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Home , Heyuannia, Nemegt Formation, Ornithurae, Oviraptorosauria

Bull. Natn. Sci. Mus.. Tokyo, Ser. C. 30, pp. 95130. December 22, 2004

New Oviraptorid (Dinosauria: ) from the of Southwestern

Junchang Lii', Yukimitsu ~omida*,Yoichi ~zuma", Zhiming ~ong~and Yuong -Nam Lee" ' Institute of Geology. Chinesc Acadcrny of Geological Sciences. Beijing 100037. '~ationalScience Museum, 3-23-1 Hyakunincho, Shinjukuku, Tokyo 169-0073, Japan 'Fukui Prefectural Dinosaur Museum, 51- 11 Terao, Muroko. Katsuyama 91 1-8601, Japan "Institute of Paleontology and Paleoanthropology. Chinese Academy of Sciences, Beijing 100044. China ' Korea Institute of Geoscience and Mineral Resources, Geology & Geoinformation Division, 30 Gajeong-dong, Yuseong-gu, Daejeon 305-350. South Korea

Abstract Nernegtia barsboldi gen. et sp. nov. here described is a new oviraptorid dinosaur from the Late (mid-) Nemegt Formation of southwestern Mongolia. It differs from other oviraptorids in the skull having a well-developed crest. the anterior margin of which is nearly vertical. and the dorsal margin of the skull and the anterior margin of the crest form nearly 90"; the nasal process of the premaxilla being less exposed on the dorsal surface of the skull than those in other known oviraptorids: the length of the frontal being approximately one fourth that of the parietal along the midline of the skull. Phylogenetic analysis shows that Nerncgtia ha,-sboldi is more closely related to osrnolskae than to any other oviraptorosaurs. Key words : Nemegt Basin. Mongolia. Nemegt Formation. . Oviraptorosauria. Nernegtia.

dae, and Caudipterygidae (Barsbold 1976; Stern- Introduction berg, 1940; Currie, 2000; Clark e/ al., 200 1; Ji er Oviraptorosaurs are generally regarded as non- a/., 1998; Zhou and Wang, 2000; Zhou el al., avian theropod (Osborn, 1924; Bars- 2000). bold 1976. 1977, I98 1, 1983, 1986, 1997; Gau- The Oviraptorosauria are mainly distributed in thier. 1986; Smith, 1992; Sereno, 1999; Barsbold Mongolia (Osborn. 1924; Barsbold 1976. 1977, Currie et a/., 2000; Barsbold Osmolska et a/., 1981, 1983, 1986, 1997; Osmolska. 1976; Norell 2000; Holtz. 2000. 2001; Norell et a/.. 1994; et al., 1994, 1995: Wester, 1996; Clark et al., Norell et a/., 1995; Norell. Clark, and 1999, 2001, 2002). China (Dong and Currie, Makovicky. 2001; Xu, Norell et a/., 2002). They 1996: Ji et a/., 1998; Lii et a/., 2000; Xu, Cheng are medium-sized theropods characterized by a et a/.. 2002; Lii, 2003) and short. deep skull with toothless jaws in derived (Sternberg, 1940; Cracraft, 197 1; Currie and forms, teeth present in primitive forms such as Russell, 1988; Sues, 1997). Additionally, ovirap- Incisivosa~u~rsgauthieri (Xu, Cheng et al., 2002) torosaurs from the Southern Hemisphere have and Caudipteyiu zoui (Ji et al., 1998). pneu- been reported (Frey and Martill, 1995; Currie et matilized caudal vertebrae present in derived a/., 1996; Frankfurt and Chiappe, 1999), al- forms, anteriorly concave pubic shaft, and a pos- though validity of the taxonomic assignment is teriorly curved ischium present in doubtful. (Barsbold and Osmolska, 1990; Barsbold In the summer of 1996, an incomplete ovirap- Osmolska et a/.. 2000; Makovicky and Sues, torid skeleton with a nearly complete skull was 1998). Oviraptorosauria is here regarded to In- discovered in the Nemegt Formation of Nemegt clude three families: Oviraptoridae, Caenagnathi- Basin, southwestern Mongolia by the Mongolian 96 Lii. Tomida. Azurna. Dong and Lce

Highland International Dinosaur Project team. It nasal process of the premaxilla, which forms part was initially described by Lii et ul. (2002) as In- of the crest, being barely exposed in dorsal view; genia sp. The skull morphology (vertical anterior the anteroposterior length of the frontal is ap- margin of the developed crest on the skull, less proximately one quarter of that of the parietals; a exposed nasal process of the premaxilla, and a presence of prefrontal; process on the quadrate process on the quadrate bearing a convex surface bears a convex surface projecting into the cotyla projecting into the cotyla on the medial surface on the medial surface of the quadratojugal. of the quadratojugal) indicates that it is distinct Mandibular condyles of the quadrate situated from named oviraptorosaurs. It represents a new rostrally to the occipital condyle. taxon from the Late Cretaceous. Description Systematic Paleontology I. Skull and mandible Marsh, 1 88 1 The skull (Fig. I) is deep, narrow and short, Oviraptorosauria Barsbold, 1976 and has a naso-premaxillary vertical crest. The jaws are toothless, and the articular has a medial Family Oviraptoridae Barsbold. 1976 process. These characters indicate that Nemegtia Nemegtia gen. nov. barsboldi is a derived oviraptorosaur. Diagnosis: As for the only known species. Skull: The premaxillae, vomers and maxillae Etymology: Nemegtia refers to the lo- form a hard palate as in other oviraptorid di- cality. the Nemegt Basin of southwestern Mon- nosaurs (Fig. lC), and the premaxillae form the golia. main part of the crest. Both premaxillae are fused. The central palatal part of the joined pre- Nemegtia barsboldi sp. nov. maxillae is strongly concave. There is a cleft on (Figs. 1, 3, 5,6) the central part of the palate. Anteriorly, small foramina distributed on the lateral sides of the lngenia sp. Lii rt 01.. 2002 suture between the premaxillae may represent Etymology: The species is named in honor of nutrient openings. The nasal processes of the pre- Dr. R. Barsbold, the Mongolian pale- maxillae are perpendicular to the palatal surface ontologist, one of the leaders of the Mongolian of the upper jaw, and extend dorsally, connecting Highland International Dinosaur Project. with the nasals at the highest point of the crest 7jpe locality and horizon: The Nemegt lo- (Fig. 1 A), and is barely exposed in dorsal view of cality of Gradzinski et al. (1968), Nemegt Basin, the skull. The crest extends posteriorly and ven- southern , Mongolia; Nemegt Forma- trally, and forms a round arc at its highest point. tion, mid-Maastrichtian (Jerzykiewicz and Rus- The antorbital cavity consists of a large inter- sell, 1991) of the Upper Cretaceous. nal antorbital fenestra posteriorly and a small Holoppe: GIN 1 0012 1 12 (formerly PC 1001 maxillary fenestra (see Witmer, 1997) anteroven- 21 12, see Lii et al., 2002): an incomplete skele- trally. The internal antorbital fenestra is sub-oval, ton with a nearly complete skull. Specimen and the long axis is vertical. The external surface stored in the Paleontological Center of the Mon- of the maxilla is smooth, and moderately con- golian Academy of Sciences. Ulan Bator, Mon- cave. Ventrally, there are two longitudinal, round- gol ia. ed ridges on each maxilla as in other ovirap- Diagnosis: An oviraptorid displaying the fol- torids. In addition to the lateral ridge, there is a lowing derived characters. The skull with a naso- longitudinal groove. Posteriorly, there is a ven- premaxillary crest, anterior margin (the nasal trally directed tooth-like process on the palatal process of the premaxilla) nearly vertical; the shelf of each maxilla as in other oviraptorids. Ncw Oviraptorid Dinosaur. Mongolia 97

antorb

Fig. I. The skull of A'etneg~inhnr-sholdi gge. et sp. nov. (G11\!100/21 12). In lateral (A), dorsal (B), and ventral views (C'). Scalc bar=5 cm. Abbreviations, as in appendix I. 98 Lii. Tomida. 4zuma. Dong and Lcc

Both nasals are completely fused posteriorly The postorbital process of the jugal inclines pos- with no trace of the suture. Anteriorly, the nasals terodorsally. and forms almost 213 of the height are not fused forming a "<" shaped cleft, which of the postorbital bar. similar to other ovirap- receives the nasal processes of the premaxillae torosaurs (Barsbold et al., 1990). The process be- (Fig. 1 B). The dorsal portion of the lacrimal is comes widened and slightly twisted and its me- crescent shaped in lateral view. The shaft of the dial surface is smooth, near the position where it lacrimal is stout, and forms the posterior margin contacts the descending process of the postor- of the antorbital cavity. There is a large lacrimal bital. The dorsal part of the postorbital process of foramen in the middle of the anterior edge of the the jugal is tightly attached to the posteroventral orbit. The posterodorsal part of the lacrimal is surface of the postorbital. The posterior branch wedged between the prefrontal and frontal. The of the jugal contacts the anterior process of the lateral surface of the lacrimal is smooth. Pos- quadratojugal. The angle of the postorbital terodorsal to the antorbital cavity, there is a large branch and posterior branch of the jugal is 90". recessed pneumatic pocket (6 mm high. 4 mm which form the anterior and ventral margins of wide), similar to that in Citipati osmolskae the large lateral temporal fenestra, respectively. (Clark et 01.. 2002). The width of the frontal is much greater than Anterodorsal to the left orbit, there is a small its length. In lateral view, the whole central part triangular bone, which is located between the of the frontal surface is convex, thus the trans- frontal and lacrimal. It should be a prefrontal, al- verse cross-section of the frontal is arc-shaped. though this bone has not been reported in other The anteroposterior length of the frontal is ap- oviraptorosaurs. proximately one quarter of that of the parietals. The jugal is a triradiate bone. The posterior Anteriorly, the suture of the frontal and the nasal part of its anterior branch is stout and plate-like. is relatively straight. while posteriorly, the suture Anteriorly. it gradually becomes rod-like and with the parietal is convex anteriorly (Fig. 19). then ends as a thin plate (Fig. 2). where it inserts There is a deep groove along the suture with the between the maxilla, lacrimal and ectopterygoid. nasal. When compared with other oviraptorids.

cross-section of the jugal

Fig. 2. Skull of Ncnqtiu htr~vholdigen. et sp. nov. (GIN 100i21 12) showing the structure of the jugal and the relationship betwcen the quadrate and the quadratojugal. New Oviraptorid Dinosaur. Mongolia 99 the frontal is very short medially and widens lat- In ventral view, the vomer has a sharp process, erally towards orbital margin. which projects ventrally. This tooth-like process The parietals are fused. Their dorsal surface is is also formed in part by processes of the maxil- smooth. A sagittal parietal ridge is present. This lae. The vomers are completely fused anteriorly. ridge extends anteriorly and disappears at the The posterior parts of the vomers are separated level of the anterior corner of the supratemporal from each other by the cultriform process of the fossa, not reaching the frontoparietal suture. parasphenoid which wedges rostrally between The right squamosal is almost completely pre- the vomers as in all oviraptorids. The middle part served. It curves posteriorly. There is a notch, of the vomer is relatively long, its surface is which holds the distal lateral head of the smooth, and its dorsal part is fused with the ante- quadrate on the middle of the lateral margin. The rior part of the cultriform process. The posterior anterior process of the squamosal forms the pos- part of the vomer is covered by the anterior part tolateral margin of the sub-oval supratemporal of the palatal ramus of the pterygoid. The fused fossa. The long axis of the suboval supratemporal part of the vomer projects ventrally, and the mid- fossa points anterolaterally, making an angle with dle part has a round ventral surface, while its the long axis of the skull of nearly 30". posterior part is relatively flat. The postorbital is a triradiate bone. The frontal The palatine forms the posterior and lateral process of the postorbital is upturned about 90 margins of the elliptic choana. It connects with degrees. Its descending process occupies 213 of the vomer medially, and with the maxilla antero- the height of the postorbital bar. The anterior sur- laterally. The posterolateral part of the palatine face of this process is smooth. The angle of the contacts the anteromedial side of the ectoptery- anteromedial process and the posterior squamos- goid and its posteromedial part contacts the a1 process is greater than 90". The frontal process palatal ramus of the pterygoid. covers the orbital process of the frontal. The Ventrally, the pterygoids have elongated squamosal process extends posteriorly with a palatal processes, which do not meet along the sharp dorsal margin and overlies the anterior middle line until they contact the vomers. Each process of the squamosal. This process does not palatal process has a longitudinal groove. which reach the posterolateral corner of the supratem- divides the palatal branch of the pterygoid into poral fossa. The upper part of the postorbital bar two parts anteriorly: the lateral part connects forms most of the anterolateral margin of the with the ectopterygoid, while the medial one con- supratemporal fossa. nects with the palatine and quadrate. A shallow groove separates the articular sur- The ectopterygoid is located rostra1 to the face into two parts in the quadrate. In ventral pterygoid. It extends vertically, similar to other view, the width of the mandibular articular sur- derived oviraptorids, whereas in other theropods, face of the quadrate is greater than its length. it extends horizontally. The posterior end of the The mandibular condyles are situated rostrally to dorsal part disappears near anterior end of the the occipital condyle. The articular surface for jugal. There is an opening within ectopterygoid the quadratojugal is convex. while that on the which is close to the maxilla. This opening may quadratojugal for the quadrate is concave. This be homologous to the dorsal ectopterygoid recess joint forms a mobile junction (Fig. 2), which is of dromaeosaurids (Witmer, 1997). similar to the condition in most and other The epipterygoid is present on both sides of oviraptorosaurs (Maryanska and Osmolska, the skull. It is a thin plate-like bone with an ir- 1997). The convex quadrate and concave quadra- regular triangular shape, whose free vertex angle tojugal joint is reversed from the condition in projects dorsally. It tightly ties to the pterygoid other oviraptorosaurs (Maryanska and Osm6lska, branch of the quadrate. Its anterior margin is 1997). sharp and free. 100 Lii. Tomida. Azuma. Dong and Lee

