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Systematics of New Information from the Chasmosaurus

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Systematics of New Information from the Chasmosaurus F1000Research 2015, 4:1468 Last updated: 16 MAY 2019 RESEARCH ARTICLE Systematics of Chasmosaurus - new information from the Peabody Museum skull, and the use of phylogenetic analysis for dinosaur alpha taxonomy [version 1; peer review: 1 not approved] Nicholas Longrich Department of Biology and Biochemistry and Milner Centre for Evolution, University of Bath, Bath, UK First published: 17 Dec 2015, 4:1468 ( Peer review discontinued v1 https://doi.org/10.12688/f1000research.7573.1) Latest published: 17 Dec 2015, 4:1468 ( Peer review at F1000Research is author-driven. https://doi.org/10.12688/f1000research.7573.1) Currently no reviewers are being invited. What does this mean? Abstract The horned dinosaur Chasmosaurus from the late Campanian Dinosaur Reviewer Status Park Formation of Alberta, is known from numerous skulls and skeletons, but over a century after its description, the taxonomy of the genus is Invited Reviewers controversial. Two species, Chasmosaurus belli and C. russelli, are 1 currently recognized, with a third species, C. irvinensis, recently placed in a new genus, Vagaceratops. Here, the Yale Chasmosaurus skull is version 1 described, and implications for Chasmosaurus systematics are explored. published report The Yale skull is intermediate between typical C. belli and C. irvinensis. C. 17 Dec 2015 belli-like features include large, triangular lateral epiparietals, large parietal fenestrae, and an emarginate parietal. Yet the skull also exhibits derived features of C. irvinensis, including a posteriorly inclined narial strut, brow 1 Jordan Mallon, Canadian Museum of Nature, horns replaced by rugose bosses, reduced parietal emargination, five pairs Ottawa, Canada of epiparietals, and epiparietals that fuse at their bases and hook forward Any reports and responses or comments on the over the frill. Specimen-level phylogenetic analysis provides a hypothesis of relationships upon which to base the taxonomy of Chasmosaurus. C. belli is article can be found at the end of the article. paraphyletic with respect to C. irvinensis, and the Peabody skull is closer to C. irvinensis than to other C. belli. The holotype of C. russelli clusters with C. belli, making C. russelli a junior synonym of C. belli. Accordingly, Chasmosaurus can be divided into three species: C. belli, C. irvinensis, and C. priscus sp. nov, including specimens previously referred to C. russelli. The systematics of Chasmosaurus show how specimen level phylogeny can provide an evolutionary framework upon which to establish taxonomies. However, the resulting phylogenies may lead to paraphyletic species and genera. Keywords Chasmosaurinae , Dinosaur Park Formation , Campanian Page 1 of 30 F1000Research 2015, 4:1468 Last updated: 16 MAY 2019 Corresponding author: Nicholas Longrich ([email protected]) Competing interests: No competing interests. Grant information: NRL was funded by a Donnelley Postdoctoral Fellowship and the Yale Institute for Biospheric Studies. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Copyright: © 2015 Longrich N. This is an open access article distributed under the terms of the Creative Commons Attribution Licence, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. How to cite this article: Longrich N. Systematics of Chasmosaurus - new information from the Peabody Museum skull, and the use of phylogenetic analysis for dinosaur alpha taxonomy [version 1; peer review: 1 not approved] F1000Research 2015, 4:1468 ( https://doi.org/10.12688/f1000research.7573.1) First published: 17 Dec 2015, 4:1468 (https://doi.org/10.12688/f1000research.7573.1) Page 2 of 30 F1000Research 2015, 4:1468 Last updated: 16 MAY 2019 Institutional abbreviations first application of this approach to Chasmosaurus. First, however, AMNH, American Museum of Natural History, New York, a history of the genus is presented. New York; CMN, Canadian Museum of Nature, Ottawa, Ontario; NHMUK, Natural History Museum, London; ROM, Royal Ontario History of Chasmosaurus Museum, Toronto, Ontario; TMP, Royal Tyrrell Museum of The genus Chasmosaurus has a complex taxonomic history. Palaeontology, Drumheller, Alberta; UALVP, University of Alberta Lambe12 originally described Monoclonius belli on the basis of a Laboratory for Vertebrate Paleontology; YPM, Yale Peabody partial parietal (CMN 491). This species would then be referred Museum, New Haven, Connecticut. to Hatcher et al.18 to the now defunct Ceratops, as Ceratops belli. Later, with more complete fossils, Lambe showed that the animal Introduction was distinct from previously known ceratopsids and created a new The Chasmosaurinae is a diverse group of horned