Villavägen 16, Uppsala SE-752 36, Sweden; 7Department of Organismal Biology, Uppsala University, Norbyvägen 18A, Uppsala SE-752 36, Sweden, [email protected]; 8Museum of Evolution, Uppsala University, Norbyvägen 16, Uppsala SE-752 36, Sweden, [email protected], [email protected] ABSTRACT—Tanius sinensis was one of the first dinosaur species to be named from China. It was established on a partial skeleton recovered by a joint Sino-Swedish expedition in 1923. The fossils were excavated from Upper Cretaceous strata of the Jiangjunding Formation (Wangshi Group) in Shandong Province, and although their discovery dates back almost 100 years, they have not been reassessed in detail since their initial description in 1929. This omission is critical because T. sinensis is now recognized as one of the stratigraphically youngest non-hadrosaurid hadrosauroid taxa. Here, we re- evaluate the postcranial osteology of T. sinensis as a prelude to an anatomical and phylogenetic revision of the species. We examined the holotype and all currently referred specimens of T. sinensis first-hand, and identified a unique postcranial character state combination incorporating tall dorsal neural spines, a reduced postacetabular ridge on the ilium, a fully enclosed flexor tunnel formed by the distal condyles of the femur, and a lunate proximal end on metatarsal III. Comparisons with other species of Tanius confirm that: (1) T. chingkankouensis is a nomen dubium erected on non- diagnostic composite material; (2) T. laiyangensis was established on indeterminate hadrosaurid remains that are not attributable to Tanius; and (3) the anecdotal assignments of Bactrosaurus prynadai and Tsintaosaurus spinorhinus to Tanius cannot be substantiated. Close inspection of the holotype caudal vertebra further reveals a possible healed bite trace consistent with a prey–predator interaction. Lastly, our calculated average body mass estimate for T. sinensis of between 2091–3533 kg suggests that it was one of the largest non-hadrosaurid hadrosauroids. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP Citation for this article: Borinder, N. H., S. F. Poropat, N. E. Campione, T. Wigren, and B. P. Kear. 2021. Postcranial osteology of the basally branching hadrosauroid dinosaur Tanius sinensis from the Upper Cretaceous Wangshi Group of Shandong, China. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2021.1914642 INTRODUCTION the hadrosauroid Tanius sinensis Wiman, 1929. Tanius sinensis The first dinosaur species named from China was ‘Trachodon’ was coined in honor of its discoverer, the Chinese geologist amurense Riabinin, 1925, which was later referred to the genus Tan Xichou, who was also often referred to (even by Tan Mandschurosaurus Riabinin, 1930, but is now regarded as a himself) using the westernized version of his name, H. C. T’an. nomen dubium (sensu Horner et al., 2004). The otherwise ear- The remains of T. sinensis were recovered from Upper Cretaceous liest documented valid Chinese dinosaur taxa were erected by (upper Campanian–lower Maastrichtian) sediments in the Jiang- the eminent Swedish paleontologist Carl Wiman in 1929. These junding Formation, which is a subunit of the Wangshi Group crop- included the titanosauriform sauropod Euhelopus zdanskyi ping out in the Shandong peninsular region of eastern China (Hu (Wiman, 1929) (see Mateer and McIntosh, 1985; Wilson and et al., 2001; Poropat and Kear, 2013b: fig. 1; Fig. 1). Upchurch, 2009; Poropat, 2013; Poropat and Kear, 2013a), and Since its initial description, the systematic relationships of T. sinensis have been debated. Wiman (1929) initially assigned T. sinensis to Hadrosauridae, a classification reiterated by Young (1958). Conversely, von Huene (1956) suggested that *Corresponding authors T. sinensis might constitute a separate family ‘Prohadrosauridae’, © 2021 Niclas H. Borinder, Stephen F. Poropat, Nicolás E. Campione, comprising both T. sinensis and Telmatosaurus transsylvanicus Tomas Wigren, and Benjamin P. Kear (Nopcsa, 1900). More recently, Weishampel and Horner This is an Open Access article distributed under the terms of the Crea- “ tive Commons Attribution License (http://creativecommons.org/licenses/ (1990:555) listed T. sinensis as: known from poor material that by/4.0/), which permits unrestricted use, distribution, and reproduction in has yet to yield phylogenetically significant information”. None- any medium, provided the original work is properly cited. theless, these authors interpreted the taxon as a basally Published online 03 Jun 2021 Borinder et al.—Postcranial osteology of Tanius sinensis (e1914642-2) FIGURE 1. Map of the Shandong peninsular region showing source localities for the Tanius sinensis holotype skeleton (PMU 24720/1–32) and referred elements (PMU 22481, PMU 22482, PMU 22483, PMU 24721). Scale bar equals 100 km. branching member of Hadrosauridae, and probably positioned literature-based comparative interpretations (Gilmore, 1933; amongst the sister lineages of Euhadrosauria (Lambeosaurinae Young, 1958; Buffetaut and Tong-Buffetaut, 1993) and phylogenies + Hadrosaurinae). This proposal was predicated upon the identi- (e.g., Sues and Averianov, 2009; Prieto-Márquez and Wagner, 2009; fication of various plesiomorphic cranial traits (see discussion in McDonald et al., 2010; Prieto-Márquez, 2010a; McDonald, 2012; Weishampel and Horner, 1990), yet the presence of tall neural Xing et al., 2014a; McDonald et al., 2017; Zhang et al., 2020). spines on the dorsal vertebrae and a robust, angular deltopec- Therefore, it is the purpose of this article to redress the shortfall toral crest on the humerus were both cited as features shared in knowledge via a comprehensive reassessment of the holotype with lambeosaurines. By contrast, the phylogenetic analyses of and all referred specimens of T. sinensis. Given the scale of such Sues and Averianov (2009) and Prieto-Márquez and Wagner a study, we have elected to present our data as a series of papers, (2009) both returned T. sinensis as a non-hadrosaurid hadrosaur- the first of which focuses on the postcranial skeleton and the oid, an affinity that has since been advocated by multiple later species-level taxonomy of Tanius. This will be followed by a synop- studies (e.g., McDonald et al., 2010; Prieto-Márquez, 2010a; sis of the historically associated Wangshi Group vertebrate assem- McDonald, 2012; Xing et al., 2014a). This topological placement blage, and lastly a reappraisal of the cranial osteology, together is significant because T. sinensis represents one of the geologically with a phylogenetic revision and contextual evaluation of non- youngest non-hadrosaurid hadrosauroids. Moreover, its fossils hadrosaurid hadrosauroid interrelationships. occur in close stratigraphic proximity to those of the lambeosaurine Institutional Abbreviations—AMNH, American Museum of (sensu Prieto-Márquez and Wagner, 2009) Tsintaosaurus Natural History, New York, U.S.A.; CMN, Canadian Museum spinorhinus Young, 1958, as well as the gigantic hadrosaurine of Nature, Ottawa, Canada; PMU, Paleontological Collections, (sensu Xing et al., 2014a, 2014b, 2017) Shantungosaurus Museum of Evolution, Uppsala University, Uppsala, Sweden. giganteus Hu, 1973, indicating that the replacement of non- hadrosaurid
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