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Postcranial Anatomy of Tanius Sinensis Wiman, 1929 (Dinosauria; Hadrosauroidea) Postkraniala Anatomin Hos Tanius Sinensis Wiman, 1929 (Dinosauria; Hadrosauroidea)

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Postcranial Anatomy of Tanius Sinensis Wiman, 1929 (Dinosauria; Hadrosauroidea) Postkraniala Anatomin Hos Tanius Sinensis Wiman, 1929 (Dinosauria; Hadrosauroidea) Examensarbete vid Institutionen för geovetenskaper Degree Project at the Department of Earth Sciences ISSN 1650-6553 Nr 320 Postcranial Anatomy of Tanius Sinensis Wiman, 1929 (Dinosauria; Hadrosauroidea) Postkraniala anatomin hos Tanius sinensis Wiman, 1929 (Dinosauria; Hadrosauroidea) Niclas H. Borinder INSTITUTIONEN FÖR GEOVETENSKAPER DEPARTMENT OF EARTH SCIENCES Examensarbete vid Institutionen för geovetenskaper Degree Project at the Department of Earth Sciences ISSN 1650-6553 Nr 320 Postcranial Anatomy of Tanius Sinensis Wiman, 1929 (Dinosauria; Hadrosauroidea) Postkraniala anatomin hos Tanius sinensis Wiman, 1929 (Dinosauria; Hadrosauroidea) Niclas H. Borinder ISSN 1650-6553 Copyright © Niclas H. Borinder and the Department of Earth Sciences, Uppsala University Published at Department of Earth Sciences, Uppsala University (www.geo.uu.se), Uppsala, 2015 Abstract Postcranial Anatomy of Tanius Sinensis Wiman, 1929 (Dinosauria; Hadrosauroidea) Niclas H. Borinder Tanius sinensis Wiman, 1929 was one of the first hadrosauroid or “duck-billed” taxa erected from China, indeed one of the very first non-avian dinosaur taxa to be erected based on material from the country. Since the original description by Wiman in 1929, the anatomy of T. sinensis has received relatively little attention in the literature since then. This is unfortunate given the importance of T. sinensis as a possible non-hadrosaurid hadrosauroid i.e. a member of Hadrosauroidea outside the family of Hadrosauridae, living in the Late Cretaceous, at a time when most non-hadrosaurid hadro- sauroids had become replaced by the members of Hadrosauridae. To gain a better understanding of the anatomy of T. sinensis and its phylogenetic relationships, the postcranial anatomy of it is redescribed. T. sinensis is found to have a mosaic of basal traits like strongly opisthocoelous cervical vertebrae, the proximal end of scapula being dorsoventrally wider than the distal end, the ratio between the proximodistal length of the metatarsal III and the mediolateral width of this element being greater than 4.5. Derived traits present in T. sinensis include curved dorsal surface of the scapula, arcuate fourth trochanter of the femur, cnemial crest of the tibia extending ventrally into the proximal half of the shaft, and the distal end of the fibula expanding into a club shape in lateral view. A potential autapomorphy is noted, being the caudal fusion of the medial and lateral condyles of the femur forming a completely enclosed “tunnel”. The body mass of the holotype of T. sinensis, is also estimated, and found to have been around 2950 kg. The phylogenetic analysis agrees with previous studies placing T. sinensis as a non-hadrosaurid hadrosauroid. Furthermore, in the phylo- genetic analysis, T. sinensis is recovered as non-hadrosaurid hadrosauroid, forming a clade with Bactrosaurus johnsoni Gilmore, 1933, which shares the following unambiguous synapomorphies; the width of the orbital margin of the jugal being almost equally wide to that of the infratemporal margin of the bone; the ratio between the mediolateral width of the skull roof across the postorbitals and that across the quadrate cotyli of the paired squamosals being more than 1.20. Keywords: Tanius, Jiangjunding, postcranial, hadrosauroid, phylogeny Degree Project E1 in Earth Science, 1GV025, 30 credits Supervisor: Nicolàs E. Campione Department of Earth Sciences, Uppsala University, Villavägen 16, SE-752 36 Uppsala (www.geo.uu.se) ISSN 1650-6553, Examensarbete vid Institutionen för geovetenskaper, No. 320, 2015 The whole document is available at www.diva-portal.org Populärvetenskaplig sammanfattning Postkraniala anatomin hos Tanius sinensis Wiman, 1929 (Dinosauria; Hadrosauroidea) Niclas H. Borinder Den här uppsatsen handlar om Tanius sinensis Wiman, 1929 som levde under Kritaperioden i Shan- dongprovinsen i nordöstra Kina. Tanius sinensis hörde till gruppen ”Anknäbbsdinosaurier” som ut- vecklades under början av Kritaperioden för mellan 130-100 miljoner år sedan. I slutet av Kritaperi- oden för ungefär 84 miljoner år sedan så blev de ”primitiva” ”anknäbbsdinosaurierna” bortträngda av de mer ”avancerade” ”anknäbbsdinosaurierna”. Tanius sinensis är viktig på så vis att den kan ha varit en ”primitiv” ”anknäbbsdinosaurie” som överlevde vid en tidpunkt när de flesta andra ”primitiva” ”anknäbbsdinsoaurier” hade trängts undan. För att få en bättre bild av T. sinensis anatomiska känne- tecken och en bättre bild av dess släktskapsförhållanden, så ombeskrivs anatomin hos den del av skelettet som inte omfattar kraniet. En släktskapsanalys