dinosaurs that genus, Protorosaurus19. After Protorosaurus proved to be preoccu- flourished during the Late Cretaceous of western North America1,2. pied, Lambe proposed Chasmosaurus20 as a replacement. And so, Chasmosaurines were especially diverse in the late Campanian. within the space of a dozen years, the species bore four different Different species are found up and down the coast of the Western genus names. Despite being fragmentary, the holotype is diagnos- Interior Seaway1,3–6, and multiple species frequently occur in a single tic, as the morphology of the posterior bar of the parietal is unique habitat. This diversity and a rich fossil record make chasmosaurines to this species5. an ideal system for studying dinosaur diversity and biogeography. Lambe named a second species in the 1902 paper, Monoclonius Among the best-known genera is the Chasmosaurus from the Dino- canadensis, which was later renamed Eoceratops canadensis21. saur Park Formation (DPF) of Alberta. Chasmosaurus is known This species was later placed in Chasmosaurus, as Chasmosaurus from numerous skulls and skeletons7–12 but despite abundant mate- canadensis22,23. As discussed below, it appears to be distinct from rial there is little agreement about how many species, or even gen- Chasmosaurus, but the type is nondiagnostic. era are present4–8,10,13. New specimens led to new species. In 1933, Lull described a skull Until recently, Chasmosaurus was split into three species: (ROM 839) as Chasmosaurus brevirostris11. The specimen has an Chasmosaurus belli, Chasmosaurus russelli, and Chasmosaurus unusually short rostrum, but this feature is typical of juvenile chas- irvinensis7. The three succeed each other in stratigraphic section, mosaurs24 and other features of C. brevirostris, including unfused with primitive species occurring at the base of the DPF and more epoccipitals and open cranial sutures24–26 indicate immaturity. As derived forms appearing higher up7, suggesting a lineage evolv- differences between C. brevirostris and C. belli likely reflect ontog- ing through time. Recently, however, it has been proposed that eny, C. brevirostris is considered a junior synonym of C. belli8,10,22. C. irvinensis represents a distinct lineage, and it was placed in a new genus, Vagaceratops4. That year, another skull, AMNH 5401, was described as Chas- mosaurus kaiseni27. AMNH 5401 has traditionally been referred Here, a skull from the Yale Peabody Museum of Natural History, to C. belli8,10, but the resemblance of the frill to that of C. belli is YPM 2016, is described. Until now, this specimen has only received the result of plaster reconstruction. As discussed below, the skull cursory description11 but its unique morphology may help sort out appears to be distinct from Chasmosaurus and has been referred to the systematics of Chasmosaurus. YPM 2016 has previously been a new genus, Mojoceratops5. referred to C. belli5,7,8,10,11, but closer examination shows apomor- phies that are absent in other specimens of this species. These Sternberg9 named a fourth species, C. russelli. The type (CMN features include five pairs of epiparietals, widely divergent lateral 8800) consists of a crushed skull. Sternberg interpreted the frill as rami of the parietal caudal bar, and a caudally inclined premaxillary having a V-shaped caudal margin; based on this and other char- narial strut. These characters occur, however, in the highly derived acters, he regarded CMN 8800 as a distinct species. Sternberg C. irvinensis7. referred three more specimens to C. russelli, with additional speci- mens referred by Godfrey and Holmes8, including CMN 2280, a This paper uses YPM 2016 as a starting point to examine Chas- largely complete skeleton. C. russelli has been regarded as a valid mosaurus’ systematics. Here, a revised taxonomy is proposed. species by all recent studies1,4,5,7,8,10,13. Rather than starting with a taxonomy and then identifying diagnos- tic characters, this paper uses a bottom-up approach to taxonomy. In 1989, a fifth species, Chasmosaurus mariscalensis, was named Specimen-level phylogenetic analysis was conducted first, then from bonebed material from the upper Campanian Aguja Forma- phylogeny is used as a framework to delimit and diagnose spe- tion of Texas22. Phylogenetic analyses fail to group this species and cies. Specimen-level phylogenies have previously been used to Chasmosaurus to the exclusion of other chasmosaurs3–6,28–30 how- examine the systematics of other groups, including apatosaurine ever, and so it was referred to a separate genus, as Agujaceratops sauropods14, thescelosaurid ornithopods15, and the leptoceratopsid mariscalensis31. Montanoceratops16. Most recently, Scannella et al.17 have used specimen-level phylogenetic analysis to examine the systematics
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