The basioccipital condyle is ball-shaped with ternal mandibular fenestra, as in other ovirap- a weak neck region. The articular surface of the torids. The ventral surface of the lower jaw is occipital condyle faces posteroventrally. The an- convex, and the mandibular symphysis is short. terior part of the basioccipital projects ventrally In dorsal view, the jaw slightly prqjects laterally and laterally, forming the posterior part of the in the middle part, and both branches of the basal tubera. The suture of the basioccipital and lower jaw are parallel posteriorly from the level basisphenoid is clear. Its middle part projects of the posterior margin of the mandibular symph- posteriorly; therefore, in the ventral view, the su- ysis (Fig. 3A). The dentaries have a dorsal pro- ture has a weak "W" shape. trusion near the posterior margin of the mandibu- The basisphenoid forms the posteroventral part lar symphysis. The large external mandibular of the floor of the braincase. The sutures with the fenestra is located at the anterior part of the pterygoid and the parasphenoid are not clear. In lower jaw. The coronoid process is posterior to ventral view, it is narrow anteriorly and broad the mandibular fenestra. posteriorly. The portion comprising the basal tu- The mandibular symphysis is short and deep. bera is strongly developed. Its middle portion is No evidence shows that a coronoid is present in strongly depressed and has a large opening. This the present specimen. CT scanning shows that opening is probably the Vidian canal for the pas- the dorsal margin of the mandibular symphysis is sage of the internal carotid artery and the palatal thin, while its lower margin (lower part) is stout, branch of nerve VII, which entered the pituitary the symphysis is highly pneumatized and the fossa through the basisphenoid. The basiptery- spaces are uniform in size and distribution (Figs. goid process is absent, similar to other derived 4A, B). There is no bony construction between oviraptorosaurs (Clark et ul., 2002). the hollow spaces of the right and left dentaries. The tall parasphenoid process is delicate. It is The cross section near the mandibular symphysis shallow anteriorly and deep posteriorly. Its depth shows that both sides are steep, displaying a "V" gradually lessens anteriorly, as it connects with shape. It is "U" shaped in Cuenagnuthus and the vomer. Its posterodorsal part extends dorsally there is a bar of bone between the dorsal and ven- between the epipterygoid and pterygoid contact- tral surface supporting the midline ridge on the ing the laterosphenoid. In ventral view, it extends lingual surface of the symphysial shelf (Currie et forward into the large space called interpterygoid a/., 1993; Figs. 4C, D). Around the anterior mar- vacuity. Its length is about 114 of that of the skull. gin of the external mandibular fenestra, the den- The laterosphenoid is a thin plate-like bone, taries are deflected inwards, forming a shallowly anterior to the floor of the braincase. The or- flattened surface, thus producing a thin and sharp bitosphenoid is also a thin plate like bone, anteri- anterior edge on the external mandibular fenes- or to laterosphenoid portion of the floor of the tra. The ventral process of the denary is relatively braincase. weak, and it tightly extends along the ventrolater- The suture between the exoccipital and al part of the anterior portion of the angular, and supraoccipital is not clear, but the trend of the su- disappears at the level of the coronoid eminence. ture shows that both exoccipitals connect above The dorsal process of the dentary is comparative- the oval foramen magnum. Thus, they separate ly massive, and dorsally, its distal end is divided the supraoccipital from the foramen magnum, into lateral and medial prongs, with the anterior which is larger than the occipital condyle. branch of the articular-surangular-coronoid com- Mandible: The lower jaw is short and deep plex (ASC) wedged between them. It disappears (Fig. 3). The dorsal margin of the external anterior to the coronoid eminence. The ventral mandibular fenestra is formed by the surangular process of the dentary extends more posteriorly and the dorsal arch of the dentary. The surangular than that of the dorsal process. Only the external has a middle branch, which projects into the ex- surfaces of both processes of the dentaries are New Oviraprorid Dinosaur. Mongolia 101

asc

.*.... 1 emf

3. Lower jaw ofrl'c.nwg/io bnrsholdi gen. et sp. nov. (GIN100121 12). In dorsal (A), ventral (B), and latcral (C) views. Scale bar=2 cm. Abbreviations as in appendix I. Lii. Tomida. Azuma. Dong and Lcc

Fig. 4. Comparison CT scans or the corresponding positions or the lower jaws of Nemrgriu (GIN10012 1 12) and . A, C: cross section through anterior part of the symphyseal region; B. D: cross section through the posterior portion or the symphyseal region. Scale bar= I0 mm (C and D are chosen from Currie et a/.,1993, Fig. 7).Abbreviations as in appendix 1. smooth. Anterior to the external mandibular fen- There is a longitudinal rounded ridge on the ar- estra and the lateral edges of the mandibular ticular surface. This ridge corresponds to the lon- symphysis, irregular small holes are densely dis- gitudinal sulcus on the articular surface of the tributed. A row of four small foramina is sym- quadrate condyle. The ridge separates the articu- metrically distributed on both sides of the serrat- lar surface into two parts: a medial one, called ed suture of the mandibular symphysis. This row the internal mandibular process by Sternberg of small foramina extends posterolaterally, form- (1940), is large and semicircular, and a lateral ing a nearly 30" angle with the serrated symphy- one, called the external mandibular process seal suture. The lateral surface of the ventral part (Sternberg, 1940). is smaller and sub-oval. The of the mandibular symphysis is convex. and there surface of the lateral one slightly slants laterally. is a process centrally. The lengths of both processes are nearly equal. The surangular and articular are fused in the The retroarticular process is slender, extending posterior part of the lower jaw, with no clear su- posteroventrally from the articular surface. Its ture between them. The fused part occupies the ventral and medial surfaces are covered by the position, which corresponds to the position of the prearticular. Its lateral surface is covered by artic- articular, surangular, and coronoid in other ular-surangular-coronoid complex. The angular theropods. Currie et al. (1993) named it as the does not cover the prearticular, and it projects articular-surangular-coronoid complex (ASC). posteromedially. The articular-surangular-coro- New Oviraptorid Dinosaur. Mongolia 103 noid complex occupies the major part, posterlor smooth. It becomes thin and sharp poster~orly, to the posterior margin of the external mandibu- and disappears at the level of 11'3 length of the lar fenestra. There is a deep. longitudinal groove lower jaw from the retroarticular process. on the lateral surface of the anteroventral part of the art~cularbone. This depression should be the 2. Postcranial skeleton ~nsertionarea for the external mandibular adduc- A nearly complete cervical series, part of the tors. Thc coronoid process projccts posteromedi- dorsal vertebrae, a nearly conlplete sacrum, both ally. ilia, the proximal ends of the and ischium, The angular wedges anteriorly between the thc proximal end of the femur, parts of the splenial and the ventral process of the dentary, humerus, and the complete sight radius are pre- and disappears anteriorly to the anterior margin served. of the extcrnal mandibular fenestra. Medially, the Cervical series: Thc cervical series (Fig. 5) suture with the preart~cularis cleal; but, laterally, includes 12 of 13 vertebrae naturally articulated the suturc with the surangular is not clear. (including a small part of the axis). .4lthough The prcarticular is located at the ventromedial somctimes, it IS difficult to distinguish the poste- surface of the articular. It 1s a thin plate-like rior cervical vertebrae from the anterior dorsal bone, which covers a part of thc medd surface vertebrae, the presence of a weak median ventral of the lower jaw. keel and a hypapophysis on the last of the natu- The splenial IS strap-like, widcr anteriorly than rally articulated 13 vertebrae indicate that this posteriorly. It contacts the dentary anteriorly, bul last should be the first dorsal vertebra. ~t does not take part in the formation of the Thus, with the atlas, the number of the cervical mandibular symphys~s. Its medial surface is vertebrae is 13. The measurements of the pre-

Fig. 5. The cervical vertebrae and the antcrior dorsal vertebra (the last one) of Ncmcgiin bnrsboldi gen. el sp. nov. (GIN100/2 1 12). In lateral view (A). and dorsal view (B). Scale bar-Scrn. Abbreviations as In appendix I. 104 Lii, Tomida. Azuma. Dong and Lee

Table I. The measurements of the 12 (including part of axis but not thc atlas) articulated cervical vertebrac and the first dorsal vertebra (mm).

C2 C3 C4 C5 C6 C7 C8 C9 ClO C11 C12 C13 Dl

Length of the centrum ? 33 29 32 31 31 33 35 37 37 36 33 3 1 Width of the centrum ? 15 27 30.5 39 39 40 42* 42* 43 48 52 52*

"*" represents estimation. "'? missing served 13 vertebrae are given in Table I. the position of the neural arch is much more for- Only the posterior part of the axis is preserved ward. and it is fused with the third cervical vertebra. The distance between the prezygapophyses Two postzygapophyseal facets are preserved and and that of the postzygapophyses of the 6th cer- the distal end of the postzygapophysis is relative- vical vertebra is nearly equal to the length of the ly sharp. There is no epipophysis on the dorsal centra, thus forming a square outline. This is dif- surfaces of postzygapophyses. ferent from that of other centrum in that the ante- The anterior end of the 3rd cervical vertebra is rior and posterior widths are equal in dorsal view, almost twice as wide as the posterior end. The the diapophyseal facets are larger than that of the central part of the lateral surface is strongly de- anterior ones, but smaller than that of the postcri- pressed with a well-developed elongated pleuro- or ones. coel. In the lateral view, the centrum is wedge- In dorsal view, the distance between the prezy- shaped. There is a weak epipophysis on the pos- gapophyses of the 7th cervical vertebra is longer terodorsal surface of the postzygapophysis. The than that of the postzygapophyses. The lateral posterior articular end of the centrum is concave. surface of the centrum is concave, with a well- The neural arch of the 4th cervical vertebra is developed pleurocoel on the central part. The an- obviously larger than that of the 3rd centrum. but terior margin of the pleurocoel is sharp, its pos- the length of the centrum is slightly shorter than teroventral margin is not clear. The size, position the anterior one. From the lateral view, the anteri- and the morphology of epipophysis are similar to or articular end slants anterodorsally at 45O, cor- that of the 6th cervical vertebra. responding to the posterior end of the anterior The neural arch of the 8th cervical vertebra is connected centrum. The anterior margin of the obviously smaller than those of the anterior ones. prezygapophysis slightly outruns the anterior The distance between the prezygapophyses is margin of the centrum, whereas the posterior slightly greater than that of the postzygapophy- margin of the postzygapophysis is at the level of ses. The epipophysis is relatively small and locat- the posterior end. ed anteriorly. The facet of the postzygapophysis In dorsal view, the distance betwcen the faces posteroventrally. postzygapophyses of the 5th cervical vertebra is In the 9th to 13th cervical vertebrae, the size greater than that between the prezygapophyses. of each vertebra is nearly equal. The position of The epipophysis is well developed. The anterior- pleurocoels on the lateral surface of the centra ly slanted spine is much lower than the previous gradually moves posteriorly from the 9th cen- one. The anterior margin of the prezygapophysis trum to the 13th centrum. It arrives at the central extends far beyond the margin of the anterior end part of the centrum in the 12th cervical vertebra. of the centrum. The posterior margin of the and the pleurocoel is oval. The radius of the pos- postzygapophysis is located anterodorsally to the terior margin of the pleurocoel is larger than posterior articular end of the centrum. Compared those of the anterior ones, while the radius of the with the 4th vertebra, the relative position of the anterior margin of the pleurocoel is larger than neural arch on the centrum is greatly changed the posterior one in the anterior vertebrae. There New Oviraptorid Dinosaur. Mongolia 105 are clear fossae posteroventral to the prezy- while the rib tubercle is not fused with the cen- gapophyses of the 1 Ith and the 12th cervical ver- trum. The rib tubercle of the 5th cervical rib is tebrae. The infradiapophyseal fossa is reduced in smaller than that of the 6th cervical rib. the posterior vertebrae. There is no infraprezy- In the 7th to 12th cervical ribs, the rib tubercle gapophyseal fossa. The distance between the and rib head are completely fused with the di- prezygapophyses of the five vertebrae is equal to apophyses and parapophyses of the centra. The that of the postzygapophyses, but this distance is tubercle of the 13th cervical rib is clearly not shorter than the length of the centrum. Therefore, fused with parapophysis, and its morphology is their outlines are rectangle. The facet of the similar to the anterior ones. postzygapophysis faces postolaterally. The poste- Do,sul ver-tehrme: Only the anterior one and rior part of the ventral surface of the centrum is a half dorsal vertebrae, and three posterior dorsal flat and slightly expanded. vertebrae and two neural arches, which articulat- The neural spines of the cervical vertebrae are ed with the sacral vertebrae, are preserved. The short and centered on neural arch, giving neural count of dorsal vertebrae is uncertain. arches an "X" shaped appearance (Fig. 5B), as The pleurocoel in the 1st dorsal vertebra is that in , Microvenutor celer-, and much larger than those of cervical vertebrae. The other oviraptorids (Makovicky and Sues, 1998). distance between the postzygapophyses becomes The vertebrae in the middle of the cervical series shorter. There is a weak ventral keel. A clear pro- are the largest (5th, 6th and 7th). The facets of jection on the middle part of the ventral margin the postzygapophysis from the 3rd to 8th cervical of the anterior end of the vertebra represents the vertebrae face posteroventrally, while those from hypapophysis. According to Sues (1997). the the 9th to 13th vertebrae face posterolaterally. An presence of the hypapophysis generally appears infradiapophyseal fossa appears in the 1 1 th and to be restricted to the first three or four dorsal 12th cervical vertebrae. In dorsal view, the out- vertebrae in non-avialan theropods. Compared line formed by the four zygapophyses in the sixth with the cervical vertebrae, the first dorsal verte- cervical is nearly square, it is rectangle (the angle bra is short and high, and the posterior articular formed by the right-left prezygapophysis with the end is flat. The spine is triangular in anterior and centered neural arch is greater than 90") in the posterior views. The anterior margin of the 4th and 5th cervical vertebrae, but the angle is prezygapophyseal facet approaches the middle of less than 90" after the seventh cervical vertebrae. the anterior vertebra, unlike that of the cervical There are also variations in the pleurocoels. They vertebrae. Only the anterior part of the 2nd dor- are relatively small in anterior vertebrae. The an- sal vertebra is preserved and the anterior articu- terior margin of the oval pleurocoel is larger than lar surface is strongly concave. that of the posterior margin from the 3rd to 9th On the five preserved posterior dorsal verte- cervical vertebrae, whereas the pleurocoel of the brae, only the neural arches of the anterior two 10th is circular. From the I lth to the last cervical posterior dorsals are preserved. The base of the vertebrae, the posterior margin of the pleurocoel transverse process is wider than on posterior ver- is larger than its anterior margin. tebra. The pleurocoel in the middle two is large Almost all the cervical ribs are preserved. The and circular. The neural spine is wide, thin and axis rib is slender, rod-like and it is the longest plate-like. Its width is nearly 213 of the length of one among the cervical ribs. It extends to the an- the vertebra. terior end of the 4th cervical rib. The 3rd cervical The last dorsal vertebra is located medial to rib is also rod-like. longer than the length of the the anterior end of the ilium, and has a free rib, 3rd centrum. but its transverse process contacts the ilium. The The 5th and the 6th cervical ribs are very simi- ventral surface of the centrum is round. There is lar. The rib head is fused with the parapophysis, a large pleurocoel in the middle of the lateral sur- 106 Lii. Tomitla, AZLI~.Dong and Lce

