The First Well-Preserved Coelophysoid Theropod Dinosaur from Asia

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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 antorbital fossa, 2) elliptical, laterally facing fenestra caudodorsal to aforementioned diagonal ridge, and 3) hooked craniomedial 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 supporting 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 IV width at midportion 3 ...... continued on the next page

FIRST COELOPHYSOID DINOSAUR FROM ASIA Zootaxa 3873 (3) © 2014 Magnolia Press · 235 TABLE 1. (Continued) Dorsals centrum length total height manual phalanx I-1 length 17 manual phalanx I-2 (Ungual) length 21 manual phalanx II-1 length 18 manual phalanx II-2 length 22 manual phalanx II-3 length 12+ manual phalanx III-1 length 10 manual pahlanx III-2 length 8 manual phalanx III-3 length 9 manual phalanx IV-1 length 4+ Iliac blade length 63+ iliac blade height above acetabulum 44 femur length 164 tibia length 182 tibia length with astralas 188 fibula length 178 metatarsal IV length 98 metatarsal V length 41 pedal phalanx III-2 length 22 pedal phalanx III-3 length 23 pedal phalanx III-4 length 17 pedal phalanx IV-1 length 18 pedal phalanx IV-2 length 17 pedal phalanx IV-3 length 13.5

FIGURE 1. Skeleton of Panguraptor lufengensis gen. et sp. nov. (LFGT-0103). a, photo; b, interpretive line drawing. Abbreviations: as, astragalus; ax, axis; ca, calcaneum; cer 3–10, cervical 3-10; dor 7, dorsal 7; dt IV, distal tarsal IV; fdt III, fused distal tarsal III; fi, fibula; hu, humerus; il, ilium; lfe, left femur; lj, lower jaw; is, ischia; ma, manus; mt IV, metatarsal IV; mt V, metatarsal V; pd III, pedal digit III; ph IV-1, phalanx 1 of pedal digit IV; ra, radius; rfe, right femur; sac 1, sacral 1; sc, scapula; sk, skull; ti, tibia; ul, ulna.

236 · Zootaxa 3873 (3) © 2014 Magnolia Press YOU ET AL. FIGURE 2. Skull and lower jaw of Panguraptor lufengensis gen. et sp. nov. (LFGT-0103) in right lateral view. a, photo; b, interpretive line drawing. Abbreviations: af, additional fenestra; ar, alveolar ridge; at, atlas; ax, axis; dr, diagonal ridge; emf, external mandibular fenestra; f, frontal; gd, groove on dentary; iaf, internal antorbital fenestra; itf, infratemporal fenestra; j, jugal; l, lacrimal; m, maxilla; n, nasal; or, orbit; pa, parietal; pf, promaxillary fenestra; po, postorbital; pr, prefrontal; q, quadrate; qj, quadratojugal; rsur, ridge on surangular; sq, squamosal; stf, supratemporal fossa.

Skull and lower jaw. The skull is almost complete, and exposed mainly in right lateral view (Fig. 2). Assuming that the missing premaxillary body contributed ~10% of the total skull length (based on skull reconstruction of the “Syntarsus”c kayentakatae holotype of Tykoski 1998, figure 6), the intact skull of Panguraptor would have measured 12.4 cm, about half as long as adult skulls of C. bauri (AMNH 7224: 20.7 cm) (Rinehart et al. 2001), C. rhodesiensis (QG 193: 22.0 cm) (Raath 1977), and “S.” kayentakatae (MNA V2623: 23.0 cm) (Rowe 1989). The preorbital length without the premaxilla (6.0 cm) is slightly more than half the preserved skull length (54%), and the maximum rostrocaudal length of the large triangular internal antorbital fenestra is about two thirds of the preserved preorbital length. In contrast, the internal antorbital fenestra is relatively small (about half as high as the orbit and half as long as the maxilla) in a similar-sized (skull length 12.3 cm) juvenile specimen of C. bauri NMMNH P-42200 (Rinehart et al. 2009). The orbit is subcircular, though with a relatively straight rostral edge, and has a maximum length of 3.0 cm and maximum height of 3.5 cm. The maximum height of the skull at the mid-orbital level is 4.8 cm, or 39% of the estimated skull length. The postorbital portion of the lateral side of the skull is almost completely occupied by the rectangular infratemporal fenestra, which is 2.3 cm long and 3.1 cm high. Most of the lateral surface of the right maxilla is exposed. The rostral process of the maxilla and the rostrodorsal edge of the maxillary body proper (the portion between the rostral process of the maxilla and the rostral rim of the internal antorbital fenestra, but not including the ascending process) have been lost, and the preserved rostrodorsal margin appears to represent the caudoventral rim of the external naris. The maxillary body proper is rostrocaudally longer than dorsoventrally high as in C. rhodesiensis (QG 193), but the opposite is true in “S.” kayentakatae (MNA V2623). The lateral surface of the maxillary body proper bears a diagonally aligned (rostrodorsal-caudoventral) ridge between the rims of the internal and external antorbital fenestrae. Caudodorsal to this ridge an elliptical fenestra opens laterally, while rostroventral to it are two small fenestrae separated by another short vertical ridge: a rostrally positioned fenestra that is more or less round in lateral view, and a caudally positioned one that is triangular. We interpret the rostral one as the promaxillary fenestra, which occupies a similar position in “S.” kayentakatae. However, the part of the antorbital fossa that lies just caudal to the promaxillary fenestra is smooth in “S.” kayentakatae, rather than interrupted by additional fenestrae. The longitudinal and very prominent alveolar ridge runs along the entire ventral border of the external antorbital fenestra and extends onto the rostral process of the jugal under the orbit. In “S.” kayentakatae the alveolar ridge extends further caudally, continuing onto the caudal process of the jugal. The ascending process of the maxilla is broken away from the rest

FIRST COELOPHYSOID DINOSAUR FROM ASIA Zootaxa 3873 (3) © 2014 Magnolia Press · 237 of the bone, and judging from the orientation of the nasal and the shape of the antorbital fenestra, it would have been elevated about 30 degrees above the horizontal in the intact skull. The orientation of the ascending process is similar in C. rhodesiensis, but steeper in “S.” kayentakatae.

