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Metatarsals of a Large Caenagnathid Cf. Anzu Wyliei (Theropoda: Oviraptorosauria) from the Hell Creek Formation in South Dakota, U.S.A

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Metatarsals of a Large Caenagnathid Cf. Anzu Wyliei (Theropoda: Oviraptorosauria) from the Hell Creek Formation in South Dakota, U.S.A Canadian Journal of Earth Sciences Metatarsals of a large caenagnathid cf. Anzu wyliei (Theropoda: Oviraptorosauria) from the Hell Creek Formation in South Dakota, U.S.A. Journal: Canadian Journal of Earth Sciences Manuscript ID cjes-2020-0171.R1 Manuscript Type: Article Date Submitted by the 04-Jan-2021 Author: Complete List of Authors: Tsujimura, Kousuke; The University of Tokyo, Dept. of Earth and Planetary Science Manabe, Makoto; National Museum of Nature and Science, Geology & PalaeontologyDraft Chiba, Yumiko; The University of Tokyo, Dept.of Earth and Planetary Science Tsuihiji, Takanobu; National Museum of Nature and Science, Dept. of Geology and Paleontology Keyword: Dinosauria, Theropoda, Maastrichtian, Hell Creek Formation Is the invited manuscript for consideration in a Special Tribute to Dale Russell Issue? : © The Author(s) or their Institution(s) Page 1 of 36 Canadian Journal of Earth Sciences 1 Metatarsals of a large caenagnathid cf. Anzu wyliei (Theropoda: Oviraptorosauria) from the 2 Hell Creek Formation in South Dakota, U.S.A. 3 4 Kousuke Tsujimura1, Makoto Manabe2, Yumiko Chiba1 and Takanobu Tsuihiji1,2* 5 1Department of Earth and Planetary Science, The University of Tokyo, 7-3-1 Hongo, 6 Bunkyo-ku, Tokyo 113-0033 Japan 7 2Department of Geology and Paleontology, National Museum of Nature and Science, 4-1-1 8 Amakubo, Tsukuba, Ibaraki 305-0005 Japan 9 *Corresponding author. Email: [email protected] 10 11 12 13 14 15 16 17 18 19 1 © The Author(s) or their Institution(s) Canadian Journal of Earth Sciences Page 2 of 36 20 Abstract 21 Isolated metatarsals III and IV of a caenagnathid theropod likely referable to Anzu wyliei are 22 described from a locality of the Hell Creek Formation in northwestern South Dakota of the 23 U.S.A. These bones are missing from the holotype and only partial shafts have been 24 described for a specimen referable to this species. Accordingly, the present description adds 25 further anatomical information on this already well-known species of Caenagnathidae. The 26 present finding also demonstrates the significance of isolated or fragmentary specimens 27 found in multitaxic bone beds. 28 Draft 29 Key Words: Dinosauria, Theropoda, Caenagnathidae, Hell Creek Formation, Maastrichtian 30 31 32 33 34 35 36 37 38 2 © The Author(s) or their Institution(s) Page 3 of 36 Canadian Journal of Earth Sciences 39 40 41 Introduction 42 The caenagnathid theropod Anzu wyliei has been described from localities of the 43 Maastrichtian Hell Creek Formation in North and South Dakota (Lamanna et al. 2014). It is a 44 large-bodied member of Oviraptorosauria and, with the femoral length ranging from 500 to 45 525 mm and the total length estimated to be 3.5 m (Lamanna et al. 2014), is surpassed only 46 by Gigantoraptor erlianensis in body size for this clade. In addition, with most skeletal 47 elements known based on multiple specimensDraft referred to the species (Lamanna et al. 2014), 48 A. wyliei is one of the reference species among Caenagnathidae for its morphological 49 information. Among the bones missing from the holotype and referred specimen of this 50 species, however, are the weight-bearing metatarsals, i.e., metatarsals II through IV. 51 Although Cullen et al. (2020) recently described associated left metatarsals II through IV in a 52 large caenagnathid referred to A. wyliei (ROM VP 65884) found in the Hell Creek Formation 53 in Montana, these bones consist mostly of broken shafts and lack morphological information 54 on their proximal and distal articular ends. 55 In describing the skeletal morphology of Gigantoraptor erlianensis, Xu et al. (2007) 56 suggested a potentially high cursorial ability of Gigantoraptor erlianensis compared to 57 theropods of similar body sizes based on a relatively slender femur and longer tibia/fibular 3 © The Author(s) or their Institution(s) Canadian Journal of Earth Sciences Page 4 of 36 58 and metatarsals relative to the femur. Although this observation was based on a limited 59 amount of data, it would suggest possible locomotor aptation specific to Caenagnathidae. To 60 further examine this hypothesis, information on the hindlimb bones of other species 61 belonging to this clade is crucial. 62 In the present paper, we describe isolated metatarsals III and IV likely referable to 63 Anzu wyliei that were found in the Hell Creek Formation exposed in northwestern South 64 Dakota of the U.S.A. These specimens contribute new anatomical information on this 65 currently well-known species of Caenagnathidae. In addition, the importance of such isolated 66 or fragmentary bone bed specimens inDraft evaluating the local species diversity of dinosaurs 67 before the K/Pg extinction event is discussed. 