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Dinosaur Park Formation Canadian Journal of Earth Sciences Anatomical, morphometric, and stratigraphic analyses of theropod biodiversity in the Upper Cretaceous (Campanian) Dinosaur Park Formation Journal: Canadian Journal of Earth Sciences Manuscript ID cjes-2020-0145.R2 Manuscript Type: Article Date Submitted by the 13-Dec-2020 Author: Complete List of Authors: Cullen, Thomas; North Carolina Museum of Natural Sciences, Paleontology; North Carolina State University, Biological Sciences; Field Museum of Natural History, Nagaunee Integrative Research Center Zanno, Lindsay;Draft North Carolina Museum of Natural Sciences, Paleontology; North Carolina State University, Biological Sciences Larson, Derek W; Philip J Currie Dinosaur Museum, Todd, Erinn; University of Toronto, Ecology and Evolutionary Biology Currie, Philip J.; University of Alberta, Biological Sciences Evans, David; Royal Ontario Museum, Department of Natural History; University of Toronto, Ecology and Evolutionary Biology theropod, dinosaur, biodiversity, Dinosaur Park Formation, troodontid, Keyword: biostratigraphy Is the invited manuscript for consideration in a Special Tribute to Dale Russell Issue? : © The Author(s) or their Institution(s) Page 1 of 52 Canadian Journal of Earth Sciences Anatomical, morphometric, and stratigraphic analyses of theropod biodiversity in the Upper Cretaceous (Campanian) Dinosaur Park Formation THOMAS M. CULLEN*,1,2,3, LINDSAY ZANNO1,2,3, DEREK W. LARSON4,5, ERINN TODD6, PHILIP J. CURRIE7, and DAVID C. EVANS6,8 1Paleontology, North Carolina Museum of Natural Sciences, 11W. Jones, St. Raleigh, NC, 27601, USA 2Department of Biological Sciences, North Carolina State University, 100 Brooks Ave., Raleigh, NC, 27607, USA 3Nagaunee Integrative Research Center, Field Museum of Natural History, 1400 S Lake Shore Dr., Chicago, IL, 60605, USA. 4Collections Care and Conservation, Royal BC Museum, 675 Belleville St, Victoria, BC, V8V 9W2, Canada. 5Philip J. Currie Dinosaur Museum, Box 328 9301-112 Avenue, Wembley, AB T0H 3S0. 6Department of Ecology and Evolutionary Biology, University of Toronto, 25 Willcocks Street, Toronto, ON M5S 3B2, Canada. 7Department of Biological Sciences, University of Alberta, Edmonton, AB, T6G 2E9 8Department of Natural History, Royal Ontario Museum, 100 Queen’s Park, Toronto, ON M5S 2C6, Canada. *Corresponding author: [email protected]; [email protected] Draft ABSTRACT— The Dinosaur Park Formation (DPF) of Alberta, Canada has produced one of the most diverse dinosaur faunas, with the record favouring large-bodied taxa, in terms of number and completeness of skeletons. Although small theropods are well documented in the assemblage, taxonomic assessments are frequently based on isolated, fragmentary skeletal elements. Here we reassess DPF theropod biodiversity using morphological comparisons, high- resolution biostratigraphy, and morphometric analyses, with a focus on specimens/taxa originally described from isolated material. In addition to clarifying taxic diversity, we test if DPF theropods preserve faunal zonation/turnover patterns similar to those previously documented for megaherbivores. Frontal bones referred to a therizinosaur (cf. Erlikosaurus), representing among the only skeletal record of the group from the Campanian-Maastrichtian (83-66 Ma) fossil record of North America, plot most closely to troodontids in morphospace, distinct from non-DPF therizinosaurs, a placement supported by a suite of troodontid anatomical frontal characters. Postcranial material referred to cf. Erlikosaurus in North America is also reviewed and found © The Author(s) or their Institution(s) Canadian Journal of Earth Sciences Page 2 of 52 most similar in morphology to caenagnathids, rather than therizinosaurs. Among troodontids, we document considerable morphospace and biostratigraphic overlap between Stenonychosaurus and the recently described Latenivenatrix, as well as a variable distribution of putatively autapomorphic characters, calling the validity of the latter taxon into question. Biostratigraphically, there are no broad-scale patterns of faunal zonation similar to those previously documented in ornithischians from the DPF, with many theropods ranging throughout much of the formation and overlapping extensively, possibly reflecting a lack of sensitivity to environmental changes, or other cryptic ecological or evolutionary factors. KEYWORDS: theropod; biodiversity; biostratigraphy; Dinosaur Park Formation; troodontid; dinosaur Draft INTRODUCTION A major component of the work of the late Dr. Dale Russell focused on the palaeontology of theropods in Western Canada. This research included a series of systematic assessments of major theropod groups, such as tyrannosaurs, including the description of Daspletosaurus torosus (Russell 1970), ornithomimosaurs, including the description of Dromiceiomimus brevitertius (Russell 1972), a detailed review and re-description of the holotype of the dromaeosaurid Dromaeosaurus albertensis (Colbert and Russell 1969), and a broad review of therizinosaurs (Russell 1997). In addition, and of particular note, was Russell’s work on troodontids from what is now the Dinosaur Park Formation (at the time a part of the Oldman Formation, later split into a separate unit by Eberth and Hamblin [1993]), which included the description of the most © The Author(s) or their Institution(s) Page 3 of 52 Canadian Journal of Earth Sciences complete specimen of Stenonychosaurus inequalis (Russell 1969) and subsequent investigations into the palaeobiology of that taxon alongside discussions of troodontid speculative evolution and intelligence (Russell and Séguin 1982). In this contribution, we build on Russell’s foundational work by reviewing and addressing a series of outstanding questions in theropod biodiversity, taxonomy, and biostratigraphy in the Late Cretaceous of North America, with a specific focus on the Dinosaur Park Formation. The Dinosaur Park Formation (or DPF) of Alberta has produced some of the most diverse dinosaur assemblages in the fossil record (Béland and Russell, 1978; Currie and Koppelhus, 2005; Brown et al., 2013) including a relatively high diversity of small-bodied theropods such as dromaeosaurs, troodontids, ornithomimids,Draft and oviraptorosaurs (Currie, 2005). Unfortunately, some theropod groups are still known from a relative dearth of skeletal materials (Brown et al., 2013). As a result, the number of named theropod taxa in the DPF has often fluctuated, with taxonomic assessments frequently based on isolated, fragmentary, and/or non-overlapping remains. This has impacted some theropod groups more than others. Recent work and the discovery of multiple new skeletons have yielded important insights into the biodiversity and palaeobiology of several of these theropod groups in the DPF. Among ornithomimids, new discoveries and re-assessments of previously collected specimens have led to the recognition of a smaller-bodied taxon, Rativates evadens (McFeeters et al 2016), distinct from the previously recognized DPF ornithomimids Struthiomimus altus and Ornithomimus edmontonicus (in the latter case, following Makovicky et al [2004]’s suggested synonymization of Dromiceiomimus samueli with O. edmontonicus). Additionally, the presence © The Author(s) or their Institution(s) Canadian Journal of Earth Sciences Page 4 of 52 of a currently unnamed large-bodied taxon (Longrich 2008) was hypothesized based on isolated caudal vertebrae, a frontal, and unguals, though the degree to which at least the morphology of the latter represents autapomorphic characters as opposed to individual, positional, or intraspecific variation has been questioned (Cullen et al 2013, McFeeters et al 2018a, McFeeters et al 2018b). Lastly, a partial skeleton and several isolated elements were recently re-examined and hypothesized to represent evidence of the Asian taxon Qiupalong in the DPF (McFeeters et al 2017). Similar to the ornithomimids, caenagnathids in the DPF have undergone a series of recent taxonomic revisions based on the discovery of new material and re-assessment of existing specimens. Throughout the last few decades,Draft Chirostenotes pergracilis and Caenagnathus collinsi have been known from what is now the DPF, with recent work referring additional specimens to the latter (Funston et al 2015). Of greater contention has been the taxonomic identification of other potential caenagnathid taxa within the formation, with this issue recently reviewed by Funston (2020), wherein the taxon Citipes elegans was described. It should also be noted that some authors have raised concerns regarding the use of size classes and grouping of non-overlapping materials in some of these referrals and descriptions (Lamanna et al 2014, Cullen et al 2020b). Dromaeosaurids represent an additional small theropod group of the DPF which are known primarily from isolated materials, but for which several well-preserved specimens exist. The earliest dromaeosaurid described from what is now Dinosaur Provincial Park, and paradoxically now one of the more poorly known, is Dromaeosaurus albertensis (Matthew and Brown, 1992; © The Author(s) or their Institution(s) Page 5 of 52 Canadian Journal of Earth Sciences Colbert and Russell 1969). This taxon is primarily known from its holotype, a nearly complete skull, which unfortunately lacks precise biostratigraphic data (Colbert and Russell 1969). Considerably more common than D. albertensis in the DPF is Saurornitholestes langstoni, described by Sues (1978). This taxon is known from multiple partial skeletons,
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