FIGURE 3. Cervical vertebrae 2-6 of Panguraptor lufengensis gen. et sp. nov. (LFGT-0103) in right lateral view. Abbreviations: a rib, axial rib; C4-C6, cervicals 4–6; cp, caudal pleurocoel; rib of cer 3, rib of cervical 3; rp, rostral pleurocoel; sg, shallow groove on axis.

A large portion of the right nasal is visible, exposed in dorsal view. A large and subcircular hole penetrates the rostral half of the bone, probably not a true feature of the animal in life. The dorsal surface of the nasal is smooth, and lacks any evidence of a parasagittal vertical crest. The transverse width of the nasal above the antorbital fenestra exceeds the width of the frontal above the orbit. The transverse width of the prefrontal exceeds its rostrocaudal length. A depression is visible rostrodorsal to the orbit, at the junction between the nasal, prefrontal and frontal. The robust and almost vertical ventral ramus of the lacrimal separates the caudal border of the antorbital fossa from the rostral border of the orbit. The caudodorsal corner of the lateral surface of the lacrimal bears a rostrodorsal-caudoventrally running groove, the caudal edge of which leads to a small foramen, which should be

238 · Zootaxa 3873 (3) © 2014 Magnolia Press YOU ET AL. the exit of the naso-lacrimal duct. The lateral lamina seems to be restricted to the caudal margin of the ventral half of the ventral ramus of the lacrimal, and lacks an expansion at the dorsal end. The rostral process seems to contribute at least one third to the dorsal border of the antorbital fenestra because its rostral end appears to be broken off and incomplete. The jugal borders the orbit ventrally, and seems to contact the maxilla ventral to the lacrimal as in C. rhodesiensis (QG 165) (Bristowe & Raath 2004). The caudal edge of the postorbital process of the jugal forms an angle of approximately 700 with the dorsal edge of the caudal process, rather than a right angle as in “S.” kayentakatae. The ventral branch of the short and forked caudal process is slightly longer than the dorsal branch, and the slot for the quadratojugal formed by the two branches extends rostrally to a point located ventral to the base of the postorbital process. The quadratojugal forms the ventral half of the caudal border of the infratemporal fenestra, and contacts the ventral process of the squamosal. This squamosal-quadratojugal contact excludes the quadrate from the border of the infratemporal fenestra, as in “S.” kayentakatae (Tykoski 1998) but in contrast to the condition in C. rhodesiensis (Raath 1977). The frontal-parietal contact region is damaged. The dorsal surface of the frontals bulges along the midline at the level of the caudodorsal corners of the orbits. The rostral portion of the well-developed supratemporal fossa is bounded ventrolaterally by a sharp rimmed edge on the rostral process of the postorbital. As preserved, the lower jaw is oriented almost perpendicular to the long axis of the skull, and the mandibular glenoid fossa is preserved close to the quadrate head and considerably displaced from the distal condyles of the quadrate. Judging from the position of the external mandibular fenestra, which is normally located underneath the orbit, the rostral end of the lower jaw comparable to the premaxilla is missing; in actuality the preserved lengths of the skull and the lower jaw are same. The caudal end of the lower jaw is overlapped by the atlas and axis. It is difficult to discern the sutures among the bones of the lower jaw. A longitudinal groove is visible on the lateral surface of the middle portion of the dentary, and a longitudinal ridge is present on the lateral surface of the surangular. Six maxillary teeth are visible, four in the rostral one-third of the maxilla and two at the caudal end of the bone under the caudal part of the orbit. Ten dentary teeth are preserved. All maxillary and dentary teeth are slightly recurved, and bear few or no serrations. Vertebral column and ribs. Ten cervicals, 13 dorsals and the first sacral vertebra are preserved (Fig. 1). The atlas is disarticulated in such a way that its individual components are hard to identify (Fig. 2). The axis is the shortest of all the cervicals, except for the atlas. The exposed right lateral side of the axial centrum does not bear any clearly visible pneumatic excavations (pleurocoels), although a shallow longitudinal groove is present (Fig. 3). Although the neural spine is not complete, it is clearly tall and craniocaudally long. The centrum of the third cervical is about 130% longer than that of the axis. Centrum lengths continue increasing steadily until cervical 7, then decrease gradually so that the last cervical is about equal in centrum length to the third one. All postaxial cervicals are elongate, with mid-central constrictions and stout prezygapophyses. The lateral surfaces of the centra of cervicals 4-5 bear deep, elongate pneumatic foramina (“pleurocoels) (Fig. 3). In cervical 4, the rostral part of the pleurocoel is larger and deeper than the caudal part, but in cervical 5 the opposite is the case. Only weak pleurocoels are present on cervicals 6-7, and the condition in the rest of the cervicals is unknown due to poor preservation. The shafts of the axial ribs are preserved ventral to cervical 3, and one of them extends far enough to lie partly ventral to cervical 4 (Fig. 3). Although it is hard to trace individual ribs in the remainder of the cervical series, the existence of bundles of elongated ribs is evident. The third cervical rib has a small capitulum and tuberculum. In contrast, the remaining cervical ribs have well-developed capitula and tubercula, and are fused to their corresponding vertebrae. The centra of the well-preserved caudal dorsals are less than twice as long as they are high, whereas the corresponding centra are about twice as long as high in C. bauri and C. rhodesiensis. The neural arches of the cranialmost seven dorsal vertebrae have sheet-like transverse processes that are strongly backswept and triangular when viewed from above. In contrast, the transverse processes of the eighth and ninth dorsals are rostrocaudally narrow and taper laterally (Fig. 1). The parapophysis is short, and situated cranioventral to the transverse process. The neural spines are longer than high and so close together that they almost form a thin, continuous wall along the dorsal midline of the axial column. Only the first sacral is exposed, while the others are covered by the right ilium. This vertebra is slightly shorter than the caudal dorsals, and has a round, flat cranial articular surface.

