Zootaxa 3873 (3): 233–249 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2014 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3873.3.3 http://zoobank.org/urn:lsid:zoobank.org:pub:4441BCDF-E4A9-4C67-AE08-5D6D418706CC The first well-preserved coelophysoid theropod dinosaur from Asia HAI-LU YOU1,5, YOICHI AZUMA2, TAO WANG3, YA-MING WANG4 & ZHI-MING DONG1 1Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, 142 Xizhimenwai Dajie, Beijing, 100044, P. R. China. Email: [email protected] 2Fukui Prefectural University, 4-1-1 Kenjojima, Matsuoka, Eiheiji-cho, Yoshida-gun, Fukui 910-1195, Japan 3 Bureau of Land and Resources of Lufeng County, Yunnan Province, 651207, P. R. China 4School of Earth Sciences and Resources, China University of Geosciences (Beijing), Beijing 100083, P. R. China 5Corresponding author Abstract Coelophysoid dinosaurs represent the earliest major radiation of neotheropods. These small-to-medium-sized agile bipeds lived throughout much of Pangaea during the Late Triassic–arly Jurassic. Previously reported coelophysoid material from Asia (excluding the Gondwanan territory of India) is limited to two specimens that comprise only limb fragments. This paper describes a new genus and species of coelophysoid, Panguraptor lufengensis, from the Lower Jurassic Lufeng For- mation of Yunnan Province, China. The new taxon is represented by a well-preserved skeleton, including the skull and lower jaw, the presacral vertebral column and partial ribs, the right scapula, a partial forelimb, part of the pelvic girdle, and an almost complete hind limb. It is distinguished from other coelophysoid theropods by the unique combination of the following three character states: 1) diagonal (rostrodorsal-caudoventral) ridge on lateral surface of maxilla, within an- torbital fossa, 2) elliptical, laterally facing fenestra caudodorsal to aforementioned diagonal ridge, and 3) hooked cranio- medial corner of distal tarsal IV. Cladistic analysis recovers Panguraptor lufengensis deeply nested within Coelophysoidea as a member of Coelophysidae, and it is more closely related to Coelophysis than to “Syntarsus”. Pan- guraptor represents the first well-preserved coelophysoid theropod dinosaur from Asia, and provides fresh evidence sup- porting the hypothesis that terrestrial tetrapods tended to be distributed pan-continentally during the Early Jurassic. Key words: Theropoda, Coelophysoidea, new genus and species, Early Jurassic, Lufeng Formation, Lufeng Introduction Coelophysoid dinosaurs are small-to-medium-sized agile bipedal meat-eaters that lived throughout much of Pangaea during the Late Triassic and Early Jurassic (Tykoski 2005; Tykoski & Rowe 2004). They are among the earliest well documented dinosaurs and represent the earliest major radiation of neotheropods (Brusatte et al. 2010; Colbert 1989; Cope 1889; Sereno 1999). A recent study indicated that late Norian–Rhaetian theropod assemblages were dominated by basal (early diverging) coelophysoids, whereas Early Jurassic ones were composed of coelophysids (Coelophysis bauri + "Syntarsus" kayentakatae and all descendents of their most recent common ancestor), dilophosaurids and basal averostrans (Ezcurra 2012). However, despite the well-documented discoveries of derived coelophysoids in North America and Africa, the coelophysoid material that has previously been reported from Asia is limited to two specimens comprising only limb fragments and perhaps belonging to one individual (Irmis 2004). Here we describe a new genus and species of coelophysoid based on a well-preserved skeleton from the same rock unit, the Lower Jurassic Lufeng Formation of Yunnan Province, China, that yielded both previously reported specimens. Our cladistic analysis shows that the new taxon is a coelophysid coelophysoid, and is more closely related to Coelophysis than to "Syntarsus". This new taxon represents the most basal theropod dinosaur currently known in China, and provides fresh evidence supporting the hypothesis that terrestrial tetrapods tended to be distributed pan-continentally during the Early Jurassic. Institutional abbreviations: AMNH, American Museum of Natural History, New York, New York, USA; Accepted by S. Brusatte: 9 Sept. 2014; published: 16 Oct. 2014 233 FMNH, Field Museum of Natural History, Chicago, Illinois, USA; LFGT, Bureau of Land and Resources of Lufeng County, Lufeng, Yunnan, China; MNA, Museum of Northern Arizona, Flagstaff, Arizona, USA; NMMNH, New Mexico Museum of Natural History and Science, Albuquerque, New Mexico, USA; QG, Natural History Museum of Zimbabwe, Bulawayo, Zimbabwe. Systematic palaeontology Dinosauria Owen, 1842 Saurischia Seeley, 1887 Theropoda Marsh, 1881 