68 69 Institutional abbreviations 70 CMN, Canadian Museum of Nature, Ottawa, Ontario, Canada; NSM, National Museum of 71 Nature and Science, Tsukuba, Ibaraki, Japan; ROM, Royal Ontario Museum, Toronto, 72 Ontario, Canada; TMP, Royal Tyrrell Museum of Palaeontology, Drumheller, Alberta, 73 Canada; ZPAL, Institute of Paleobiology, Polish Academy of Sciences, Warsaw, Poland. 74 75 Materials and methods 76 Measurements on the specimens described herein were made using a digital caliper 4 © The Author(s) or their Institution(s) Page 5 of 36 Canadian Journal of Earth Sciences 77 to the nearest hundredth mm and rounded to the nearest tenth millimeter (for those smaller 78 than 200 mm) or with a taper measure to the nearest mm (for those larger than 200 mm). 79 These measurements are summarized in Table 1. 80 The specimens were also subjected to X-ray computed tomographic (CT) imaging. 81 They were scanned on a Shimadzu inspeXio SMX-225 CT FPD HR scanner at the National 82 Museum of Nature and Science, Tsukuba, Ibaraki, Japan, with a slice thickness of 365 μm at 83 200 kV and 70 μA. Observations on image slices and three-dimensional (3D) visualization 84 were done using the software package Amira 6.4.0 (FEL, Hillsboro, Oreg.). 85 Draft 86 Geological setting 87 The specimens described herein were discovered from a single fossil locality in the Hell 88 Creek Formation called the Sandy Site (Triebold, 1997; Russell and Manabe, 2002; Bartlett, 89 2004) situated in Harding County in northwestern South Dakota, U.S.A (Fig. 1). The section 90 cropping out at this locality is located approximately 50 m below the boundary between the 91 Hell Creek and Fort Union formations and belongs to the megafloral zone HCII (Johnson, 92 2002; K. Johnson, pers. comm., cited in Bartlett, 2004). A recent survey, however, found that 93 the fossiliferous horizon is located only 30 m below the formational contact (T. R. Lyson, 94 pers. comm., October 2020). 5 © The Author(s) or their Institution(s) Canadian Journal of Earth Sciences Page 6 of 36 95 These bones described herein are among more than 3000 isolated or fragmentary 96 vertebrate skeletal elements collected from a single layer. This fossil-bearing layer is spread 97 out over an approximately 50-m-long outcrop. It overlies a tabular mudstone and varies in 98 thickness between approximately 1 and 1.5 m. The fossiliferous horizon consists of a 99 structure-less unit with layers of rip-up clasts of 1–2 cm in diameter and is overlain by a 100 coarse-grained, yellowish sandstone. The fossils were found on top of, and mixed in with, the 101 rip-up clast horizon (Bartlett, 2004). These sedimentological observations suggest a crevasse 102 splay deposit within a meandering river system (Fastovsky, 1987; Bartlett, 2004). Such 103 multitaxic bonebed localities are not uncommonDraft within the Hell Creek Formation and provide 104 important insights into the paleoecology of the latest Cretaceous North American ecosystems 105 (Lyson and Joyce, 2009; Horner et al., 2011; Lyson et al., 2019). 106 The specimens described herein, as well as other bones collected at this fossiliferous 107 unit by Triebold Paleontology, Inc., are housed at the National Museum of Nature and 108 Science, Japan (Russell and Manabe, 2002). In addition, specimens collected at this locality 109 after acquisition of the collection by the National Museum of Nature and Science are housed 110 at the Denver Museum of Nature & Science, Denver, Colorado, U.S.A. (with the locality 111 number DMNH Loc. 1492; T. R. Lyson, pers. comm., October 2020). 112 113 Systematic paleontology 6 © The Author(s) or their Institution(s) Page 7 of 36 Canadian Journal of Earth Sciences 114 Dinosauria Owen, 1842 115 Theropoda Marsh, 1881 116 Maniraptora Gauthier, 1986 117 Oviraptorosauria Barsbold, 1976 118 Caenagnathidae Sternberg, 1940 119 Cf. Anzu wyliei Lamanna, Sues, Schachner, and Lyson, 2014 120 121 Material—NSM PV 21086, isolated left metatarsal III, and NSM PV 21055, isolated right 122 metatarsal IV. Draft 123 Locality—Sandy Site, Harding County, South Dakota, U.S.A. 124 Formation/Age—Hell Creek Formation (Maastrichtian). 125 126 Description 127 Third metatarsal (NSM PV 21086; Figs. 2, 4A) 128 This specimen, especially the distal portion, is fractured. A small portion of the 129 proximal end is missing whereas the proximal end of the distal condyle is badly damaged and 130 partially restored with plaster. This specimen is identified a bone of the left side for the 131 following reasons. Firstly, Currie (1989), Currie et al. (2016) and Funston et al. (2016) 132 described that the articulation between the metatarsals III and II is more intimate than the one 7 © The Author(s) or their Institution(s) Canadian Journal of Earth Sciences Page 8 of 36 133 between the metatarsals III and IV in the caenagnathids Elmisaurus rarus and Citipes 134 elegans. As described below, the articular surface on the left side seen in dorsal or anterior 135 view is better developed than the one on the right.
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