FIRST COELOPHYSOID DINOSAUR FROM ASIA Zootaxa 3873 (3) © 2014 Magnolia Press · 239 Appendicular skeleton. An almost complete right scapula is exposed in lateral view, and is broken only near the articulation with the coracoid (Fig. 1). The scapula is long. Its blade is expanded distally, and constricted proximally. The caudal edge of the blade is straight, while the cranial edge is concave. The coracoid is not preserved, indicating that it is not fused with the scapula. In describing the forelimb, we orient it so that it is held vertically with the radius positioned cranial to the ulna and the flexor surface of the manus facing medially. The distal portion of the right humerus, including what seems to be the part of the deltopectoral crest that lies distal to the apex, is preserved (Fig. 4). The distal end of the humerus curves slightly cranially. In distal view, the medial condyle is slightly mediolaterally wider than the lateral one. The right radius and ulna are preserved, and are exposed mainly in caudal view (Fig. 4). The well-preserved proximal end of the radius is well preserved, and can be seen in proximal view. The proximal surface is subrectangular in shape, being craniocaudally elongated, and protrudes slightly caudally, unlike the strongly caudally protruded condition in the Lufeng specimen FMNH CUP 2089 (Irmis 2004). The distal end of the radius is not exposed. The ulna is broken into four pieces, the distalmost of which is detached from the others (Fig. 1).

FIGURE 4. Forelimb of Panguraptor lufengensis gen. et sp. nov. LFGT-0103. Abbreviations: dhu, distal end of humerus; dpc, deltopectoral crest; mtc I, metacarpal I; mtc IV, metacarpal IV; ph I-1, phalanx 1 of digit I; ph III-3, phalanx 3 of digit III; ph IV- 1, phalanx 1 of digit IV; pra, proximal end of radius.

The right manus is represented by four digits, exposed mainly in medial and caudal views (Fig. 4). Metacarpals I and II are more than twice as wide as metacarpals III and IV. Metacarpal I is less than half the length of metacarpal II, which is about the same length as metacarpal III. The slender metacarpal IV is preserved between metacarpals II and III, and measures about two thirds the length of metacarpal III. Phalanx 1 of digit 1 is not completely exposed, but seems to be as long as phalanx 2 of digit 2, one of the longest phalanges in the manus. The claw of digit 1 is almost the same length as phalanx 2 of digit 2, and is transversely compressed, strongly recurved, and equipped with a large flexor tubercle. The claw of digit 2 is not completely exposed, but seems less well developed than that of digit 1. Three slender phalanges of digit 3 are visible, but there is no trace of a claw. Only

240 · Zootaxa 3873 (3) © 2014 Magnolia Press YOU ET AL. one small phalanx appears to be present in digit 4, although an additional small terminal nub may also exist as in “S.” kayentakatae (Tykoski 1998). Therefore, the manual phalangeal formula is 2-3-3-1/2-X. Part of the right ilium is preserved (Fig. 1). Most of the pre- and postacetabular processes are missing, but the portion surrounding the acetabulum is well preserved. The pubic peduncle is much stouter than the ischial peduncle, and bears an approximately triangular articular surface for the pubis in distal view. The acetabulum is bordered by a prominent supracetabular crest craniodorsally. The bone surface that forms the caudal and caudodorsal parts of the wall of the acetabulum is relatively narrow transversely and faces slightly laterally. The ischial peduncle is fused to the ischium, although a suture can be vaguely discerned. The distal portions of the ischia are preserved (Fig. 5). They are straight and mutually appressed, with moderately enlarged ends that are about two times broader craniocaudally than the shaft.

FIGURE 5. Distal portions of ischia and right hind limb of Panguraptor lufengensis gen. et sp. nov. LFGT-0103. Abbreviations: as, astragalus; ca, calcaneum; dt IV, distal tarsal IV; fdt III, fused distal tarsus III; fi, fibula; lis, left ischium; mt IV, metatarsal IV; mt V, metatarsal V; ris, right ischium; ti, tibia.

