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Dental Anatomy of the Apex Predator Sinraptor Dongi (Theropoda: Allosauroidea) from the Late Jurassic of China

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Dental Anatomy of the Apex Predator Sinraptor Dongi (Theropoda: Allosauroidea) from the Late Jurassic of China Canadian Journal of Earth Sciences Dental anatomy of the apex predator Sinraptor dongi (Theropoda: Allosauroidea) from the Late Jurassic of China Journal: Canadian Journal of Earth Sciences Manuscript ID cjes-2019-0231.R1 Manuscript Type: Article Date Submitted by the 06-Feb-2020 Author: Complete List of Authors: Hendrickx, Christophe; University of the Witwatersrand, Evolutionary Studies Institute Stiegler, Josef; The George Washington University, Department of Biological Sciences Currie, PhilipDraft J.; University of Alberta, Biological Sciences Han, Fenglu; China University of Geosciences, 4School of Earth Sciences Xu, Xing; Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology; CAS Center for Excellence in Life and Paleoenvironment Choiniere, Jonah; University of the Witwatersrand, Evolutionary Studies Institute Wu, Xiao-Chun; Canadian Museum of Nature Keyword: tooth, crown, denticle, dinosaur, theropod Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue? : https://mc06.manuscriptcentral.com/cjes-pubs Page 1 of 64 Canadian Journal of Earth Sciences 1 Dental anatomy of the apex predator Sinraptor dongi (Theropoda: 2 Allosauroidea) from the Late Jurassic of China 3 4 Running title: The dentition of Sinraptor. 5 6 CHRISTOPHE HENDRICKX1*, JOSEF STIEGLER2, PHILIP J. CURRIE3, FENGLU 7 HAN4, XING XU5, 6, JONAH N. CHOINIERE1, and XIAO-CHUN WU7 8 9 1Evolutionary Studies Institute, University of the Witwatersrand, Private Bag 3, Johannesburg 10 WITS 2050, South Africa. [Current address: Unidad Ejecutora Lillo, CONICET-Fundación 11 Miguel Lillo, Miguel Lillo 251, San Miguel de Tucumán 4000, Tucumán, Argentina]. 12 2Department of Biological Sciences, TheDraft George Washington University, Washington, DC 13 20052, USA. 14 3Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9, 15 Canada. 16 4School of Earth Sciences, China University of Geosciences, Wuhan, PR China. 17 5Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate 18 Paleontology and Paleoanthropology, Chinese Academy of Sciences, 142 Xiwai Street, 19 Beijing 100044, China. 20 6CAS Center for Excellence in Life and Paleoenvironment, 142 Xiwai Street, Beijing, 21 100044, China. 22 7Canadian Museum of Nature, PO Box 3443 STN ‘D’, Ottawa, ON K1P 6P4, Canada. 23 24 Emails: [email protected]. *Corresponding author 1 https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 2 of 64 25 Research Highlights 26 A thorough description of the dentition of the theropod dinosaur and apex predator Sinraptor 27 dongi from the Shishugou Formation (Oxfordian, Upper Jurassic) of China is provided for the 28 first time. The dental morphology of Sinraptor shows clear adaptations towards a predatory 29 lifestyle including crowns capable of enduring bone-to-bone contact and inflicting wide cuts 30 and open wounds. This study will facilitate the comparison and assignment of isolated 31 theropod teeth to metriacanthosaurid theropods and ultimately help to understand how the 32 different-sized theropods from the Shishugou Formation divided the carnivorous 33 ecomorphospace. Draft 2 https://mc06.manuscriptcentral.com/cjes-pubs Page 3 of 64 Canadian Journal of Earth Sciences 34 Abstract 35 The gross dental morphology of the holotype of the theropod Sinraptor dongi from the 36 Jurassic Shishugou Formation of China is comprehensively described. We highlight a 37 combination of dental features that appear to be restricted to Sinraptor: i) crowns with 38 denticulated mesial and distal carinae extending from the root, and an irregular surface texture 39 on the enamel; ii) D- to salinon-shaped cross-sectional outline at the crown-base in 40 mesialmost teeth; iii) mesial crowns with mesial carinae spiraling mesiolingually and 41 lingually positioned longitudinal groove adjacent to the mesial carina; and iv) labiolingually 42 compressed lateral teeth with weakly labially deflected distal carinae, flat to concave 43 basocentral surfaces of the labial margins of the crowns, and horizontally elongated distal 44 denticles showing short to well-developed interdenticular sulci. Using cladistic, multivariate, 45 discriminant and cluster analyses, we demonstrateDraft that the dentition of Sinraptor is relatively 46 similar to that of ceratosaurids, megalosauroids and other allosauroids, and particularly close 47 to that of Allosaurus. The dental anatomy of Sinraptor and Allosaurus, which mainly differs 48 in the labiolingual compression of the lateral crowns and the number of premaxillary teeth, 49 shows adaptations towards a predatory lifestyle, including premaxillary teeth capable of 50 enduring bone-to-bone contact, and crowns with widely separated mesial and distal carinae 51 capable of inflicting widely open wounds. 