A New Oviraptorosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of Southern China and Its Paleoecological Implications
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A New Oviraptorosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of Southern China and Its Paleoecological Implications Junchang Lu¨ 1*, Laiping Yi2, Hui Zhong2, Xuefang Wei3 1 Institute of Geology, Chinese Academy of Geological Sciences, Beijing, China, 2 Ganzhou Museum of Natural History, Ganzhou, Jiangxi Province, China, 3 Cores and Samples Center of Land and Resources, China Geological Survey, Sanhe, Hebei Province, China Abstract A new oviraptorosaur Nankangia jiangxiensis gen. et sp. nov. is described on the basis of a partial postcranial skeleton with a partial lower jaw collected from the Upper Cretaceous Nanxiong Formation of Ganzhou, in Jiangxi Province of southern China. The new taxon is diagnosed by: (1) a mandibular symphysis that is not turned down; (2) neural spines of the cranial caudal vertebrae that are wider transversely than anteroposteriorly, forming a large posterior fossa with rugose central areas; (3) a femoral neck extending at an angle of about 90 to the shaft; and (4) a ratio of femur to tibia length of 0.95. Phylogenetic analysis recovers Nankangia as basal to the oviraptorid Yulong, but more derived than Caenagnathus, which also has a mandibular symphysis that is not turned down. The coexistence of Nankangia jiangxiensis, Ganzhousaurus nankangensis, Jiangxisaurus ganzhouensis, an unnamed oviraptorid from Nanxiong Basin and Banji long suggests that they occupied distinct ecological niches. Nankangia may have been more herbivorous than carnivorous. Citation: Lu¨ J, Yi L, Zhong H, Wei X (2013) A New Oviraptorosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of Southern China and Its Paleoecological Implications. PLoS ONE 8(11): e80557. doi:10.1371/journal.pone.0080557 Editor: Peter Dodson, University of Pennsylvania, United States of America Received April 29, 2013; Accepted September 24, 2013; Published November 27, 2013 Copyright: ß 2013 Lu¨ et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: J. Lu¨ was supported by Natural Science Foundation of China grants (41272022). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: The authors have declared that no competing interests exist. * E-mail: [email protected] Introduction To evaluate the systematic position of Nankangia within Oviraptorosauria, 182 characters were scored for 20 taxa (data Oviraptorids are specialized non-avian theropod dinosaurs that matrix based on [31]) for phylogenetic analysis. The analysis was have been reported from Mongolia [1–12] and China [13–23]. performed using the branch-and-bound search algorithm of The closely related caenagnathids are best known from North PAUP* 4.0 b10 [32]. Running the entire data matrix of 20 taxa America 31 [24–29]. Oviraptorosaurs are known from many (including Herrerasaurus, Archaeopteryx, and Velociraptor as outgroups, regions of China [13–18,21–23,30,31]. Recently, many oviraptor- plus most known oviraptorosaurs) and 182 characters produced osaur nests and eggs, along with at least ten oviraptorosaur two most parsimonious trees, each with a length of 370 steps skeletons buried with the eggs (All oviraptorosaur skeletons that (consistency index of 0.58, homoplasy index of 0.42, and retention have been published from the Ganzhou area thus far are index of 0.69). incomplete because, during quarrying of the rocks for building material, most specimens are destroyed) has made the Ganzhou Nomenclatural Acts district one of the most productive localities for oviraptorosaur fossils. Here we report on a new oviraptorosaur, Nankangia The electronic edition of this article conforms to the requirements jiangxiensis gen. et sp. nov., from the Upper Cretaceous Nanxiong of the amended International Code of Zoological Nomenclature, Formation of Nankang City, Jiangxi Province (Figure 1). Its and hence the new names contained herein are available under that discovery plays a key role in our understanding of the distribution Code from the electronic edition of this article. This published work and paleoecology of oviraptorosaurian dinosaurs. and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The Methods ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser This study was based on specimen GMNH F10003, housed in by appending the LSID to the prefix ‘‘http://zoobank.org/’’. The the Ganzhou Museum of Natural History, in Ganzhou City, LSID for this publication is: urn: lsid: zoobank.org: pub: C9F22590- Jiangxi Province. The specimen was donated to the Ganzhou 8438-427A-9E46-BE2A8CF7FCD2. The