The right femur is broken into two portions. The proximal portion is exposed in cranial and lateral views, and the distal portion in lateral view (Fig. 1). The femoral head is enlarged, well offset from the shaft, and directed craniomedially, with its end turned slightly downward. In proximal view, no clear constriction is apparent, and the proximal surface tapers caudolaterally. A longitudinal bulge is present near the proximal end of the caudolateral surface of the shaft. The anterior trochanter begins at the level of the ventral edge of this bulge, centered transversely on the craniolateral surface of the femur, as a prominent boss that extends ventrally. A tubercle lying caudal to the ventral end of the anterior trochanter may be homologous to the trochanteric shelf. The area proximal to the anterior trochanter is shallowly depressed. The right tibia and fibula are preserved, and exposed mainly in proximal and lateral views (Fig. 1). In proximal view, they combine to form a flat and roughly square articular surface for the femur. The cnemial crest forms the craniolateral corner of this surface, while the craniomedial corner consists of two small tubercles of the tibia that protrude slightly above the surface. The shafts of both the tibia and fibula are straight, and the fibula is closely appressed to the lateral surface of the tibia. The distal ends of both bones are enlarged. The astragalus and calcaneum are clearly not fused to each other, or to the tibia and the fibula (Fig. 5). The astragalus is visible in caudodistal view, and the calcaneum is slightly displaced from its original position. The articular surfaces between the astragalus and calcaneum are flat. In lateral view, the calcaneum acquires a semilunate shape.

FIRST COELOPHYSOID DINOSAUR FROM ASIA Zootaxa 3873 (3) © 2014 Magnolia Press · 241 One distal tarsal is visible, and is very likely to be distal tarsal IV based on its position and shape (Fig. 5). Its proximal surface is slightly depressed, and it has an irregular outline with an expanded cranial portion. The craniomedial corner of the bone is hooked, rather than a simple right angle as in “S.” kayentakatae (Tykoski 1998). Right metatarsals III, IV and V are partially exposed (Fig. 5). The proximal end of metatarsal IV is positioned slightly distal and dorsal to that of metatarsal III, indicating that the latter incorporates a fused distal tarsal III. The proximal end of metatarsal III extends ventrally well beyond the metatarsal shaft, and bears a clear ventrolateral process. The proximal surface of metatarsal IV has a triangular shape matching that of distal tarsal IV, with an expanded dorsal portion and a tapering ventral end. The fifth metatarsal is splint-like, with a proximal end at the same level as that of metatarsal III and a distal end that tapers abruptly to a point. The four phalanges of pedal digit III are preserved, as are the proximal three phalanges of pedal digit IV (Fig. 1).

Cladistic analysis

A cladistic analysis was performed in order to assess the phylogenetic position of Panguraptor. The data matrix was based on that of Ezcurra & Brusatte (2011), which consists of 339 characters and 43 dinosaur and other archosauriform taxa. In our analysis, Panguraptor was added, and two characters (222 and 316) were recoded from state ‘0’ to state ‘1’ for “S.” kayentakatae (see Appendix for data matrix). Erythrosuchus was used as an outgroup. A traditional search in TNT (Goloboff et al. 2008) with 100 Wagner tree replications and tree bisection and reconnection (TBR) branch swapping (saving 100 trees per replicate) recovered 8 MPTs (most parsimonious trees) of 1011 steps each. The CI (consistency index) was 0.400, and the RI (retention index) was 0.691. The strict consensus of these eight trees is shown in Fig. 6, with absolute bootstrap frequencies and all decay index values equal to or greater than one.

FIGURE 6. Phylogenetic placement of Panguraptor lufengensis gen. et sp. nov., based on strict consensus tree obtained in this study. Absolute bootstrap frequencies and decay index values are indicated.

242 · Zootaxa 3873 (3) © 2014 Magnolia Press YOU ET AL. The strict consensus recovered Panguraptor as a member of Coelophysidae, and placed Panguraptor more closely related to the grouping of C. bauri + (C. rhodesiensis + Camposaurus arizonensis) than to “S.” kayentakatae. The strict consensus also recovered Liliensternus liliensterni and Zupaysaurus rougieri as members of Coelophysoidea. The monophyly of Coelophysoidea is supported by four unambiguous synapomorphies: 17(1>2): more than 18 maxillary teeth; 24(0>1): sharp longitudinal ridge present on lateral surface of the maxilla; 194(0>1): caudal margin of postacetabular process of ilium notched or indented; and 320(3>2): sublacrimal part of jugal squared off rostrally with small dorsally directed prong, slightly overlapping lacrimal. Of these character states, only 24(1) is clearly present in Panguraptor. Coelophysidae is supported by eight unambiguous synapomorphies: 47(0>1): width of ventral process of squamosal less than one quarter of its length; 129(0>1): cervical vertebrae with rimmed depression on caudal part of centrum; 139(0>1): sacral centra co-ossified at ventral edges; 230(1>2): dorsolateral margin of proximal portion of femur bearing rounded ridge; 234(0>1): trochanteric shelf present proximal to fourth trochanter on femur; 240(0>1): infrapopliteal ridge present between medial distal condyle and crista tibiofibularis on femur; 265(0>1): distinct deep fossa on medial side of fibula; and 323(1>0): small contact between squamosal and quadratojugal. Of these character states, 47(1), 129(1), 230(2), 234(1), and 323(0) are present in Panguraptor. The Panguraptor + Coelophysis clade is supported by four unambiguous synapomorphies: 316(0>1): angle between ascending and horizontal processes of maxilla less than 35; 318(0>1): lateral surface of maxillary antorbital fossa at base of ascending process bears at least one deep, large, blind pocket that is subcircular or oval in shape; 319(3>2): lateral lamina of lacrimal with no interruption of the lacrimal antorbital fossa and restricted to the caudal margin of the ventral ramus, and 321(0>1): angle between ascending process and longitudinal axis of jugal less than 75, with ascending process strongly caudodorsally oriented. All four of these features are present in Panguraptor. The Coelophysis bauri + (Coelophysis rhodesiensis + Camposaurus arizonensis) clade is supported by five unambiguous synapomorphies: 27(1>0): absence of promaxillary foramen; 322(1>0): tapering ventral process of squamosal; 323(0>2): no squamosal-quadratojugal contact; 324(0>1): no postorbital participation in supratemporal fossa; and 329(1>2): significantly elongated caudal dorsal vertebrae, with centrum length at least 1.33 times height of cranial articular surface. All these features are absent in Panguraptor.