Fig. 6. Sacrum and pclvic girdle of ~Vetnegtiahaisbold; gen. el sp. nov. (GIN100121 12) face of the centrum. The posterior articular end part and its neural arch. Whether their spines are 1s not fused to the first sacral vertebra and the ar- fused together or not is uncertain, due to their ticular surface is slightly concave. being covered by the matrix, but it is clear that Sacral vertebrae: There are 8 sacral vcrte- thc distal ends of the spincs do not rise above the brae (Fig. 6). The anterior six are completely pre- dorsal marglns of the ilia. The measurements of served, the 7th sacral vertebra is partly preserved, the sacral vertebrae are given in Table 2. and the 8th has preserved only a small antcrlor The width of the anterior end of the 1st sacral New Oviraptorid Dinosaur. Mongolia

Table 2. The measurements of the sacral vertebrae (mm).

SI S2 S3 S4 S5 S6 S7 S8

Length of the ccntrum 32 33 29 29 29 32 40 ? Widthoftheccntrum 22 23 22 20 21 21 ? ?

vertebra is greater than its posterior end. The an- inwards, while the postzygapophysis faces later- terior articular end is strongly concave. The ven- ally. tral surface of the centrum is smooth. The 2nd und the,/iwe linzh: The proxi- sacral vertebra is similar to the Ist, except that mal end of the left is preserved. It is sim- the pleurocoel is elongated and enlarged and it ilar to those of other oviraptorosaurs. Distal parts occupies nearly 112 the length of the centrum. of both humeri are preserved (Figs. 7A. B). The The width of the anterior end of the 3rd centrum preserved portions are similar to other ovirap- is larger than its posterior end while in the 4th torosaurs. but there is a fossa on the anterior sur- and 5th centra, the width of both ends is equal. face (Fig. 7A, fmb) occupying a similar position There is a shallow groove on the anterior portion to the,fixsu m. hruchiulis (Baumel and Witmer, of the ventral surface in the 4th centrum. This is 1993) in modern birds. This is not observed in clearly different from other centra. The pleuro- other oviraptorosaurs. coel becomes smaller, and the sacral rib of the The right radius is completely preserved (Figs. 4th vertebra is located at the same level as the is- 7C, D). It is straight, and the cross section ofthe chiadic peduncle. The anterior articular end is shaft is oval. Both ends are slightly expanded but larger than the posterior in the 6th centrum. and the proximal is larger. the contact area of the rib with the centrum is the Pelvic girdle und the hind linzh (Fig. 6): The largest among the sacrals, occupying nearly the preserved length of the ilium is 280 mm. In later- whole length of the centrum. Only a small part of al view. the dorsal margin of the ilium is straight, its anterior end is fused with the posterior end of and the depth of the preacetabular process is the the 5th centrum. The small pleurocoel is located same as that of the postacetabular process. The on the mid-posterior part of the centrum. The anterodorsal part of the preacetabular process is distal end of the 7th sacral rib is divided into two concave, while its ventral part is convex. The parts, which contact the ilium. Posterior to where pubic peduncle is slender and projects down- the sacral rib is fused with the sacral centrum, wards and its articular end is triangular. The is- there are two fenestrae, which are nearly parallel chiadic peduncle is relatively stout with a round to each other, and they are much larger than the articular end. Dorsally, the ilia nearly meet. but anterior pleurocoels. The right sacral rib of the are not fused. The same condition is also found 8th sacral vertebra is fused with the transverse in other okiraptorosaurs, such as Non~ingiugohi- process of the centrum and is nearly rod-like. ensis (Barsbold, Osmolska et ul., 2000). The middle of the rib shaft is relatively slender. Only the proximal part of both ischia is pre- but both ends are widened. Its distal end tightly served. The process that contacts the pubis is contacts the internal surface of the ilium. The smaller than that which articulates with the ilium. preserved part shows that the fusion with the The proximal ends of both pubes are preserved. connected centra is weaker than that of other cen- Only the proximal end of the femur is pre- trum. served. There is a clear neck between the femoral Cu~idulvertrhrue: Only the neural arches of head and the shaft of the femur. The angle of thc the first two caudal vertebrae are preserved. The neck and the shaft is about 90". There is a shal- nearly vertical facet of the prezygapophysis faces low groove between the greater trochanter and Lii. Tomida. Azuma. Dong and Lee

Fig. 7. The left humerus in anterior view (A)and posterior view (B)and the right radius in medial view (C) and lateral view (D) of Nernqyitr huncsholdi gcn. ct sp. nov. (GIN100121 12). Scale bar=4cm. Abbreviations as in appendix I. the femoral head. The lesser trochanter is smaller the crest is located above between the lacrimals and fingerlike. It connects tightly with the greater in 0. mongoliensis, although the anterior margin trochanter. The broken surface shows that many of the crest is nearly vertical (Barsbold and bony struts are distributed near the lateral wall of Osmolska, 1990). Dorsally, the premaxillae are the shaft. barely exposed and the anteroposterior length of the parietal is greater than the frontal in Nemeg- Comparison and Discussion tia harsboldi, but the exposed nasal process of the premaxilla is much larger than the nasals, and The high, narrow and short skull with tooth- the lengths of the parietal and the frontal are less jaws and the medial process of the articular nearly equal in 0. philoceratops (Barsbold and indicate that Nemrgtia barsholdi is a derived ovi- Osmolska, 1990). An additional opening appear- raptorosaur (Barsbold and Osmolska, 1990; ing on the wall within the antorbital cavity in Ne- Barsbold 1997). Nemegtia is different from Ovi- megtia barsholdi is smaller than that of the 0. raptor- in the skull crest. The anterior margin of philoceratops. The posterior margin of the the skull crest is vertical and the highest point of quadrate condyle is distinctly posterior to the an- the crest is located between the nasals and pre- teroventral margin of the occipital condyle in 0. maxillae in Nemrgtia harsholdi. The dorsal ante- philoceratops. The shape of the ilium, and the rior margin of the premaxilla projects forward number of the sacral vertebrae are also different and the ventral anterior margin of the crest is betwccn Nemegtia and . concave posteriorly in the specimen GIN 100142, Nenwgtia hursboldi has eight sacral vertebrae which was described as Oviraptor- philoce~ntops whereas evunshini has six or seven. Ne- by Barsbold et al. ( 1990). The highest point of megtia havsholdi has a distinct crest in contrast New Oviraptorid Dinosaur. Mongolia 109 to lngenia junshini. The ischiadic peduncle of skull. the occiput and the quadrate sloped an- the ilium is relatively stout, and the pubic pedun- terodorsally in Citiputi osmolskue (Clark el al., cle is relatively weak in Nemegtia bur:sboldi, as 2001; 2002), the width and the length of the cer- opposed to the condition in lngmia uvunLshini. vical vertebrae are not so different in Nemegtiu Nenwgtia bur-sholdi is different from Con- hursholdi, but the length of the cervical vertebrae choraptor gracilis in having a developed crest, is about two times longer than their width in Citi- while the skull has no crest in gra- puti osmolskue. The phylogenetic analysis shows cilis. that Nemrgtiu bursholdi is more closely related Nemegtiu hursholdi is different from Citipati to Citipati osmolskue than to any other ovirap- osmolskue, in that the convex crest is higher than torosaurs (Fig. 8). the dorsal surface of the skull, and the occiput Nemegtia hursboldi differs from Khaun and the quadrate are vertical, while the top of the mckennai (Clark rt ul., 2001) in having both a skull crest is below the dorsal surface of the distinct skull crest and a parietal crest, which are

Allosauroidea

Archaeopteryx lithographica

A vimimus portentosus

Chirostenotes pergracilis

Nomingia gobiensis

"Oviraptor" mongoliensis

lngenia yanshini

Conchoraptor gracilis

Heyuannia huangi

Nemegtia barsboldi

Citipati osmolskae

GIN 1 00142

Oviraptor philoceratops

Khaan mckennai

Microvenator celer

Caudipteryx zoui

lncisivosaurus gauthieri

Fig. 8. One most parsimonious tree found by PAUP. 110 Lii. Tomida. Azuma. Dong and Lee lacking in Khuun. The dorsal process of the pre- margin of the antorbital cavity (Fig. 9). such as in maxilla is vertical in Nemegtiu hucsholdi, but it 1ncisivosaurv.s guuthieri. In Cuudipteiy sp. extends posterodorsally in Khuun mckennui, the (IVPP V 12430, Zhou et al., 2000). although the jugal extends further anteriorly in Nemegtia haw skull is heavily crushed it still can be inferred holdi, but it extends further posteriorly in Khaun that the dorsal margin of the external nasal open- mckennui. ing is below the dorsal margin of the antorbital Nemegtia hursholdi is different from Nomingiu cavity. Derived forms have the dorsal margin of gohiensis in that having eight sacral vertebrae, the external nasal opening above the dorsal mar- compared to five in gohiensis; the dor- gin of the antorbital cavity, a larger distance sal margin of the ilium is straight in Nemegtiu being more derived. This condition in Nemegtiu hursholdi, but convex in Nomingia gohiensis; the harsholdi shows that Nemegtiu hursholdi is a pubic peduncle is more slender and weaker in more derived oviraptorosaur. Nemegtiu hursholdi, compared to Nomingiu gob- The morphology of the lower jaw of Nemegtiu iensis. Nomingiu has a (Barsbold Cur- shows that it is similar to those of derived ovirap- rie et ul., 2000). torosaurs, Oviruptor phi1ocemfop.s and 0. won- Nentegtiu bar-sholdi differs Microvmafor celer goliensis, Ingeniu yanshini (Barsbold et a/., in having one pleurocoel on the cervicals and hy- 1 !NO), Citipuri o.smolskue (Clark el ul.. 2002). pophysis on the ventral surface of dorsal verte- Khuun mckennui (Clark et ul.. 2001). and brae in Nemegtiu, compared with two pleuro- Hg:l,uunniuhuungi (Lii. 2003). in which the lower coels and no hypophysis in Micramutor cder: jaw is short and deep, the upper margin of the ex- the number of sacral vertebrae is at least 8 in Ne- ternal mandibular fenestra is formed by the megtia hursholdi, it is less than 6 in Micrownu- surangular and the dorsal branch of the dentary, tor celer. the surangular has a middle branch, which pro- Nemegtiu hur.sholdi differs from Cuudipter:w jects into the external mandibular fenestra. The zoui in that Nernc>gtiubur-sholdi has no premaxil- ventral surface of the lower jaw is convex. and lary teeth, while Cuudiptenx zoui has premaxil- the mandibular symphysis is short. This is differ- lary teeth. Nemegtiu has a well-developed crest ent from that of Cuenugnuthus collinsi (Stern- in the skull, but the skull of Cuudipter:w zoui berg. 1940; Cracraft. 1971; Currie et ul.. 1993). lacks a crest. Additionally, the pubic peduncle is where the lower jaw is slender and shallow, the slender and weaker than ischiadic peduncle in mandibular symphysis is also longer, the external Nemegtiu hursholdi, but it is stronger in mandibular fenestra is elongate with a dorsal Cuudipter;~~,~zoui. margin, wHich is formed by the surangular, and Nemegtiu hursholdi differs from the surangular had no middle branch. Inci- huungi (Lii, 2003) in that the articular surface for sivo.suurus guuthieri (XU, Cheng et 01.. 2002) the quadratojugal is convex, while it is more also has a longer lower jaw. similar to that of groove-like in Hqvuunniu huangi. Nemegtiu Cuenugnuthus collinsi, but it bears teeth. It hursholdi has a developed skull crest. whereas may represent a primitive caenagnathid ovirap- Hqaranniu huungi does not. In the lateral view. torosaur. the dorsal margin of the ilium is straight through- Eight sacral vertebrae are present in Nemegtiu out its length. while it is convex in Hejwunniu hur.sholdi, similar to some derived but huungi. more than most known non-avian theropod The relative position of the external nasal dinosaurs. More basal theropods. such as opening and the antorbital cavity varies in differ- Compso,qnuthu.s longipes (Ostrom, 1978). and ent oviraptorosaurs. The apparent primitive state Dilophosuzrrzrs wetherilli (Welles, 1984) have is with the dorsal margin of the external nasal four sacrals, five in the relatively advanced opening below or near the level of the dorsal theropods-Ceruto.su~rrz~.s(Gi lmore, 1920). Al- New Oviraptorid Dinosaur. Mongolia 111