52 53 Keywords: Tooth, crown, denticle, dinosaur, theropod 3 https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page 4 of 64 54 Introduction 55 Sinraptor dongi Currie and Zhao 1993 was one of the apex terrestrial carnivores in the 56 vertebrate ecosystem of the ‘upper’ Shishugou Formation, Xinjiang Uygur Autonomous 57 Region, Xinjiang Province, north-western China (Currie and Zhao 1993, Xu and Clark 2008; 58 Figure 1A.3). The formation has yielded one of most diverse dinosaur faunas from the 59 Jurassic of Asia and is currently divided into lower, middle and upper beds (Choiniere et al. 60 2014a, 2014b, Moore et al. 2018). The upper section of the Shishugou Formation (here 61 referred as the ‘upper’ Shishugou Formation), which includes the upper part of the middle 62 beds and the upper beds (Figure 1A), is the richest taxonomically, with a dozen dinosaur 63 species recorded hitherto (Currie and Zhao 1993, Eberth et al. 2010, Choiniere et al. 2014a, 64 2014b, Qin et al. 2019). The fauna is dominated by herbivorous dinosaurs belonging to five 65 clades (Figure 1A) – SauropodomorphaDraft with Mamenchisaurus sinocanadorum Russell and 66 Zheng 1993, Tienshanosaurus chitaiensis Young 1937, and Bellusaurus sui Dong 1990 (and 67 see Dong 1992, Moore et al. 2018), Theropoda with Limusaurus inextricabilis Xu et al. 2009 68 (and see Wang et al. 2016, 2017), Ornithischia with Gongbusaurus wucaiwanensis Dong 69 1989, Stegosauria with Jiangjunosaurus junggarensis Jia et al. 2007, and Marginocephalia 70 with Yinlong downsi Xu et al. 2006b (and see Han et al. 2016, 2018) and Hualianceratops 71 wucaiwanensis Han et al. 2015. Besides the large-bodied predator Sinraptor dongi (i.e., ~7m 72 in length; Tanke and Currie 1998), carnivorous theropods are represented by five small to 73 medium-sized dinosaurs (Figure 1A), the basally branching coelurosaur Zuolong salleei 74 Choiniere et al. 2010a, the proceratosaurid tyrannosauroid Guanlong wucaii Xu et al. 2006a, 75 and the early alvarezsauroids Aorun zhaoi Choiniere et al. 2014a (and see Xu et al. 2018), 76 Haplocheirus sollers Choiniere et al. 2010b (and see Choiniere et al. 2014b) and 77 Shishugounykus inexpectus Qin et al. 2019. Stratigraphically, the Shishugou Formation 78 correlates with the upper Callovian or lower Oxfordian to the upper Oxfordian, an assessment 4 https://mc06.manuscriptcentral.com/cjes-pubs Page 5 of 64 Canadian Journal of Earth Sciences 79 based on 40Ar/39Ar radiometric ages (Eberth et al. 2010, Choiniere et al. 2014b). Because 80 confident radiometric ages are not yet available for the underlying Xishanyao Formation or its 81 lateral equivalents, it is however possible that the Shishugou Formation is entirely Oxfordian. 82 Sinraptor dongi is one of the best-known members of the ‘family-level’ clade 83 Metriacanthosauridae, a Laurasian radiation of allosauroid theropods closely related to 84 Allosaurus and extending from the Middle Jurassic to the Early Cretaceous of Europe and 85 Asia (Figure 2A; Currie and Zhao 1993, Carrano et al. 2012). S. dongi is presently known 86 from a nearly complete skull (Figure 1B-C, 2B) and a partial skeleton missing the tail and 87 several bones of the forearm (Figure 2C). The material was discovered by Haijun Wang in 88 September 1987, 25 km northeast of Jiangjunmiao, Junggar Basin, Xinjiang Province. The 89 skeleton was unearthed by a team of Chinese and Canadian paleontologists in 1987 and 1988 90 during the Sino-Canadian expedition knownDraft as the ‘Dinosaur Project’ (Dong et al. 1988, 91 Currie 1991, Dong 1993). S. dongi is characterized by a postorbital horn, an enlarged lateral 92 temporal fenestra and a highly pneumatized internal naris, maxilla and postpalatine (Figure 93 2B; Carrano et al., 2012; Currie & Zhao, 1993). 94 Whereas the cranial and postcranial material of S. dongi has received a comprehensive 95 description from Currie and Zhao (1993), the dentition morphology has been surprisingly 96 succinctly described, with information on the mesial dentition, crown ornamentation, carinae 97 and denticle morphology being omitted. The teeth of this taxon are, nonetheless, particularly 98 well-preserved, and most of them are still positioned in their alveoli (Figure 1A). The scarcity 99 of information available on the dentition of Sinraptor, and metriacanthosaurids in general, led 100 several authors (e.g., Han et al., 2011; Xu & Clark, 2008) to tentatively assign isolated 101 theropod teeth from the Shishugou Formation either to more inclusive clades (i.e., Tetanurae, 102 Tyrannosauroidea) or to metriacanthosaurids. 5 https://mc06.manuscriptcentral.com/cjes-pubs Canadian Journal of Earth Sciences Page
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