Discussion

There are three other Early Jurassic coelophysoid specimens not included in the above cladistic analysis, and we compare them with Panguraptor lufengensis. Irmis (2004) referred FMNH CUP 2089 (distal humerus, proximal radius and ulna, fragments of metacarpals and phalanges) and FMNH CUP 2090 (right distal tarsals II and III, and proximal metatarsals II and III) from the Early Jurassic Lufeng Formation in Lufeng Basin to cf. Megapnosaurus (=Syntarsus). Comparable portions of FMNH CUP 2089 and the holotype of Panguraptor lufengensis are similar in morphology, and both specimens share a cranially curved distal end of the humerus and uneven distal humeral condyles. Distal tarsal III of the Panguraptor lufengensis holotype possesses a ventrolateral process that is not present in FMNH CUP 2090, but this feature may have been lost in the latter specimen due to poor preservation. However, pending further discoveries here we still refer FMNH CUP 2089 and 2090 to cf. Megapnosaurus. Carrano et al. (2005) restudied the holotype and only known specimen of Segisaurus halli Camp 1936, a small coelophysoid from the Lower Jurassic of North America. Segisaurus is distinctive in having a very long and slender scapula, measuring 93 mm long and 20 mm wide at the distal end. The equivalent measurements are 86 mm and 20 mm in Panguraptor lufengensis. Histological study indicates that the holotype individual of Segisaurus halli was a subadult at the time of death (Carrano et al. 2005). Tykoski (2005) argued for the existence of a distinct ‘Shake-N-Bake’ coelophysoid species from the Kayenta Formation, which also yielded “S.” kayentakatae, and showed that the ‘Shake-N-Bake’ species was a small coelophysoid. The proximal tarsals, the astragalus and calcaneum, are clearly well fused to the tibia, and to each other in the small ‘Shake-N-Bake’ species, but such fusion is not apparent in the holotype of Panguraptor lufengensis despite the larger size of this specimen.

FIRST COELOPHYSOID DINOSAUR FROM ASIA Zootaxa 3873 (3) © 2014 Magnolia Press · 243 The Lufeng Saurischian Fauna (Young 1951), or more appropriate the Early Jurassic Lufeng Dinosaur Fauna, represents one of the richest dinosaur faunas in the world, and provides critical information regarding dinosaur evolution and biogeography during the Early Jurassic. Eight species of sauropodomorphs (Lufengosaurus huenei Young, 1941, L. magnus Young, 1947, “Gryposaurus” sinensis Young, 1941, Yunnanosaurus huangi Young, 1942, Y. robustus Young, 1951, Jinshanosaurus xinwaensis Zhang and Yang, 1995, Chuxiongosaurus lufengensis L et al., 2010, Xixiposaurus suni Sekiya, 2010) and two species of ornithischians (Tatisaurus oehleri Simmons, 1965 and Bienosaurus lufengensis Dong, 2001) have been reported from the Lower Jurassic Lufeng Formation in the Lufeng area. In contrast, only one theropod, Sinosaurus sinensis Young, 1948 has been previously confirmed to exist in these strata (Xing 2012). The discovery of Panguraptor lufengensis adds a second theropod to the known fauna of the Lufeng Formation.

Acknowledgements

We are grateful to the crew of the Lufeng World Dinosaur Valley for discovering, excavating, and preparing the specimen, to Ms. Jin-Ling Huang for producing the line drawings, and to Dr. Corwin Sullivan for detailed review of the first draft. Thanks also go to reviewers (Dr. Ronald Tykoski and Mr. Martin Ezcurra) and editor Dr. Stephen Brusatte. Funding was provided by the Hundred Talents Project of the Chinese Academy of Sciences, the National Natural Science Foundation of China, the National Basic Research Program of China (973 Program), and the Bureau of Land and Resources of Lufeng County.