Fig 0 Skulls of (A) Inciti~~oso~irusg(~cru/l~ieri. (B) GIN 100121 12. (C) Khuon tr/ckeunai (from Clark ct ul.. 2001). (D) Oviruplor philocevotops (revcrsed. from Clark c.1 01.. 2002). (E) C'UI~IO~O~~O~gi.ircilis, (FI "01:i- /.iip/oi"' 111011go1ic~nsi.s.(GI Citipcili o.sn~ol.skuc~(from Clark er a/.. 2002), and (H) Ovirupfor pkilocerrr~ops (GIN 100142) (from Barsbold. 1986). Scale bar=5 cm in A. B: E and F, 2cm in C and D. 4cm in H. 112 Lii, Tomida. Azuma. Dong and Lee losau~*zu,/ragiis (Madsen, 1976). Nomingia gob- crovenator celer and Troodontidae were exclud- iensis (Barsbold Osmolska el a/., 2000). six or ed in Maryanska et al.'s analysis due to the large seven in Oviraptor (Barsbold et al., 1990). and amount of missing data they thought (Maryanska eight in Hqwannia huangi (Lu. 2003). et al., 2002). taxa should not be excluded a priori Nernegtia barsboldi independently acquired from phylogenetic analysis based only on the the avian characters such as the fused premaxil- number of preserved characters (Kearney and lae, as in Confzicizisornis sanctus (Chiappe et al., Clark, 2003). Therefore, in the present analysis, 1999). toothless jaws, and the presence of nutri- these taxa are still employed. Most characters are ent openings on the premaxilla and the maxilla as from Maryanska et al. (2002). New characters in other oviraptorids. and their sources in appcndix 2 arc in bold. Sharp ventrolateral margins of the premaxillae Phylogenetic analysis was performed using indicate that a keratinous structure probably cov- MacClade 3.08 (Maddison and Maddison. 1992) ered the end of the rostrum, as in Er1icosaurv.s and PAUP 4.0b (Swofford 1998). Because of the undrewsi (Clark et al., 1994), ornithomimids large data set (20 taxa) and many missing charac- (Norell, Makovicky, and Currie, 200 1 ; Kobayashi ter states, a Heuristic Search was used (Swofford and Lu, 2003), and birds. The rod-like jugal is and Begle, 1993), with branch-swapping options similar to that of sunctus and of the TBR swapping algorithm method. The other birds (Elzanowski, 1999). the mobile con- analysis resulted in one most parsimonious tree dition between the quadrate and the quadratoju- (tree length=48l; consistency index=0.5073; re- gal is similar to the condition of most birds, and tention index=0.6269) (Fig. 8). This tree shows so are the increased cervical and sacral vertebral that Oviraptorosauria forms a monophyletic counts. group. Nineteen unambiguous synapomorphies support this . These characters are 1(1), Phylogenetic Analysis 9(1), 23(1), 37(1), 40(1), 41(2), 431)- 46(1), 47(1), 55(1), 58(1), @(I), 71(1), 76(1), 77(1), In order to determine the phylogenetic status 81(1), 84(1), 86(1) and 88(1). The primitive of Nemegtia barsboldi among oviraptorosaurs, forms, such as gauthieri, 20 taxa and 200 characters (107 cranial and 93 zoui, Microvenaior celer-, and Avim- postcranial characters) (Appendices 2 and 3) are imus portentosus form an ascending sequence used for this analysis. All characters are equally from most basal to more derived in the tree. weighted and unrooted. The character/matrix are portentosus is a basal form of Caenag- modified from Maryanska et (11. (2002). Five new nathoidea. Following Barsbold ( I981 ; 1983), genera of Oviraptorosauria are added to the ma- Maryanska et al. (2002) used GIN 100142 (IGM trix; these include mckennai (Clark et ul., 100142) as the representative of Ovir-aptor philo- 2001), Citipati osmolskae (Clark et al., 2001, ceratops in their phylogenetic analysis. The pre- 2002), Incisivosaur-us gauthieri (Xu, Cheng et sent analysis shows that GIN 100142 and Ovirap- al., 2002), Nemegtia barsboldi, and Heyannia tor philoceratops are closely related. GIN 100142 huangi (Lii, 2003). The aim of this analysis is to either belongs to a different species of the same determine the phylogenetic position of Nemegtia as 0. philoceratops or the same species, bur*sholdi among oviraptorosaurs, so the most the latter differs from Clark et ul.3 judgment primitive forms such as Hc.rwrusuirr~irs is- (Clark et al., 2002). Nemegtia harsholdi is close- chigualastensis, Coeloph~~sisbauri, and the spe- ly related to Citipati osmolskae. cialized forms such as and Al- varezsauridae, which were used by Maryanska et Conclusion al. (2002) as the outgroup in their analysis, are excluded from the present analysis. Although Mi- Nemegtia barsholdi is distinguished by at least New Oviraptorid Dinosaur. Mongolia 113

leotrrologic~heskoiEkpd tsii, 15: 28-39. five autapomorphies from other known ovirap- Barsbold R.. 1983. Carnivorous dinosaurs from the Cre- torosaurs (vertical skull crest, anteroposterior taceous of Mongolia. Trudy Sovmestnoi So~wtsko-Mon- length of the frontal approximately 25% of that golSkoi Pa/eonrologicheskoi Ekspeditsii. 19: 5- 120. of the parietals in dorsal view, less exposed nasal Barsbold, R.. 1986. [Raubdinosauricr Oviraptoren]. In process of the premaxilla on the dorsal surface of Vorobyeva. E. I. (ed). Herperologisc~heUntws~ich~ingcn in dcr Mongolischm Volksrsrrpublik.Pp. 2 10-223. Akad. the skull, a process on the quadrate projecting Nauk. S. S. S. R. Inst. Evolyucionnoy Morfologii i into the cotyla of the quadatojugal, mandibular Ekologii Zhivotnykh im. A. M. Severtsova, Moskva (in condyles of the quadrate situated rostrally to the Russian with German summary). occipital condyle). Phylogenetic analysis shows Barsbold R.. 1997. Oviraptorosauria. In Currie. P. J. and that Nemrgtia har~sholdiis closer to Citipati os- Padian. K. (eds.). Enc.vclopedia of' Dinosaiirs. Pp. 505-509. Academic Press. San Diego. molskue than to other oviraptorosaurs. Barsbold, R.. Currie, f? J.. Myhrvold, N. P., Osmolska. H.. Tsogtbaatar. K. & Watabe, M., 2000. A pygostyle from Acknowledgments a non-avian theropod. Nature, 403: 155- 156. Barsbold. R.. Maryanska, T. & Osmolska, H.. 1990. Ovi- The authors would like to thank Drs. Louis L. raptorosauria. In Weishampel. D. B., Dodson. P. and Jacobs, Dale A. Winkler (SMU, USA), H. Osmolska. H. (eds.). The Dinosauria. Pp. 249-258. Osmolska (Poland), R. E. Molnar (USA), R. University of California Press, Berkeley. Barsbold, R. & Osmolska. H., 1999. The skull of Vsrrloci- Barsbold (Mongolia), and Y. Kobayashi (Japan) raptor (Theropoda) from the Late Cretaceous of Mon- for their valuable comments on the early versions golia. Acta Palaeontologica Polonica, 44(2): 189-2 19. of the manuscript, and to acknowledge the mem- Barsbold R.. Osmolska. H.. Watabe. M.. Currie. F! J. & bers of the Mongol Highland International Di- Tsogtbaatar, K., 2000. A new oviraptorosaur (dinosaur, nosaur Project. Special thanks go to Y. theropod) from Mongolia: the first dinosaur with a py- gostyle. Acla Palaeontologica Polonica. 45(2): 97- 106. Kobayashi, who found this beautiful specimen in Baumel, J. J. & Witmer, M., 1993. Osteologia. In Baumel. 1996. Thanks also go to Drs. Zhou, Z. and Xu, J. J., King, A. S. et a/. (eds.). Handbook of'avian anaro- X. (IVPP of Chinese Academy of Sciences, Bei- mr: nomina anatomica a~~iurn.second edition. Pp. jing, China), for access of the specimens in their 45- 132. Cambridge, Massachusetts. care. This project is supported by Chunichi Shin- Bonaparte. J. F.. Novas, F. E. & Coria, R. A., 1990. bun Co. Ltd., Kyoto Kagaku Co. Ltd., Chukyo Carnotaunrs sastvei Bonaparte, the horned, lightly built carnosaur from the Middle Cretaceous of Patagonia. TV Broadcasting Co. Ltd., and Tokai Bank Ltd.; Contributions in Science Nut. Histons Museum Los An- plus Institute for the Study of Earth and Man at geles County. 416: 142. Southern Methodist University. the Chiappe. L. M., 1996. Late Cretaceous birds of southern Foundation, and the Chang Ying-Chien Science South America: Anatomy and systematics of enantior- Grant for USA-China Collaborative Field Re- nithes and Patagoptetyx dgftrrariisi. Mwzchnrr Ge- owiss. Abh. (A),30: 203-224. search to Junchang Lii. Chiappe. L. M., 2001. Phylogenetic relationships among basal birds. In Gauthier. J. and Gall. L. F. (eds.). New References persppccrivc.~on the origin and earl^, evol~rrionof'birds. Proceedings of'thr International Svrnposi~imin Honor Barsbold R.. 1976. [On a new Late Cretaceous family of of'John H. Ostrom. Pp. 125- 139. Yale Peabody Muse- small theropods (Oviraptoridae fam. n.) of Mongolia]. um. Doklad~.Akademia Nauk SSSR. 226(3): 685-688 (in Chiappe. L. M., 2002. Basal phylogeny: Problems Russian). and solutions. In Chiappe L. M. and Witmer L. M. Barsbold R.. 1977. Kinetism and peculiarity of the jaw (eds.). hirh above the heads of'dinosaurs. apparatus of oviraptors (Theropoda. ). Tmns- Pp. 448472. University of California Press. Berkeley. actions. Joint Sovier-Mongolia Gcolog~calExpedition. Chiappe, L. M.. Ji, S-A,, Ji, Q. & Norell, M. A..1999. 4: 37-47 (in Russian). Anatomy and systematics of the Confuciusornithidac Barsbold R., 198 1. Toothless carnivorous dinosaurs of (Theropoda: Aves) From the late Mesozoic of north- Mongolia. Tr~tdj.Sovmesrnoi So~~c~.vko-i~iongoIkkoiPa- eastern China. Bulletin of' the American Mziseum of 114 Lii. Tomida. Azuma. Dong and Lee