görs också baserat på de kännetecken som jag själv och tidigare forskare har observerat i skelettet. De anatomiska observationerna avslöjar en mosaik av ”primitiva” och mer ”avancerade” karaktärer, som tillsammans med släktskapsanalysen pekar på att T. sinensis var en sent överlevande ”primitiv” ”anknäbbsdinosaurie”. Nyckelord: Tanius, Jiangjunding, postcranial, hadrosauroid, fylogeni Examensarbete E1 i geovetenskap, 1GV025, 30 hp Handledare: Nicolàs E. Campione Institutionen för geovetenskaper, Uppsala universitet, Villavägen 16, 752 36 Uppsala (www.geo.uu.se) ISSN 1650-6553, Examensarbete vid Institutionen för geovetenskaper, Nr 320, 2015 Hela publikationen finns tillgänglig på www.diva-portal.org Table of Contents 1. Introduction ………………………………………………………………………………… 1 2. Aim …………………………………………………………………………………………… 2 3. Background ………………………………………………………………………………… 3 3.1 Geological context……………………………………………………………… 3 3.2 Paleoenvironmental context…………………………………………………… 5 3.3 Historical context ……………………………………………………………… 5 4. Methodology ………………………………………………………………………………… 6 4.1 Osteological comparisons …………………………………………………… 6 5. Results ……………………………………………………………………………………… 7 5.1 Systematic palaeontology ……………………………………………………… 7 5.2 Holotype ...……………………………………………………………………… 7 5.3 Locality and horizon …………………………………………………………… 8 5.4 Diagnosis ..……………………………………………………………………… 8 5.5 Description .….….….…………………………………………………………… 9 5.5.1 Axial skeleton…………………………………………………………… 9 5.5.2 Appendicular skeleton …………………………………………………… 13 5.6 Referred material PMUR235 and PMUR239 ……………………………… 18 5.7 Locality and horizon …………………………………………………………… 18 5.8 Comments ……………………………………………………………………… 18 5.9 Description……………………………………………………………………… 18 5.10 Referred material PMUR271 and PMUR27………………………………… 19 5.11 Locality and horizon …………………………………………………………… 19 5.12 Comments ……………………………………………………………………… 19 5.13 Description ……………………………………………………………………… 19 5.14 Referred material small vertebra without registration number …………… 20 5.15 Locality and horizon …………………………………………………………… 20 5.16 Description ……………………………………………………………………… 20 5.17 Referred material, two fragmentary vertebrae centra ……………………… 21 5.18 Locality and horizon …………………………………………………………… 21 5.19 Description……………………………………………………………………… 22 5.20 Referred material, two pubic bones PMUR245, and one without number 22 5.21 Locality and horizon …………………………………………………………… 22 5.22 Description ……………………………………………………………………… 22 5.23 Comparisons between the two humeri ……………………………………… 22 6. Phylogenetic systematics…………………………………………………………………… 23 6.1 Methods and matherials ……………………………………………………… 23 6.2 Results of phylogenetic tree ………………………………………………… 24 7. Discussion …………………………………………………………………………………… 27 7.1 Postcranial character complex in Tanius sinensis ………………………… 27 7.2 Paleobiological implications ………………………………………………… 30 7.3 The size of Tanius sinensis …………………………………………………… 31 8. Conclusions ………………………………………………………………………………… 32 9. Acknowledgements ……………………………………………………………………… 32 10. References ………………………………………………………………………………… 34 11. Plates ……………………………………………………………………………………… 43 Appendix 1 Axial skeleton measurements ……………………………………………… 109 Appendix 2 Appendicular skeleton measurements …………………………………… 111 1. Introduction Hadrosauroids, here defined as the as the least inclusive taxon containing Equijubus normani You et al 2003a and Parasaurolophus walkeri Parks, 1922, where a major group of ornithopods that first appears in the fossil record of the Early Cretaceous, with the earliest member E. normani (Norman 2004; Horner, Weishampel and Forster 2004) from China being found in deposits dated as Late Barremian-Albian, upper Lower Cretaceous (Tang et al. 2001). By the end of the Cretaceous they had spread to all continents except Africa and Australia, and especially in Laurasia, becoming one of the most diverse and abundant large-sized tetrapods of the Late Cretaceous (Horner, Weishampel and Forster 2004; Lund and Gates 2006). From the Santonian and onward, the hadrosauroids outside the family of Hadrosauridae became replaced by the members of the Hadrosauridae (Norman 2004; Prieto-Márquez 2010b; Campione et al. 2013; Xing et al. 2014b). Tanius sinensis Wiman, 1929, is a hadrosauroid from the Jiangjunding Formation of the Upper Cretaceous Wangshi Group, Shandong Province, China (Hu et al. 2001; Poropat and Kear 2013). Tanius sinensis is not the only hadrosauroid known from the Wangshi Group; Young (1958) described and named Tanius chingkankouensis and the lambeosaurine Tsintaosaurus spinorhinus. The validity of T. chingkankouensis is generally not accepted, and, for example, it is regarded as “doubtful” by Buffetaut (1995) or treated as synonymous with T. sinensis (Horner, Weishampel and Forster 2004; Lund and Gates 2006). Other hadrosauroid taxa from the Wangshi Group include the giant hadrosaurine Shantungosaurus giganteus Hu, 1973. Both Zhuchengosaurus maximus Zhao et al., 2007 and Huaxiaosaurus aigahtens Zhao, Wang and Li, 2011, have recently
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