References

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Erythrosuchus_africanus 00010001200002000100000000000000000010000000000000001010000000001000000000000000000000001000000000000 100000000000110000000?00000000010?000000000000000??1000?001?000101?000?00??01??0???000?0000001000000000 000??000000000010001000000000001??00000000000000000000010000010000000?00?00?0000000000?000000000001000 ?0???????000010110101001000001000 Euparkeria_capensis 000100000000?000100000000100000000000000000100000000[01]0002000000010100000000000001000000000000000000 011000000000001000000000000000010000010000000000000001000?00000010000000000??010?000?00000000000000000 000000??0000000000100010001000000000000000000000000000000000?00000000000?00?0000000000010?000000000000 0000101000001002100000110??000001000 Revueltosaurus_callenderi 000000002000030110101001020101000000000010010100011012002000000011211000000?20????0?001?001?0000000011 0000110000?1110000000001000020100011000000000000?01001010000000110110?0???000?0?0?????00000000000000000 00??000000000000011000100000000000000000000010000001000010000?000100?00?0001101101021010?0000?000000?? 10012112?2022001??01???00000100? Aetosaurus_ferratus 10100000111000010110011002010?00?10000000011010?011012002011000??12?1?00000?????1???001000110??21??1??00 01000010?111??????????????????????0?000000???0001?01010000000110110?00??00000???00000?[01]000?00000000000? ??00?0000000?0011000100000000000000?00000?10?000010??0????0??????????????????1?1?21?????????0??0???2100121 12210221??0110???000001000 Arizonasaurus_babbitti ??????1????00?00??00011000000100???????000?100000000?000200000001001100000000011?010001?001?000001101100 000000000111000010?0000000010?0000?11000100???0010?1010?????????????????????????????00100010020000?0011100 21111100011?10000100000001001000100?0????????????????????????????????????????????????????????????0???????200? 20??00?10110???????? Effigia_okeeffeae 011000104??0?20???01?111000000000000000011?000120010???000?1000001211010?????111?0?0001?001000?010010001 02????????11???000?01?0?1101??000111111011?0110?00?1001??0?10?1?000100??00?000??????10110000021010?0011110 2122100001??30100??0000011??101110000001?0100010010100010000100?0??00011011?1?11?10000001010100?10?????? 00102200??10000?000001000 Batrachotomus_kuperferzellensis 2?0?000?10010100011001100000111101001001100110011000?00020000001112110011000?01111110011101?00000110110 000000000011100000000000000211000100010001001000010?101000000011110??????????????????0010001000000010011 00010010100011010000100000001001010000000010000001010010000??????0??1????110?101021?100000??000?10?2111 ?10?012021001000101100000100? Postosuchus_kirkpatricki 0001000010000101100000020101111101001001101110011100111031000001112110?110012011??1?0011111100020110110 0000000000111??00???????000?1?00000000000000??0??1011011001000111?00100??00???0??0000001?0010000000?????? ??1??1110001?010000100000000001010?00000010000001011011000?????00?01?0001102101111?10000001010110?0111?1 00?120031010110011000001000 Dromicosuchus_grallator 2000000?200011111100000001001?1000010001111110?1?10011102110????????????100?????????001?011?000?021011000 000000001?1????????0??0000?1???000?0000000?00201011111001110011?00100?10???????????00110010000100?00????01 ??????????010000100000100001010?00000010000001001011000?????00??1????11021011?1?11?00???????????1110100012 0220??01?1?210???????? Eudimorphodon_ranzii 1110???010100?0?2000011000000??0??0?000???0?????0?00000?0??1????????????????????????10??0?100?000000?00?001 ?0000?101????1???0000000?1?0?0?00?????0?01000????110000110?0?001??????00000110?110?0100???0?00?00000??00?? 0?0000001????????0?00??2??????0?0??????0?????????????1?0???0?1??1????1?01?????11?001?1??0110000??????1200031? 211000210000???0? Dimorphodon_macronyx 2100??1?2010000?11?0011000000?????00000???010?0?00000??00??0????????????????????????101?20?00??00??0?00??01 ?0000?1?1????1?1??????00?1??000?????????010??????110000110?????1?0???00000?110?1?000100?000100?00000??00??? ?0000?0110000?00000000??001020000000?00?000?0??1?00?11000?0?1??101??1??1?????110001?001??1?000???????2000 ?0?21???011000011000 Lagerpeton_chanarensis ?????????????????????????????????????????????????????????????????????????????????????????????????????????????????? ?????????????????????0??000000000?????????????????????????????????????????0000000000000000000??00000000000011 11001100010?0100010?0010100?0000000010100011100110?1001011?1?01???2011100000000001200???????????????????? 210?0000001