.Vtrr~rrtrlHistor:\.. 242: 1 -89. from the Judith River (Oldman) Formation of Alberta. Chiappc. L. M.. Norell. M. A. & Clark. J. M.. 1996. Phy- Canada. Curintiion Jolrrnnl of'Eurrh Scierzcw, 25: 972 logenetic position of Mononlkxv (Avcs: Alvarezsauri- 986. dac) from the Late Cretaceous of the Gobi Desert. Currie. I? J.. Vickers-Rich. F? & Rich. T. H., 1996. Possi- .Mcwoir:v of'tlic~Q~rcwr.rltrncl Mirscwrn. 39( 3 ): 557 582. ble oviraptorosaur (Theropoda. Dinosauria) spccimcns Chiappc. L. P.. Norell, M. A. & Clark. J. M.. 1998. The from the Otway Group of Dinosaur skull ol'a relative of the stem-group bird Mononyklrs. Cove. Australia. .4lclieringn. 20: 73-79. Nuiirrc~.392: 275-278. Currie, P. J. and Zhao. X.-J.. 1993a: A new carnosaur (Di- Clark. .I. M.. Altangercl. P. & Norcll. M. A,. 1994. The nosauria: Theropoda) from the Jurassic of Xiniiang, skull of Erlic~ostrrrr?~.~crr~tlrewvi. a Late Cretaceous seg- People's Republic of China. Cunudiu~lJo111.nu1 ~f'Enrth nosaur (Theropoda: Therizinosauridea) from Mongolia. ScVencw. 30: 2037-208 1. .4rnericrrn Mir.vcwni NoiVttrrc,.~.3 1 15: 1-39. Currie. P. J. & Zhao. X.-J.. 1993b. A new troodontid (Di- Clark. J. M.. Norell, M. A. & Barsbold. R.. 2001. Two nosauria. Theropoda) braincase from the Dinosaur Park new oviraptorids (Thcropoda: Oviraptorosauria). Upper Formation () of Alberta. Cu~~trtiitmJo~rrnal Cretaceous D,iadokhta Formation. Ukhaa Tolgod. Mon- of'EarTh Sc;c~ncw.30: 223 1 -2247. golia. Jo~rr/ltrlN/' Ciw~hrorc~Puleonrolog!~. 21(2): 209~ Dong, Z.-M. & Currie. I? J.. 1996. On the discovery of an 213. oviraptorid skeleton on a nest of at Byan Man- Clark. J. M.. Norcll. M. A. & Chiappe. L.M.. 1999. An dahu. Inner Mongolia. People's Republic of China. oviraptorid skeleton from the late Cretaceous of Ukhaa Cu~~udiur~Jo~rrnuI of'Eurth Sciencev. 33: 63 1-636. Tolgod. Mongolia. prcservcd in an avian-like brooding Elzanowski. A,. 1999. A comparison of the jaw skeleton position over an oviraptorid nest. .4nierictrri M~rscwrtl in theropods and birds. with a description of the palate Nolituiev. 3265: I - 36. in the Oviraptoridae. In Olson, S. L. (cd.).Avian Pale- Clark. J. M.. Norell. M. A. & Rowc. T.. 2002. Cranial ontology at the Close of the 20'" Century: Proceedings anatomy of Citil~uiiosrnolskoc (Theropoda. Ovirap- of the 4"' International Meeting of the Society ot' Avian torosauria). and a reinterpretation of the holotype of Paleontology and Evolution. Washington. D.('.. 4-7 01~ir.upro1.philocc,rutops. .411ic~ictrnM~rsc~~~rn Nol.iitrie.v. June 1996. Sn~ithwninnContrih~rtion.~ to Pu/c~o/~io/og~~. 3364: 1 24. 89: 3 1 1 323. Colbcrt. E. H. & Russell. D. A,. 1969. The small Creta- Frankfurt. N. G. & Chiappc. L. M.. 1999. A possible ovi- ccous dinosaur Dro~~~ueo.vcr~rrlr.~..A~II~I.~~.UII MIISCWII raptorosaur from thc Late Crctaccous of northwcstcrn No~itrrtc~.~.2380: 1 -49. Argentina. Jolrrnul of' krrehruir, Ptrleo~~/olog~~.19( 1 ): Cracraft. J.. 197 1. Caenagnathiformes: Cretaceous birds 101- 105. convergent in jaw mechanism to dicynodont reptiles. Frcy. E. & Martill. D. M.. 1995. A possible ovirap- Jotrrr~trlOf'Pt~/~On/0/0gJ', 45(2): 805- 809. torosaurid theropod from the Santana Formation Cracrati. J. 1986. The origin and early diversification of (Lower Cretaccous. ?Albian) of Brazil, ~VcwevJtrl~~h~rcA birds. Ptr/c~oho/ogl~.12: 383-399. ,fii'r Gt~ologiclrnd Pu/uonto/ogic rW~i~utshc.fic.7: 397 Curric. P. .I.. 1985. Cranial anatomy of Sic~r~o~~~u~l~o.str~~r~~.~4 12. i~rry~rcrlis(Saurischia. Thcropoda) and its bearing on Gauthier. J.. 1986. Saurischian ~nonophylyand the origin the . Ctrnutlitm Joirn~ul of' Earili of birds. 111 Padian K. (cd.). Thr origin of hi1.d~ul~d the Scicwcw. 22: 1643- 1658. r,~d~~iion~f:fligh/. Mc~~noir.~ (?/'the CuIifOrniu .Acodewri~ C'urric, I? J.. 1995, New information on the anatomy and of'Scionc.e\.8. San Franc~sco.Pp. 1 55. relationships of D~.oniuc-oscllrr.lr.s ulher~/c~i.~~s(Di- Gilmorc. C. W.. 1920. Osteology of the carnivorous Di- nosauria: Thcropoda). JolrrwuI of' Ertchcrtc' PUICOII~OI- nosauria in the United States National Museum. O'ql.. 1s: 57659 1. with special reference to the genera Antrodcwi~rs(.I/- Curric. P. .I.. 2000. Theropods from the Cretaccous of loscr~rr~irs)and Ccwtostrirrr~..U~itcvi Siutec. hiutiorttrl Mongolia. 111 Bcnton. M. J.. Shishkin. M. A,. Unwin. D. IWI~S~~L~II.B~rIl~~rir~. 1 10: I - 154. M. and Kurochkin. E. N. (cds.). Tlw trgc of tli~io.str~rr:vin Gradzinski. R.. Kazmicrczak. J, & Lctkld. J.. 1968. Geo- R~a.vitrtr~~cl ~Mr~~~golitr. Pp. 434455. Cambridge Uni- graphical and geological data from the Polish-Mongo- versity Press. Cambridge. lian Palacontological txpeditions. Ptrltrcontologitr Currie. P. .I.. Godfrcy. S. .I. & Nessov. L.. 1993. New cac- Polonictr. 19: 33- 92. nqnathid (Dinosauria: Thcropoda) specimens from thc Holmgren. N.. 1955. Studies on the phylogeny of birds. Upper Cretaceous of North America and Asia. Crr~~trtli- .4cirr Z,~ologic~~.36: 2.13-328. url Jorrrntrl of Eor.t11Scitvtw. 30: 22552272. Holtz. T. R. Jr.. 1994. The arctomctatarsalian pes. an un- Currie. P. .I. & Russell. D. A,. 1988. Ostcology and rela- usual structure of the metatarsus of Cretaceous tionships of Cl~i,n.vtcr~otc>s(Saurischia. Theropoda) Theropoda (Dinosauria: Saurischia). Jolrrnul of l.i,rie- New Oviraptorid Dinosaur. Mongolia

l~ru/cP(~lcon/ologi~. 14(4 ): 480- 5 19. h,rr/u Pc~l.l.sicr/iccr.38(3): 99. Holtz. T. R. Jr.. 2000. A new phylogeny ofthc carnivorous Maddison. W. P & Maddison. D. R.. 1992. iMtrc,(%clc,: dinosaurs. Guiu. 15: 5-6 I. untr/~:sisof'pl~~~logcv~~ trnd chr.trctc,r c~idtrtion,version Holtz. T. R. Jr.. 2001. Arctometatarsalia revised: The 3. Sinaucr Associates. Inc. Publishers. Sunderland. problem of homoplasy in reconstructing theropod phy- Massachusetts. logeny. In Gauthier. J. and Gall. L. F. (eds.). N~M.p,r- Madscn. J. H. Jr.. 1976. ..ll/o.sn~r~~tr.r,fi~ugiIi.~:a revised oste- .spcc.ti~v.son the origin and Eur!~.c~dtrtion of' hilrls: ology. Urtrh Geologic~rluntl hlinerul Strrlyi'. B~dlctin. Pmcrcding.~of the In~rrncrtionulS~wposilrn~ in Honor 109: 1 163. of John H. 0st1- on^. Pp. 99- 122. Peabody Museum of Makovicky. P. J. & Sucs. H.-D.. 1998. Anatomy and phy- Natural History. Yale University. New Haven. logenetic relationships of the theropod dinosaur M- Hwang. S.-H.. Norell. M. A,. Ji. Q. & Gao. K.-Q.. 2002. c~ro~~c~t~utorcder tjo~nthe Lower Crctaccous of Mon- New specimens of' Microruptor rlinoiunlrs (Theropoda: tana. An~eric,trnM~tseLrn~ Novitutc,.~. 3240: 1 27. Dromacosauridae) from Northeastern China. An~nicun Marsh. 0. C.. I88 1. Classification of Ihc Dinosouria. The Mlr.scwn ,Vo~irures.338 1: 1-44 .4nlel.ic.trnJotrrnul o/Scio~cc.Thid swics. 23: 8 I -86. Hwang. S.-H., Norell. M. A,. Ji. Q. & Gao. K.-Q., 2004. Maryanska. T. & Osm6lska. H.. 1997. The quadrate of' A large compognathid from the Early Cretaceous Yixi- oviraptorid dinosaurs. .-Icttr Prrltri~ontologicuPolonictr. an Formation of China. Jotrr11~1of'S~:st~rntrtic Pult~(w- 42(3): 361371. rolog?,.2( 1 ): 13- 30. Maryanska. T., Osmdska. H. & Wolsan. M.. 2002. Jerzykiewicz. T. & Russell. D. A,. 199 1. Late Mesozoic Avialan status for Oviraptorosauria. .4cttr P~rlrrc~ontolog- stratigraphy and of the Gobi Basin. Cretu- iccl Polonit~r.47( 1 ): 97- 1 16. cw)tr.s Restwrch. 12: 345- 377. Molnar. R. E.. 1973. The cranial morphology and mc- Ji, Q.. Curric. P. J.. Norell. M. A. & Ji. S-A,. 1998. Two chanics ol' ~w~~nostrrrrtrsre.r (Rcptilia: Saurischia). feathered dinosaurs from northeastern China. Ntrtlrrc. PhD. Diss.. University of California at Los Angclcs. 393: 753-76 1. Molnar. R. E.. 199 1. The cranial morphology of ~~'IYIII- Ji. Q.. Ji. S.-A,. You. H.-L.. Zhang. J.-P. Yuan. C.-X.. Ji. nosunr~rsre.r. P~rlurontogrupI~icu.2 17(4-6): 137 - 176. X.-X.. Li. .I.-L. & Li. Y.-X.. 2002. Discovery of an Norell. M. A,. Clark. J. M.. Chiappe. L. M. & Dashzevcg. bird Shenzholr,rrp/or.sincw.sis gen. et sp. nov. - D.. 1995. A nesting dinosaur. X~IIIII-C,.378: 774-776. from China, G~~ologictrlB~rllr/in of China 21 (7): 363 Norell. M. A.. Clark. .I. M., Dashzcvcg. D., Barsbold R., 369. Chiappe. L. M.. Davidson. A. K., Mckcnna M. C.. Alt- Ji. Q.. Ji. S.-A,. You, H.-L., Zhang. J.-P.. Zhang, H.-B.. tancrel. I? & Novacck. M. J.. 1994. A thcropod dinosaur Zhang, N.-J.. Yuan. C.-X. & Ji. X.-X.. 2003. An Early embryo ;~ndthe affinities of the Flaming ClitTs Di- Cretaccous avialan bird. Shenrho~rruprorsin(w.sis from nosaur eggs. Sr~cwc*.266: 779-782. western Liaoning. China. ,4ctu Geologicu Sinica. 77( 1 ): Norell. M. A,. Clark. J. M. & Makovicky. I? J.. 2001. 2 1 -26. Phylogenetic relationships among coelurosaurian Kearney. M. & Clark, .I. M.. 2003. Problems due to miss- theropods. 111 Gauthier. J. and Gall. L. F. (4s.). .M,u ing data in phylogenetic analyses including : a per.specti~won the origin crnd Eu,!I e~ulrrtionof'hirds: critical revlew. Jolrrnul o/' I+rtchtr/e Puloontologj~. P~~ocwtlingvof' the lntwnurionul St.np).siun~in Honor 232): 263-274. of'John H. Ostroni. Pp. 49-67, Peabody Mus. Nat. Hist. Kobayashi. Y. & LC. J.-C., 2003. A new ornithomimid di- Yale University. New Haven. nosaur with gregarious habits from the Late Cretaceous Norell. M. A,. Makovicky. P. .I. & Currie. P. J.. 2001. The of China. ,4ctu Pularontologictr Polonica, 48(2): 235- beaks of ostrich dinosaurs. Nutlrw. 412: 873-874. 259. Novas, F. E.. 1993. New information on the systematics Lii. J.-C.. 2003. A new oviraptorosaurid (Theropoda: Ovi- and postcranial skeleton of' Horlrrzr.sutr~~tr~ischig11~11u~1- raptorosauria) from the Late Cretaceous of southern cwsis (Theropoda: Hcrrerasauridae) from the Is- China. Jolrrnul of knchrarc, Pul~ontologl~.22( 4): chigualasto Formation (Upper ) of Argentina. 871 875. Jo~r~mluf'I~i~~~eh,zr/e Puleon/ologi~. 13: 400423. Lii. J.-C.. Dong. Z.-M.. Azuma.Y.. Barsbold. R. & Tomi- Osborn. H. F.. 1924. Three new theropoda, protoceratops da. Y., 2002. O~iruptomsaurscompared to birds. In zone. central Mongolia. .4n1cricun Mtr.scwi~No1i1ure.s. Zhou, Z.-H.. and Zhang, F.-C. (eds.). Proceedings of' 144(7): 1 12. the 5th Symposium of the Society of Avian Paleontol- Osmolska, H.. 1976. New light on the skull anatomy and ogy and Evolution. Science Press. Beijing China. Pp. systematic position of O~irzrp/or.Nurtr~~c~. 262: 683- 175- 189. 684. Lii. J.-C.. Huang. D. & Zhang, S.. 2000. The first discov- Osmolska. H.. Ronicwicz. E. & Barsbold. R.. 1972. A cry of oviraptorosaur from Guangdong Province. Vertc,- new dinosaur Gtrllin~im~rsh1rl1ut11.s. n. gcn. n. sp. (Or- I lh Lii, Tomida, Azuma. Dong and Lee