Neotheropoda Bakker, 1986, sensu Sereno, 1998 Coelophysoidea Nopcsa, 1928, sensu Sereno, 1998 Coelophysidae Nopcsa, 1928, sensu Holtz, 1994 Panguraptor lufengensis gen. et sp. nov. Holotype: Bureau of Land and Resources of Lufeng County LFGT-0103, an articulated partial skeleton that includes the skull and lower jaw, the presacral vertebral column, part of the ribs, the right scapula and partial right forelimb, part of the pelvic girdle and parts of both hind limbs, the right hind limb being almost complete. Type locality and horizon. Lufeng County, Yunnan Province, China. Shawan Member of the Lufeng Formation (Fang et al. 2000). This is equivalent to the Dull Purplish Beds of the Lower Lufeng Formation (Series) as defined by Bien (1940). In the Lufeng area, the Lufeng Formation overlies the Lower Jurassic Yubacun Formation and is disconformably overlain by the Middle Jurassic Chuanjie Formation (Cheng et al. 2004). The age of the Lufeng Formation is probably early - middle Early Jurassic (Hettangian - Pliensbachian), based on biostratigraphic correlations (Luo & Wu 1995). Magnetostratigraphic analysis has indicated late Sinemurian – ?Toarcian for the age of the Lufeng Formation (Huang et al. 2005). Etymology. The genus name is from “Pangu” (Chinese), well known in Chinese mythology as the first living being and the creator of all reality, and “raptor” (Latin), meaning “thief” or “robber”. The specific name refers to “Lufeng County”, one of the world’s richest sources of Early Jurassic terrestrial vertebrate fossils. Differential diagnosis (for genus and species by monotypy). A coelophysid theropod with the unique combination of the following three character states (autapomorphies marked with *): 1*) diagonal (rostrodorsal- caudoventral) ridge on lateral surface of maxilla, within antorbital fossa; 2) elliptical, laterally facing fenestra caudodorsal to diagonal ridge mentioned in previous character state, also present in Zupaysaurus rougieri (Ezcurra 2006); 3*) distal tarsal IV with hooked craniomedial corner. Description and comparisons. LFGT-0103 is an articulated partial skeleton, exposed mainly in right lateral view (Fig. 1). The specimen is small, with a preserved skull length (taken from the rostral end of the maxilla to the ventral end of the quadrate because the premaxilla is missing) of 11.1 cm, presacral vertebral column length of ~58 cm (trunk/neck length ratio about 1.3), scapula length of 8.6 cm, femur length of 16.4 cm, and tibia length of 18.2 cm (Table 1). Early theropods normally had a long tail contributing more than half of the total body length, implying that this dinosaur may have measured around two metres long at the time of death. Height at the hip was probably about half a metre. LFGT-0103 probably represents a sub-adult individual, as indicated by the relatively small body size of the specimen, the large orbit, the lack of fusion between the scapula and coracoid, the lack of ossification of distal tips of the ischia, and the separation of the astragalus and calcaneum. However, the neurocentral sutures are fused, cervical ribs are fused to their respective centra, the ilium is fused to the ischium, and distal tarsal 3 is probably fused to metatarsal III, indicating that the specimen may be close to an adult individual. Growth curves of femur 234 · Zootaxa 3873 (3) © 2014 Magnolia Press YOU ET AL. length vs. age for C. rhodesiensis (Chinsamy 1994) and C. bauri (Rinehart et al. 2009) suggest, respectively, that this dinosaur died at a sub-adult age of around four years or around two years. TABLE 1. Measurements of selected elements of Panguraptor lufengensis gen. et sp. nov. LFGT-0103. All right side, in mm. skull length (from preserved rostral end to ventral end of quadrate) 111 preorbital length (preserved) 60 length of internal antorbital fenestra 37 height of antorbital fossa along at caudal margin along lacrimal 20 orbit length 30 orbit height 35 maximum height of skull across orbit midpoint 48 length of infratemporal fenestra 23 height of infratemporal fenestra 31 width of skull above middle antorbital fossa 12 narrist width of skull above orbit 10 length of low jaw (preserved) 111 Cervicals centrum length total height axis 18.5 22 C3 24 14.5 C4 28 16 C5 29 15 C6 31 23 C7 30.5 22 C8 28.5 - C9 28 - C10 23 - Dorsals centrum length total height D9 24 32 D10 26 36 D11 26 35 D12 25 34 D13 23 23+ sacral 1 length 22+ scapula length 86 maximum width of scapular blade 20 minimum width of scapular blade 11 humerus length 58+ metacarpal I length 10+ metacarpal II length 36 metacarpal II width at midportion 5 metacarpal III length 34 metacarpal III width at midportion 3.5 metacarpal IV length 22 metacarpal