electronic edition of this Museum of Natural History three years ago by a local farmer. work was published in a journal with an ISSN, and has been Two of the authors (L. Yi and H. Zhong) from the Ganzhou archived and is available from the following digital repositories: Museum of Natural History joined the study of the specimen, and PubMed Central, LOCKSS. the senior author (J. Lu¨) obtained permission from the museum to access the collections. PLOS ONE | www.plosone.org 1 November 2013 | Volume 8 | Issue 11 | e80557 New Oviraptorosaur from Southern China Figure 1. Map of the fossil locality, near Ganzhou, Jiangxi Province, southern China. The solid pentagon represents the fossil site. doi:10.1371/journal.pone.0080557.g001 Results right pubis and most of the left pubis, complete right ischium and a partial left ischium, both femora, right tibia, and some dorsal ribs. Systematic Paleontology All specimens are housed at the Ganzhou Museum of Natural Theropoda Marsh, 1881 [33]. History, Ganzhou City of Jiangxi Province. Oviraptorosauria Barsbold, 1976 [2]. Type locality and horizon. Longling of Nankang, Ganzhou City; Nanxiong Formation (Upper Cretaceous) [34]. Nankangia gen. nov. (Figures 2, 3, 4, 5, 6, 7). Diagnosis. Nankangia is distinguished by the following com- Etymology. Nankangia refers to the Chinese administrative bination of characters: (1) the rostral end of the mandibular unit Nankang City in Jiangxi Province. symphyseal region is not downturned (shared with caenagnathids, Type species. Nankangia jiangxiensis gen. et sp. nov. urn:lsid: Incisivosaurus, Luoyanggia and Ganzhousaurus); (2) two infradiapophy- zoobank.org:act: D3C8151C-970B-4944-AA4C-E913E73F64D8. seal fossae on the ventral surface near the base of the transverse Diagnosis. As for the type and only known species. process of the dorsal vertebrae; (3) pneumatic fossae on the sacral Etymology. The specific name refers to Jiangxi Province, vertebrae slit-like; (4) neural spines of the cranial caudal vertebrae where the holotype site in Nankang City is located. wider transversely than anteroposteriorly, forming a large poste- Holotype. The holotype specimen (GMNH F10003) includes rior fossa with a rugose central area; (5) large fossa on the anterior a partial lower jaw, five dorsal vertebrae, one and one-half sacral surface (infraprezygapophyseal fossa) and another (infradiapophy- vertebrae, nine complete and two partial caudal vertebrae, both seal fossa) on the ventral surface of the base of the transverse scapulocoracoids, incomplete furcula, a nearly complete right process of the cranial caudals; (6) femur longer than ilium (shared humerus, complete right ilium, most of the left ilium, complete with Yulong and Khaan); (7) ratio of height to length of ilium 0.36; (8) PLOS ONE | www.plosone.org 2 November 2013 | Volume 8 | Issue 11 | e80557 New Oviraptorosaur from Southern China Figure 2. Lower jaw of Nankangia (GMNH F10003) in right lateral (A) and ventral (B) views. Abbreviations: d., dentary; sura., surangular. Scale bars = 5 cm. doi:10.1371/journal.pone.0080557.g002 femoral neck extending dorsomedially at about an angle of 90 to symphyseal suture is nearly straight. There is a distinct opening on the shaft; (9) femur and tibia approximately the same length. the suture near the rostral end of the lower jaw. The margins of the rostral end are sharp. In posterior view, the sutural portion is Description convex, forming a distinct ridge. In ventral view, the symphyseal Lower jaw. Both sides of the rostral part are preserved region is also U-shaped (Fig. 2B). (Figure 2). In lateral view, the rostral end of the jaw is not Dorsal vertebrae. (Fig. 3 A; see Table S1). Five dorsal downturned and the ventral margin of the dentary is nearly vertebrae are preserved. According to the position of the straight (Fig. 2A). Posteriorly, the dentary is divided into two parapophysis, these vertebrae are mid-dorsal vertebrae. But their branches that extend posterodorsally and posteroventrally, form- exact positions in the vertebral column are not clear, and thus ing the anterior margin of the external mandibular fenestra. Many these five dorsal vertebrae are referred to here as dorsal 1 to dorsal small nutrient openings are irregularly distributed on the lateral 5. The anterior articular end of the centrum is much more surface of the mandibular symphyseal region. In dorsal view, the concave than the posterior articular end, which is moderately cranial margin of the rostral end of the lower jaw is nearly straight; concave. The articular end is higher than wide and nearly thus the mandibular symphyseal region is U-shaped, similar