nithomimidae) from the Upper Cretaceous of Mongo- ty of archosaurs: a study in soft-tissue rcconstruction in lia. Pala~ontol.Polonica. 27: 103- 143. the fossil record with an analysis of the function of Ostrom, J. H.. 1978. The osteology of Compsognathus pneumaticity. Society of Vertebrate Paleontology mem- longiprs Wagner. Zittdiana, 4: 73- 1 18. oir 3. Journal o/' krtrbratt~pale onto log^^. 17: supple- Rauhut, 0. W. M., 2003. The interrelationships and evolu- ment to number I. tion of basal theropod dinosaurs. Special Papem in Xu. X.. Cheng, Y.-N., Wang, X.-L. & Chang, C.-X., 2002. Palaeontology, 69: 1 -2 1 3. An unusual oviraptorosaurian dinosaur from China. Russell, D. A. & Dong, Z.-M., 1993. The affinities of a Nature, 419: 291 -293. new theropod from the Alxa Desert. Inner Mongolia, Xu. X.. Norell. M. A,. Wang. X.-L.. Makovicky. P. J. & Pcoplc's Republic of China. Canadian Jo~irnalof'Earth Wu, X.-C.. 2002. A basal troodontid from the Early Sciorccs. 30: 2107 -2 127. Cretaceous of China. Nulure. 415: 780-784. Sanz. J. L. & Bonapartc. J. F. 1992. Zhou, 2.-H., & Wang, X.-L.. 2000. A new species of romemli. a fossil small bird articulated skeleton from Cairdipta;vx from the of Liaoning, the Early Cretaceous of Spain. Proceedings of the 11 In- Northwest China. I+rtebrata PalA.siatica. 38(2): 1 1 1- ernational Symposium of Avialian Paleontology, 1988, 127. Pp. 39-49. Zhou. Z.-H.. Wang, X.-L., Zhang. F.-C., & Xu. X., 2000. Sereno, I? C., 1999. The evolution of dinosaurs. Scirncr, Important features of Caudiptery-evidence from 284: 2137-2147. two nearly complete new specimens. krtc,bra/a PaL4si- Sereno, P. C.. 2000. lberonmornis romemli (Aves. Or- atica. 38(4): 24 1 - 254. nithothoraces) reevaluated as an Early Cretaceous enan- Zhou. Z.-H. & Zhang, F.-C., 2002. A long-tailed seed- tiornithine. Neurs Jahrbuch ,fur Grologie und eating bird from the Early Cretaceous of China. Nahrrp. Polaontologi~~Ahkandlimgen, 2 IS(3):365 - 395. 418: 405409. Screno. P. C. & Novas. F. E., 1993. The skull and neck of the basal theropod Ho-rerasuirrus ischiguaIastm.si.\. Appendix 1 Journal of Vertrhratr Palrontologl:. 13(4):45 1476. Abbreviations: An: angular; ar: articular; Sereno, P C. & Rao. C.-G.. 1992. Early evolution or avian Hight and perching: new evidence from the Lower Cre- asc: articular-surangular-coronoid complex; bs: taceous of China. [email protected]: 845-848. basisphenoid: cav antorb: antorbital cavity; ch: Smith. D.. 1992. The type specimen of Oviraptor philo- choana; CH: chamber; d: dentary; ec: ectoptery- c~ernrops.a theropod dinosaur from the Upper Creta- goid; emf: external mandibular fenestra: eo: ex- ceous of Mongolia. Nrires Jahrbuch fur Gcologir und occipital; f: frontal; fmb: ,fi.ssa m. hrachialis; j: Paluontologie Abhandl~mgen.186(3): 365 - 388. Sternberg. R. M., 1940. A toothless bird from the Creta- jugal: I: lacrimal; lat: laterosphenoid; m: maxilla; ceous of Alberta. Journal o/'P~~leonrology.14( 1 ): 81- mf maxillary fenestra; na: nasal; nar: narial 85. opening; oc: occipital condyle; o, orbit; or: or- Sues. H.-D.. 1997. On Chirostenotrs. a Late Cretaceous bitosphenoid; SP: supporting pillar; p: parietal: oviraptorosaur (Dinosauria: Theropoda) from western pa: palatine; par: parasphenoid; pm: premaxilla; North America. Joiirnul~/'Kwrbrute pale onto log^.. 17: po: postorbital pre: prearticular; pt: pterygoid; q: 698-7 16. SwoRord, D. L.. 1 998. PAC'P*: phvlogenrtic urrtil~:sis quadrate; qj: quadratojugal; sof: suborbital fenes- using parsimonj. (and orher rnethod.~).Version 4.0. Sin- tra; sq: squamosal; stf: supratemporal fenestra; t: auer. Sunderland, Mass. tooth-like process formed by maxillae and Swofford. D. L. & Begle. D. P.. 1993. PAW: phvlogc~nctic~ vomers: v: vomer. una11:c.i~using parsimony. 1.iwion 3. 1. L'ser $ manual. Illinois Natural History Survey. Champaign. Illinois. Appendix 2. Welles. S. P., 1 984. Dilopho.suirru.s ~~etherilli( Dinosauria. Theropoda). Osteology and comparisons. Palurontolo- Characters for phylogenetic analysis of the re- gruphicu A, 185: 85- 1x0. lationships among Oviraptorosauria (modified Wester. D.. 1996. Dinosaurs of the Gobi. National Geo- from Maryanska et a[., 2002 with additional taxa graphic. 190( 1): 70-89. and recoded and additional characters as noted in Witmer. L. M.. 1990. The craniofacial air sac system of bold) and data matrix: O=plesiomorphic charac- Mesozoic birds (Aves). Zoological Journal of'thc, Lin- netrrr Socic*/~.(?/'London. 100: 327-378. ter state; 1.2.3 =derived character states; Witmer. L. M.. 1997. The evolution of the antorbital cavi- ? =missing data: p=character not applicable. New Oviraptorid Dinosaur. Mongolia 117

Preorbital skull length to basal skull illa at least twice the length of the pre- length ratio: 0.6 or more (0); 0.5 or less maxilla (0); shortened, nasals subequal (1). in length to frontals or shorter, maxil- Longitudinal pneumatized crest-like lary length less than twice the length of prominence on the skull roof absent (0), the premaxillary body (I) (Rauhut, present ( 1 ). 2003). In the majority of theropods, the Premaxilla main body length (ventral) to preorbital part of the skull forms an elon- height (below the naris) ratio: 1 .O-1.4(0); gate snout. with maxilla and nasal being more than 1.7(1); 0.7 or less (2). two of the longest bones of the skull roof. Otosphenoidal crest vertical on ba- In the basal oviraptorosaur Incisivosuurtrs sisphenoid and prootic, and does not gauthier-i and in Nemrgtia barsholdi, the border an enlarged pneumatic recess nasal is longer than the frontal (Xu et al., (0) or well developed, crescent shaped, 2002). In Cuudiptery zoui (Ji et ul., thin crest forms anterior edge of en- 1998) and most derived oviraptorosaurs, larged pneumatic recess (1). This struc- the nasal is shorter than the frontal. ture forms the anterior, and most distinct, Maxillary process of premaxilla con- border of the "lateral depression" of the tacts nasal to form posterior border of middle ear region (see Currie, 1985; Cur- nares (0) or maxillary process reduced rie and Zhao, 1993) of troodontids and so that maxilla participates broadly in some extant avians (Hwang et al.. 2004). external nares (1) or maxillary process Subnarial (maxillary) process of the pre- of premaxilla extends posteriorly to sep- maxilla: contacts the nasal, the maxilla ex- arate maxilla from nasal posterior to cluded from the narial border (0); does not nares (2) (Hwang et al., 2004). contact the nasal, the maxilla participates Nasal recesses: absent (0); present (I). in formation of the narial border (1 ). Caudal margin of the naris: rostral to the Premaxillae in adult: unfused (0); fused rostral border of the antorbital fossa (0); (I) (Chiappe, 1996). nearly reaching or overlapping the rostral Pneumatization of the premaxilla: absent part of the antorbital fossa (I); overlap- (0);present ( 1 ). ping most of the antorbital fossa (2). Basisphenoid recess present between Ventral margin of the external naris: at the basisphenoid and basioccipital (0) or level of the maxilla (0); dorsal to the max- entirely within basisphenoid (1) or ab- illa ( l ). sent (2) (Hwang et al., 2004). Prefrontal: present (0); absent or fused Subantorbital portion of the maxilla: not with the lacrimal (I). In the majority of inset medially (0); inset medially (I). theropods, the prefrontal is large. Pre- Palatal shelf of the maxilla with two longi- frontal may be absent in all ovirap- tudinal ridges and a tooth-like process: ab- torosaurs, including the primitive Inci- sent (0); present ( 1 ). sivosaurus guuthieri. Rim around the antorbital fossa: well pro- Lacrimal recess: absent (0); present ( I). nounced (0); poorly delimited ( 1 ). Premaxillary symphysis acute, V- Antorbital fossa: not bordered rostrally by shaped (0); or rounded, U-shaped (I) the premaxilla (0): bordered rostral ly by (Hwang et al., 2004). the premaxilla ( I ). Pronounced, round accessory antorbital Preorbital region of the skull in post- fenestra absent (0); or present (I). A hatchling individuals: elongate, nasals small fenestra. variously termed the acces- considerably longer than frontals, max- sory antorbital fenestra or maxillary fen- Lii, Tomida. Azuma. Dong and Lee estra, penetrates the medial wall of the an- or more of the infratemporal fenestra (2); torbital fossa anterior to the antorbital fen- absent (3). estra in a variety of coelurosaurs and other 34. Dorsal part of the quadrate: erect (0); di- theropods (Hwang et ul., 2004). rected backwards ( 1 ). Parietal length to frontal length ratio: 0.6 35. Otic process of the quadrate: articulating or less (0); 1.0 or more (I ). only with the squamosal (0); articulating Narial region apneumatic or poorly with the squamosal and the lateral .wall of pneumatized (0); or with extensive the braincase ( I ). pneumatic fossae, especially along pos- 36. Pneumatization of the quadrate: absent terodorsal rim of fossa (1) (Hwang et (0); present (1). In many theropods, the a/., 2004). quadrate is a solid bone and it is not in- Sagittal crest on the parietals: absent (0); vaded by a diverticulum of the tympanic present (I). pneumatic system, but the quadrate is Jugal pneumatic recess in posteroven- pneumatized by the quadrate diverticulum tral corner of antorbital fossa present in most modern birds. The diverticulum (0); or absent (I) (Hwang et a/., 2004). enters the bone ventromedially, dorsome- lnfratemporal fenestra: ventrally nearly as dially or caudally (Witmer, 1990). In dro- long as high rostrally (Oj; shorter ventrally maeosaurids ( klocivaptor rnongoliensis) than (I); large, square (2); not separate and other non-avian theropods. the from the orbit (3). quadrate lacks pneumatic foramina (Col- Descending (prequadratic) process of the bert and Russell, 1969; Madsen. 1976; squamosal: constricting the dorsal part of Bonaparte et al., 1990; Barsbold and the infratemporal fenestra (0); not con- Osmolska, 1999). Among non-avian stricting the infratemporal fencstra ( 1 ). theropods, troodontids (Currie and Zhao, Jugal and quadratojugal separate (0); 1993b) and tyrannosaurids (Molnar, 1 99 1 ) or quadratojugal and jugal fused and are the exception, in which the quadrate is not distinguishable from one another pneumatic. A large pneumatic foramen is (1) (Hwang et a/., 2004). placed anteromedially and somewhat Suborbital part of the jugal: deep below the mid-height in some oviraptorids dorsoventrally and flattened lateromedially (Maryanska and Osm6lska, 1997). Some (0); shallow dorsoventrally or rod-shaped birds, such as Archuroptervx lithographi- (1 ). ca, Hespe~*ornisregalis and Puruhesperor- Jugal-postorbital contact: present (0); ab- nis alexi (Chiappe, 1996) also lack pneu- sent ( 1). matic foramina. The quadrate diverticu- Quadratojugal process of the jugal in lat- lum perforates the lateral surface of the eral view: forked (0); tapering (I); fused quadrate in Patagoptriyx defkvruriisi (Chi- with the quadratojugal (2). appe, 1996), and a similar condition is Quadratojugal-squamosal contact: tips present in Hqumnniu huungi (Lii, 2003). of the bones widely separated (0); the but it is absent in "Ovivap/or" rnongolien- contact present (I) (modified). sis. The lateral position of the entrance of Ascending (squamosal) process of the the quadrate diverticulum is regarded as quadratojugal: massive, bordering about an autapomorphy of Patugop/~'~:~:r1kf21.- the ventral half of the infratemporal fenes- rariisi (Chiappe, 1996). but the similar po- tra (0); slender, bordering the ventral half sition in Hevuanniu indicates an indepen- or less of the infratemporal fenestra (I); dent acquisition. Witmer ( 1990) consid- slender, bordering the ventral two-thirds ered the presence of quadrate pneumatici- New Oviraptorid Dinosaur. Mongolia I 19