246 · Zootaxa 3873 (3) © 2014 Magnolia Press YOU ET AL. Dromomeron_romeri ?????????????????????????????????????????????????????????????????????????????????????????????????????????????????? ???????????????????????????????????????????????????????????????????????????????????????????????????????????111100 1100010102?001110111100000000000101?00??????10?1001011?1?01???2?1???????????????0???????????????????????0000 010? Dromomeron_gregorii ?????????????????????????????????????????????????????????????????????????????????????????????????????????????????? ???????????????????????????????????????????????????????????????????????????????????????????????????????????111100 1?01010100000111011110000000000?????00??????1????????1??????????????????????????????????????????????????000??? ?? Marasuchus_lilloensis ????????????????????????????????????????????????????????????????01210000?001100????0???????????????????????????? ???101000?0000000000?00000000000000000??1?????1000?101???0??????????????????0000000000100000010??11000000 01011100001001001100000?0?00001000000000101010000110010101001001?010000100111000000000012?0???????00??? ???????010000011000 Eucoelophysis_baldwini ?????????????????????????????????????????????????????????????????????????????????????????????????????????????????? ???????????????????????????????????????????????????????????????????????????????????10????????????????????0??12001 ??1110001???0??1100010?100??????????????????????????????????????????????0??????????????????????????????????????? Sacisaurus_agudoensis ??????????????010?00000000000?????????????????????????????????????????????????????????1?????0?101????????101001 110?1????????????????????????????????????1?????1?????????????????????????????002???010?0000100?0??1???????????? 100111011100010000???100010010110001?????????????????????????????????????????????????????????200??????????010 000????? Silesaurus_opolensis 0?0?000?20?0000100000000000001?000?0???00?0100100000?10020001??00021010000011001?001001?001001121001010 00100001010111100100000000001000000000000110000??110101100001010?000?00??0???????????002000010100001001 0??110110000111012011101110[0 1]01000010110001001011000101110001?1???0101001010?0?01???2011110?00010101210????????00?20????000110000111 0? Eocursor_parvus ???????????????????????????????????????????????0????????????????01210100??????????????????1??100?001??10?0?1011 ?11?1????????0?0001?0??00?10?001?1?0?????11????11?0?101??????????????????????00210001020000?11????01??10?002 1?110002101200011110010?000020000?2010101?10??1??????1???????????????????11???000???????0?????????????????1? ???0100???0? Pisanosaurus_mertii ???????????????11??01??0????????????????????????????????????????????????????????????0?????????000001?01200?1111 111?1?????0?0???0000???00??????????????????????????????????????????????????????????0?????????0?????????????2???? ???????????????001??0000?0000010001011100?1?????01010?111??0?01???21111???0?01??????0????????0??????????12?? 0000010? Heterodontosaurus_tucki 000100000011101100001000000001000000000011010002000003001000101?0121?110000111???0??001101111?0[02]0001 00120[02]11100111111000000001000000?000211?0011100001?111?0?1110011010001011000100001120000?021000002100 011110??01??100002011?0??????20?010110010?0000200001101??0110???111000?1?10?1?1??1?01?????111100?00101?120 0??????20001100101000100?00???00 Lesothosaurus_diagnosticus 00010?0?3000001110001000000001000000000011010000000003000010101001210100000??1??10??0011011111000001011 2000101101111????000001000000??0001??0?111?0??1??110???1100?1000?01010???000?00??000000210?0102100011110? ?01??1000021111000210120001011?0100000020000020101?111?00?????001?100111??0?01???21111100000101???00????? ??0001010000000??010????00 Scutellosaurus_lawleri ??0??0?????????1?0?01??000?????????????00??10??2???????00??????0???????0?????1??????0?1?????110[02]0????1????01 011011?10???00?00??0?000??00?00??0???00?00?11??0?000001101000???????0???????????0?21000102?100111?????1??0? ????1111000?1?12000?0110010000002000002010?01?10001?????010100111????01???21110?????0??????01???000?????10 ??????01?0?000100? Saturnalia_tupiniquim ???????????????????0????0?????????11??????????1??????????????????1????????0??????????1?????????0?????????0010010 01?1???0???0??100001??0000000000001001??1000?111000111??010?????????????????00200001011000100110011001210 0211010002102100111110011?0000200001111010111001111110010101111??0?01???2011110000011101?00?????????????? ?????020000001000

FIRST COELOPHYSOID DINOSAUR FROM ASIA Zootaxa 3873 (3) © 2014 Magnolia Press · 247 Plateosaurus_engelhardti 01001010200101012000001001000100111100000101001000000301000010100121010000001101?20?0111?1100101000110 001001011001111100001001100001001000000001001000011000?11100111100010101010111111200000020000002100010 0110011001210021101000210210001111001100000200000111010111000111110010101111??0?01???201101000001110110 0???????2001010?000001000000[01]000 Efraasia_minor 0?001?10100101012?0000?000000100??11000001?1001????00??1?000????01210100000011??1???011?01??000000?1?000? 001001001?11??010?001100?01000000000000001001?110?0?1110011110001010?01011?111?000?0020000002100010010? ?1100121002110100021021000111100110000020000011101011100011111001010?111??0?01???2011010000011101100???? ??120010?0?00000000000???00 Herrerasaurus_ischigualastensis 20010?00100101001000000000000?0000100000?10100000000030010001?1?012111000000?1??1???001101100??00000?10 0?000000001111000?01001100101??0020000000001011?11?????111011010000011201100110130101002000000210001001 110110112000201010102102100110110011?0000200100101010111001111110010101111??0?01???2011110000011101100?? ?????0001301?0101001000001[01]00 Staurikosaurus_pricei ????????????????????????????????????????????????????????????????????????????????????????????00000?00??00?0000000 01?1???0????01?00?????000000000?00101???????????????????????????????????????002000000210001001110110111?00?0 11?000210110011111001??00002001001000101110001?????????????????????????????????????????0????????????????0??00 10000???0? Chindesaurus_bryansmalli ?????????????????????????????????????????????????????????????????????????????????????????????????????????????????? ?1?????0?????200?1??00??00?0????100????????????????????????????????????????0???0???2?0??100??????1??????????100 02102100110110000?000020011010001?????????????0101?11?1???????????????????????????????????????????????00?0000 00?0? Eoraptor_lunensis 00010?00100112001000000100000?0011111000?10100?0000013001000101?012?1?0?00??????????001?011000?00000???? ?000?00?0101???00??001?0??0?0000200000?000?0?1?110?0?11110110100010111??1001110??1010020000112?00110011?? ?10011000??11??0??1?110?0???100????00020010?1000101?10??11111??10?0??1???0?0????2?11110000?1?1?1?00??????10 102010??00?0110???00?00 Tawa_hallae 1110000000111?0010000?0000000100??110000010100000001?30000?01?1001211?000000?11?1001001?011000000000??0 0?000000001?111?011111111010100??000000?000?011?111????11101101000001120010011012010101200001120001100?1 10?101?1???2??12000220110001210001110000200110100010111000111110010101111??0?01???2111110000010101200??? ???1000?3011?0010110????1?00 Coelophysis_bauri 1110111110111?0020000101100001?010111000010100100001130010?0101001211110000111???00?0011011000?00000?10 0?0000000011111001111111110010000001000111010111111?0?11110110100010112111001111301011121110212111110011 1011?11111021?12000210210011210011110000210111111010111011111010010100111??1?01???21111110100101012?0???? 0??1212210210001201?0000011 Coelophysis_rhodesiensis 1110111111111000200001111000010010111000010100100001?3001000111?012111100001111?1???0011011000020000??00 000000000111110?1?1111111?010000001100111010011111010111101101000101121110011113010211211102121111100111 010011111121012000210210011210011110000210111111010111011111010010100111??1?01???2111111010010101200???? ???111201021?00120101100111 Panguraptor_lufengensis ??????????????00?0000??1101?0?0???1110???10?0?100001130010?0????????????????????????001?01?0???????0?1001?00 000?0??11?0?1?11?1111?0?00??00??00???????1??11??????????0??????11???????111?0?0????????????111?????????????1?? ???1??0?2?021?01??????????0??21011?11?0101110??1?10100???0??11?????1???211111??1?0?0?0120????????1212011000? ??10?0??????? 'Syntarsus'_kayentakatae 1110111110111?002000010110100?00??1110?0010100100000130020?0???????0???0000111??1???001101100??00????1001 000000001?1?10011111111100?0?00??1??01111???11111?10111101?01??0?01????1?????12010?1?2?1102121??1100111011 ?1?111?21?120002?0210011210011110000210111111010110?111?101001010?111??1??1????1111110?00101???0???????102 0320100011?????0000011 Camposaurus_arizonensis ?????????????????????????????????????????????????????????????????????????????????????????????????????????????????? ?????????????????????????????????????????????????????????????????????????????????????????????????????????????????? ?????????????????????????11101???????1?????01010?111??1?01???211???????????????????????????????????????1111111? Liliensternus_liliensterni ??????????????00?0000??1100?0???????????????0?????01?3?020??1110??????????????????????1?????000?0?????00100000