ty as a synapomorphy of the Carinatae Zhao, 1993a). the mandibular condyle of [Ichthyornithi tomes+ Aves], but Chiappe the quadrate is caudal to the occipital ( 1996) thought that a pneumatic quadrate condyle. In Archuropte~y.~lithogruphicu, was present at least in the common ance- lncisivosaurus guuthieri and Cuudipter:w stor of Putugoptery di?f~rrurii.siand the sp. (IVPP V 12430), the mandibular and it might have evolved as condyle of the quadrate is rostral to the early as at the origin of the ornithotho- occipital condyle. Although there are no races. The pneumatic quadrate in ovirap- completely three-dimensionally preserved torosaurs is considered here as indepen- skulls of Cuudipte~vx,the preserved skull dently acquired. is little disturbed in Cuudipteiyx sp. (IVPP 37. Accessory process for a contact with the V 12430). It shows clearly relative posi- quadratqjugal on the distal end of the tion between the mandibular condyles of quadrate: absent (0); present ( 1 ). the quadrate and the occipital condyle. In 38. Quadratojugal sutured to the quadrate Nemegtia hu~,sholdiand "Oviruptor" mon- (0); or joined through a ligamentary ar- go1ien.si.s and Hqwunnia huangi, the ticulation (1) (Chiappe, 2001, 2002). In mandibular condyles of the quadrate are Archueopte~xand most non-avian thero- in the same vertical plane as the occipital pod dinosaurs, the quadratojugal is su- condyle. tured to the quadrate, in almost all de- 40. Enlarged foramen or foramina opening rived oviraptorosaurs, a cotyle is present laterally at the angle of the lacrimal, ab- either on the quadrate ("Ovirapror" mon- sent (0); or present (1) (Hwang et a/., goliensis, GIN 100142. and Conchorapror 2002). gradis, Maryanska et al.. 2002) or on the 4 1 . Paroccipital process directed: laterad (0); medial surface of the quadratojugal GIN lateroventrad ( 1 ), ventrad (2). In both Her- 100121 12 (Lii et ul., 2002). Heyuunniu rerusaurus ischigualus/ensis and AI-

huangi has a groove-like structure on the lo.suunu , fkgilis, the paroccipital process quadrate (Lii, 2003). The presence of a is directed lateroventrally. but it is directed cotyle or a groove-like surface on the much more strongly so in A. fi-agilis than quadrate or quadratojugal is interpreted in H. i.schigzrulasrensis. The paroccipital here as a ligamentary articulation between process directed ventrad in Incisivosuur~rs the quadrate and quadratojugal. In Ciripati gauthieri, Chirostmotes pergrucilis. and osmolskar, the quadrate is tightly sutured derived oviraptorosaurs. to the quadratqjugal (Clark et al., 2002). Dorsal surface of parietals flat, lateral A shallow depression is present on the lat- ridge borders supratemporal fenestra eral condyle of the quadrate in Confuciu- (0); or parietals dorsally convex with sornis sanctus (Chiappe et al., 1999), very low sagittal crest along midline (I); therefore it is interpreted as having a liga- or dorsally convex with a well-devel- mentary articulation between the quadrate oped sagittal crest (2) (Hwang et a/., and quadrato.jugal. 2004). 39. Mandibular condyles of the quadrate situ- Foramen magnum: smaller than or equal ated: caudal to the occipital condyle (0); to the occipital condyle (0); larger than the in the same vertical plane as the occipital occipital condyle ( I). condyle (I); rostral to the occipital Basal tubera: modestly pronounced (0); condyle (2). In Herrerasuurus is- well pronounced, widely separated ( 1 ). chiguulastensis (Sereno and Novas, 1993), Pneumatization of the basisphenoid: weak A1losuurw.s and sinraptorids (Currie and or absent (0): extensive ( 1 ). Lii. Tomida. Azuma. Dong and Lee

Basipterygoid process: well developed (2004), in which IGM 1 OOM2 was coded as (0); strongly reduced (1 ); absent (2). having a tetraradiate palatine, may be Posterior end of dentary without pos- wrong. terodorsal process dorsal to mandibular Pterygoid wing of the palatine situated: fenestra (0); or with dorsal process dorsal to the pterygoid (0); ventral to the above anterior end of mandibular fenes- pterygoid ( I ). tra (1); or with elongate dorsal process Maxillary process of the palatine: shorter extending over most of fenestra (2) than vomeral process (0); longer than the (Hwang etal., 2004). vomeral process ( I). Parasphenoid rostrum: horizontal or di- Vomer: distant from the parasphenoid ros- rected rostrodorsad (0); slanting rostro- trum (0); approaching or in contact with ventrad ( 1 ). the rostrum ( I ). Tibiofibular crest in the lateral condyle Suborbital (ectopterygoid-palatine) fenes- of femur: absent (0); present (1) (Ji et tra: well-developed (0); closed or reduced a/., 1998). (1). Parietals separate (0) or fused (1) Pterygopalatine fenestra: absent (0); pre- (Hwang etal., 2004). sent ( 1 ). Foramen magnum subcircular, slightly Jaw joint: distant from the skull midline wider than tall (0); or oval, taller than (0); close to the skull midline (I ). wide (1) (Hwang et a/., 2004). Occipital condyle without constricted Medially extended pterygoids meeting neck (0); or subspherical with constrict- each other along the midline and ventrally ed neck (1) (Hwang et al., 2004). underlying the basisphenoid and parasphe- Mandibular symphysis: loose (0); tightly noid: absent (0); present ( I ). sutured ( I ); fused (2). Quadrate wing of the pterygoid: distinct Extended symphyseal shelf at the from the braincase wall (0); overlapping mandibular symphysis: absent (0); present the braincase ( I). (1). Pterygoid basal process for contact with Downturned symphyseal portion of the the basisphenoid: absent (0);present (I ). dentary: absent (0); present ( I). Ectopterygoid situated: lateral to the U-shaped mandibular symphysis: absent pterygoid (0); rostra1 to the pterygoid ( 1 ). (0); present ( I). Ectopterygoid contacts with the maxilla Retroarticular process' length to total and lacrimal: absent (0); present ( I ). mandibular length ratio: less than 0.05 or Hook-like jugal process on the ectoptery- the process absent (0);about 0.10 ( 1 ). goid: present (0); absent ( I ). Mandible maximum height to length ratio: Massive pterygoid-ectopterygoid longitu- about 0.2(0);about 0.1 ( I); 0.3-0.4(2). dinal bar: absent (0); present ( 1). External mandibular Fenestra's height to Palate cxtending below the cheek margin: length ratio: 0.2-0.5(0); 0.7-1.0 (I); the absent (0); present ( 1 ). fenestra absent (2). In Cuenugnuthws Palatine: tetraradiate or trapezoid (0); tri- collinsi and Incisivosaurus guuthieri, the radiate, without a jugal process ( 1); devel- ratio is less than 0.25. but in the derived oped in three plane perpendicular to each oviraptorosaurs such as Ingeniu yunshini, other (2). Triradiate palatine is present In- Hqvuanniu huangi, Ovirupror philocer- cisi~vsaurusguuthieri (Xu, Cheng et ul., atops. "Ovmptoi' mongoliensrs, Con- 2002). Citiputi osmolskue (Clark et al., chovaplor grwcilis. Citiputi osmolskar, 2002) and GIN 100\21 12. So Hwang et 01. this ratio is greater than 0.25. New Oviraptorid Dinosaur. Mongolia 121

External mandibular fenestra's length External mandibular fenestra oval (0); to total mandibular length ratio: or subdivided by a spinous rostral 0.15-0.2(0), 0.1- or less (1); 0.21 or process of the surangular (I) (Hwang et more (2); the fenestra absent (3) (modi- al., 2002). The external mandibular fenes- fied). tra is elongate, and no spinous rostral Coossification of the articular with the process of the surangular is present in In- surangular: absent (0); present (I ). cisivosaurus guuthieri (Xu, Cheng et a/.. Mandibular rami in dorsal view: straight 2002) and Caudipteryx sp. (IVPP V (0);bowed laterad at the mid-length (I ). 12430. pers. observation). In derived ovi- Rostrodorsal margin of the dentary: raptorosaurs, the external mandibular fen- straight or weakly convex (0); deeply estra is subdivided by a spinous rostral concave (I) (modified). In most theropods process of the surangular. and Oviraptor philoceratops, the ros- Mandibular articular facet for the trodorsal margin of the dentary is straight. quadrate: formed of the surangular and ar- In the basal oviraptorosaur lncisivosaurus ticular (0); formed exclusively of the artic- gauthieri, it is slightly convex. It is deeply ular (1 ). concave in Oviraptor philoceratops, Cae- Mandibular articular facet for the nagnuthus collinsi, "Oviruptor" mon- quadrate: with one or two cotylae (0); con- goliensis, Hquznniu huungi, Citipati os- vex in lateral view, transversely wide ( I ). molskue, Conchoruptor gracilis, Nemegtia Articular facet for the mandibular joint harsholdi, and Ingenia ~xznshini. positioned: below the dorsal margin of the Caudal margin of the dentary: incised, caudal part of the mandibular ramus (0); producing two caudal processes (0); above this margin (1). oblique (1 ). The posterior end of the den- Rostral part of the prearticular: deep, ap- tary is strongly forked to form the anterior proaching the dorsal margin of the margin of the external mandibular fenestra mandible (0); shallow, strap-like, not ap- in oviraptorosaurs. In several theropods, proaching the dorsal margin of the such as Allosuurw, dromaeosaurids, or- mandible ( 1 ). nithomimosaurs, therizinosauroids and Splenial: subtriangular, approaching the tyrannosaurs, and Archueopte~a,the cau- dorsal margin of the mandible (0); strap- dal margin of the dentary is oblique. like, shallow, not approaching the dorsal Long and shallow caudodorsal process of margin of the mandible ( I ). the dentary: present (0); absent ( 1 ). Mandibular adductor fossa: rostrally de- Long and shallow caudoventral process of limited, occupying the caudal part of the the dentary, extending caudad at least to mandible (0); large. rostrally and dorsally the caudal border of the extcrnal mandibu- extended not delimited rostrally ( I ). lar fenestra: absent (0); present ( I ). Coronoid bone: well developed (0); re- Pronounced coronoid eminence: absent duced (1) or absent (2) (modified). The (0); present (1) (modified). The coronoid coronoid bone is present in most eminence on the surangular is absent in theropods, but it is weakly developed and most theropods. In lncisivosaur~us guu- strap-like in Incisi~~osuurusguuthirr-i (Xu thieri the coronoid eminence is weakly de- er ul., 2002b). Cuudip/el:\x sp. (IVPP V veloped so it is coded here as (0). The 12430: Pers. observation) and Citipati os- pronounced coronoid eminence is present molskue (Clark et ul., 2002). It is also pre- in Avimimus portentosirs and derived ovi- sent in therizinosauroids (Clark rt ul.. raptorosaurs. 1994). It is absent in birds (Archueo- Lii. Tomida. Azuma. Dong and Lee ptecix), ornithomimosaurs (Osmolska el vsis huuri, Avimimtrs portenfoszrs, al., 1972) and other derived ovirap- Archaeoptei~vx lithogruphica, Ornithomi- torosaurs including Nemegtiu barsboldi, mosauria, , Oviruptor Conchoruptor grucilis, Ingenia and Ovi- philoceratops, Caudiptei?:~sp., Nemegtiu ruptor. barsboldi, Hqvuunnia huungi, Conchorap- Premaxillary teeth: present (0); absent ( 1 ). tor gracilis, Ingeniu junshini. The epiphy- Premaxillary teeth are present in Inci- ses are only present in the third and fourth sivosuurus, Caudipteqx, .4rchaeopreiy cervicals of "Oviiuptor" nzongo1ien.si.s. and most theropods. But premaxillary They are very developed prong-shaped teeth are absent in therizinosauroids and usually extending over the distal end (Clark et ul., 1994). derived ovirap- of the postzygopophyses in Herrerusazrrtrs torosaurs and advanced birds. ischigt~alastensis, A1losuurtr.s , fi.agi1i.s. Maxillary tooth row: extends at least to Dromaeosauridae, Troodontidae and the level of the preorbital bar (0); does not Tyrannosauridae. reach the level of the preorbital bar (I); Cervical ribs: loosely attached to verte- maxillary teeth absent (2). Maxillary teeth brae in adults (0); firmly attached ( 1). are present in lncisivosaurzrs gauthieri, Shafts of the cervical ribs: longer than the and the majority of theropods. Maxillary respective centra (0); not longer than the teeth are absent in Caudiptei?.~ zoui and respective centra ( 1). derived oviraptorosaurs. Pleurocoels or lateral excavations on the Dentary teeth: present (0); absent (I ). dorsal centra: absent (0); present ( I). Dentary teeth are present in Inci- Postzygapophyses on the dorsals: not ex- sivosazrrtrs gauthieri, but absent in de- tending beyond the respective centra (0); rived oviraptorosaurs, celer markedly extending beyond the centra ( I ). (Makovicky and Sues. 1998), Caudiptery Number of the vertebrae included in the zoui (Ji et ul., 1998), Avinzimtrs portento- sacrum in adults: not more than five (0); sus, advanced ornithomimosaurs and more than five (I) (the term sacrum is birds. used here instead of synsacrum in Number of the cervicals (excluding the Maryanska et al.3 original description). cervicodorsal): not more than 10 (0); more Sacral spine in adults: unfused (0); fused 10 (I). (1). Cranial articular facets of the centra in the Continuous sulcus along the ventral side anterior postaxial cervicals: not inclined of the mid-sacral centra: absent (0); pre- or only slightly inclined (0); strongly in- sent ( I ). clined ventrocaudal, almost continuous Pleurocoels on the sacral centra: absent with the ventral surface of the centra (1): (0); present ( I ). ball-shaped (2). Scapula and separate (0), or Anterior cervical centra: not extending fused into scapulacoracoid (1) (Hwang posteriorly beyond the respective neural et al., 2004). The scapula and coracoid are arches (0); extending posteriorly beyond separate in most theropods. including Al-