248 · Zootaxa 3873 (3) © 2014 Magnolia Press YOU ET AL. 000111???011??11010001??010?0?00??10???1??1??101111011010001011???1???????????1121110212001110011101101111 ??21?020002101200012100011?00002101111110101110001?????010100111??1??1???21111???000101???0?????????2??201 ??1?101?0000000?? Zupaysaurus_rougieri ??????????????0021000?0110100????11110100100?000000113010000????0?211?00000???????????11011?0??000?0?10??00 0?00?01??????1??????????????????????????????????????????????????????????????????????????????????????????????????? ?????????????????????????????????11101????0??1?????010100111??1??????2?????????????????????????020320111110???? 000000?? Dilophosaurus_wetherelli 11101?11100?1?001100??00101001?0??11110??101000??00113000000????012111000101110??2??001111?00000001001?01 0000000011110000111111200010?0100000011101??1?111?1011110010100010112??10011112010111211002121011100?110 ?1011111121?0200021?120001210001110?002101112110101110?1111010010100111??0?01???2111111000010101200?????? 102033112?0110110000001?1 Cryolophosaurus_ellioti ???????????????0?100???1???0??0??11111101100??000000?3000?001?10?1?????00101????0??????1111??????010?10????0 ??0?01?1??????????12?001000100?????1???0?????????????0???1????????????????????????????02?21?11??0????1???111?? ????2000210?200112100011?0?00??????1??0????????1?????01?10?1????1??1???211??????????????????????1???330??0?01 00?0?0??0??? Ceratosaurus_nasicornis 2000101100010000100001000000010011111110010100120000130000?01?10012111000001111010??001111100??00000?10 010000000011010010111111200010101011??01111?110?10111011??????10???011???10???1??010?01211?0212??111000??? 110??111?21?020002100200010100111?00002100012110?01100100?????02110?111??1???????1110110100101???????????? 02132011000100?01000001? Piatnitzkysaurus_floresi ??????????????001?000?00001001?????????00???????????????????????01211100?11111???2?1??1??1????????????????0000 0001?1?011?01111120001??11001?0011111??1??1??1011110010100????????????????????0?211002021111?00?1101??1111? 121?02000220020001?100011?00002101112110100110101?10100?1???????????????2??1??????1????????????????10?????? ???001000000?0? Allosaurus_fragilis 210010112001011011?00010021001011111111001010000000013000000111001211100011111??02??0011111000000010010 01000000001111011101011120001011100000011111111110101011110110100010112111101111?1??10121100202101110011 0111021111121?02000220020001010001110000210111211010111010111010021100111??0?01???2111011000110101200??? ???1020330110001001010000000 Velociraptor_mongoliensis 01001011100100101110000010100100111111100101001000000310000011100121?1001111111?????0011011000000000?10 0100000000111??11???????????10?01?01001111110?11101?1111100110100000012111000111?1??101211002021011110111 011?20010020102010220?200010100011100002101?12110?01?0010?11???02110?111??1?01????111011000010101200???? ??10103110100?00??000000000