the respective neural arches ( I). losaurzls , fmgilis, 7)rwmo.saums /a, Epiphyses on the postaxial cervicals: in untirrhopus, form of a low crest or rugosity (0); prong- pe-gmcilis, and Cuudipteiyx sp. They are shaped ( I). The epiphyses on the postaxial fused in monpliensis, Ar- cervicals are in form of a low crest in Mi- chaeoptervx lithographica, Confuciusor- crovniator celer, a rugosity in Coeloph- nis suncttrs. Ingeniu junshini, "Oviruptor" New Oviraptorid Dinosaur, Mongolia 123

mongoliensis, Oviraptor philoceratops, Rao, 1992), but is absent in non-avian Conchoruptor grucilis, and Hejumnia theropod dinosaurs, Archaeoptervx litho- huangi. Because both character states graphic~(Chiappe, 1996) and the basal are present in dromaeosaurids and or- bird Shenzhourxzptor (Zhou and Zhang, nithomimids, this character is coded as Oil 2002; Ji et a/., 2003). The presence of a in them. pygostyle in Nomingiu gohiensis is of 107. Hyposphene-hypantrum articulations great interest in the study of the relation- in trunk vertebrae present (0); or ab- ship between birds and non-avian sent (1) (contrary to the code of Hwang theropods, although it is most parsimo- et al., 2004). These articulations are pre- nious to consider it a convergence sent in He~~~wasaurusischigualastensis (Osmolska, pers. comm., 2003). Some (Novas, 1993). Allosatrrirs .fragilis (Mad- have even regarded the pygostyle in mod- sen, 1976), ,4vimimus portmtostrs. Mi- ern and carinate birds, to be not ho- crownator celer, Chirostenotes pergra- mologous (Holmgren, 1955). The struc- cilis, Conchoraptor grucilis, ornitho- ture of pygostyle In Nomingia gohiensis mimids, troodontids, tyrannosaurids. They (Barsbold Osmolska et al., 2000) and the are absent in Confirciusornis sanctus and ostrich (Holmgren, 1955) are very similar advanced birds. But it is not clear in (fused dorsally in the vertebrae rather than lithoyraphica. In "Ovirup- ventrally in carinate birds). The complete tor" mongoliensis, and Heyuanniu huungi, distal tail of Heyanniu huungi shows that these articulations are very weak, so it was there is no pygostyle. It is also absent in coded here as absent (0). Cuudiptrryr zoui (Ji et al., 1998; Zhou et 108. Pleurocoels on the caudal centra: absent al., 2000). The development of a py- (0); present at least in the proximal part of gostyle was regarded as a synapomorphy the tail ( I ). of the clade composed of all birds more 109. Number of caudals: more than 35 (0); advanced than Archaeopteryv lithographi- 35-25 (I); fewer than 25 (2) (modified ca (Gauthier, 1986). Cracraft (1986) con- from Lii et al., 2002). Most theropods sidered it as synapomorphy of Ornithurae, have more than 35 caudals. 22 caudals are whereas Sanz and Bonaparte ( 1992) con- present in Cuudipte~yrsp. (Zhou et ul., sidered it as synapomorphic for Iberonw- 2000). 24 caudals in Non~ingiugohiensis sornis plus Ornithurae. In agreement with (Barsbold Osmolska er al., 2000), about Sanz and Bonaparte ( 1992). Chiappe 33 in "Oviraptor" mongoliensis. There are (1996) considered it as a synapomorphy about 26 caudals in zhaoiunus of the . The discoveries of and 25 in Shenzhouraptor (=, Jeholornis and further supports Zhou and Zhang, 2002; Ji et al., 2002; Sanz and Buscalioni ( 1992) and Chiappe 2003). (1 996)'s hypotheses (Chiappe, 2002). 110. Pygostyle: absent or rudimentary (fewer Scapular caudal end: blunt and much than three elements) (O), present (I). The expanded (0); tapered to a sharp point last three to six caudal vertebrae fuse to or slightly expanded (1) (modified from form a pygostyle in modern birds (Baumel Chiappe, 2002). and Raikow. 1993) that support the tail Scapular and coracoid: nearly in the . A pygostyle is present in the or- same plane (0); forming a distinct angle nithurines, hesperorniforms and ichthyor- (1) at the level of the glenoid cavity nithiforms, enantiornithines (Iheromesor- (Chiappe el al., 1998). nis. , Sereno, 2000; Sereno and Distal caudal prezygapophyses: overlap- Lii. Tomida. Azuma. Dong and Lee ping less than a half of the centrum of the about the proximal third of the humerus preceding vertebra (0); overlapping at length or less (0); about 40-50% of the least a half of the preceding vertebra ( I ). humerus length ( I). Hypapophyses in the cervicodorsal verte- Epicondyles on the humerus: absent or bral region: absent (0); small (I ); promi- poorly developed (0); the ectepicondyle nent (2). The hypapophyses are absent in more prominent than the entepicondyle Coeloph~vsis bauri, Herrerasaurus is- ( 1 ); the entepicondyle more prominent chigualastensisi, Allosaurus ,fragilis, Mi- than the ectepicondyle (2); the ectepi- crovenator celer, ornithomimids, tyran- condyle and entepicondyle about equally nosauridae. Because it is absent in Micro- prominent (3). r-aptor. present in Velociraptor mongolien- Shaft of the ulna: straight (0); bowed con- sis, this character is coded here as 011 in vex caudally (1 ); bowed concave caudally Dromaeosauridae. It is not prominent in (2). Heyuannia huangi, Caudipter.vx sp., and Radius length to humerus length ratio: Nemegtia barsboldi. It is well developed 0.80 or less (0); 0.85 or more (I ). in "Oviraptor" mongoliensis, Conchorap- Distal carpals: flat, mostly separate (0); tor gracilis, Chirostenotes pergracilis, carpals 1 and I1 separate, carpal I with Avimimus portenrosus, and troodontids. the proximal trochlea (1); carpals I and Distal chevrons: deeper than long (0); I1 fused, half-moon-shaped, with the longer than deep ( I ). trochlea on the proximal surface, cover- Sternum: unossified or small (0): ossified, ing metacarpals I and 11 (2). The ex- large ( I). panded semilunate (half-moon-shaped) Scapula length to humerus length ratio: carpal block completely caps the proximal 0.8-1 .I (0); 1.2 or more (I); 0.7 or less surface of metacarpals I and I I in ovirap- (2). torosaurs, dromaeosaurids and avialians. It Acromion: projecting dorsad (0); everted is regarded as diagnostic of . laterad ( I); projecting cranial (2). The absence in ornithomimosaurs and Caudoventral process on the coracoid: ab- adult tyrannosaurids are treated as rever- sent (0); short, not extending beyond the sals (modified from Holtz, 200 1 ). glenoid diameter (I ); long, caudoventrally Combined lengths of manual phalanges extending beyond the glenoid (2). 111-1 and 111-2: greater than the length of Orientation of the glenoid on the pectoral phalanx 111-3 (0); less than or equal to the girdle: caudoventral (0); lateral ( I). length of phalanx 111-3 (I). Deltopectoral crest: low, with the width Metacarpal I length to metacarpal I1 equal to or smaller than the shaft diameter length ratio: 0.5 or more (0); less than 0.5 (0); expanded wider than the shaft diame- (1 1. ter ( I). Proximal margin of the metacarpal I in Internal tuberosity on humerus: weakly dorsal view: straight, horizontal (0); an- pronounced or absent (0); well pro- gled due to the medial extent of the carpal nounced but low (I); subtriangular, dis- trochlea ( I). tinctly extended medially (2); in form of a Metacarpal 11 relative to metacarpal 111: longitudinally short, tuber-like extension, shorter (0); subequal (I); longer (2). In sharply delimited from the shaft and usu- Heyuannia huangi, and lngenia j~anshini, ally also from the humeral head (3). the length of metacarpal 111 is shorter than Deltopectoral crest (measured from the that of metacarpal 11; In Conchoraptor humeral head to the apex) extending for: gracilis, the length of metacarpal 111 is New Oviraptorid Dinosaur. Mongolia 125 longer than that of metacarpal II; In Supracetabular crest: well developed (0); Cuudiptry~s,metacarpal I1 is subequal to reduced or absent ( 1 ). metacarpal I I I. Craniocaudal length of the pubic pe- Metacarpal I1 length to humerus length duncle: about as long as the ischiadic ratio: 0.4 or less (0);more than 0.4 ( 1). peduncle (0); distinctly longer than the Metacarpal 111: unmodified (0); very slen- ischiadic peduncle (1); distinctly short- der (1). er than the ischiadic peduncle (2) (mod- Lip or nubbin on the proximodorsal edge ified). of the manual unguals: absent (0); present Dorsoventral extension of the pubic pe- (1). duncle: level with the ischiadic peduncle Manus length to humerus length plus ra- (0); deeper than the ischiadic peduncle dius length ratio: 0.50-0.65 (0); more than (1). 0.65 ( I); less than 0.50 (2). Brevis fossa: absent or small (0); large ( I ). Manus length to femur length ratio: 0.3- Antitrochanter on the ilium: present (0); 0.6 (0); more than 0.7( 1 ); less than 0.2 (2). absent ( 1 ). Humerus length to femur length ratio: Ilium length to femur length ratio: 0.5-0.7 0.5-0.6 (0); less than 0.4 ( I); 0.7 or more (0); 0.8 or more ( I). (2). Hypocleidium on absent (0); or Dorsal margins of the opposite iliac present (I). The hypocleidium is a blades: well separated from each other (0); process extending from the ventral mid- close to or contacting each other along line of the furcula, and is attached to their medial sections ( I). the sternum by a ligament in extant Dorsal margin of the ilium along the cen- birds (Hwang et al., 2004). tral portion of the blade: straight (0); Pubic shaft: straight (0); concave cranially arched ( I). (1 ). Preacetabular process relative to the Pubic foot: with the cranial and caudal postacetabular process (the lengths mea- processes being about equally long (0); sured from center of the acetabulum): with the cranial process being longer shorter or equal (0); longer ( 1 ). than the caudal process (1); with the Preacetabular process: not expanded or cranial process being shorter than the weakly expanded ventrally below the level caudal process (2); absent (3) (state 2 is of the dorsal acetabular margin (0); ex- modified). panded ventrally well below the level of Dorsoventral length of the pubic apron: the dorsal acetabular margin ( 1 ). longer than half total length of the pubis Morphology of the ventral margin of the (0); not longer than the half of total length preacetabular process: the cuppedicus of the pubis ( 1 ). fossa absent, the margin transversely nar- Caudal margin of the ischiadic shaft: row (0); the cuppedicus fossa or wide straight or almost straight (0); strongly shelf present ( I ); the margin flat, wide at concave (1 ). least the base of the pubic peduncle (2). Position of the obturator process on the is- Cranioventral process on the preacetabular chium: proximal (0); at about mid-length blade: absent (0); rounded (I); hook-like ( 1); distal (2); obturator process lacking (2). (3). Distal end of the postacetabular process: Distal end of the ischium: not expanded truncated or broadly rounded (0); nar- (0); expanded ( I). rowed or acuminate ( 1). lschium length to pubis length ratio: 0.75 Lii. Tomida, Azuma. Dong and Lee or more (0);0.70 or less ( I). (1). Because the metatarsi of some or- Posterior (greater) trochanter: weakly sep- nithomimosaurs (Carudimimus and arated or not separated from the femoral Harpymimus) are not strongly pinched head (0); distinctly separated from the as in more typical ornithomimids, this femoral head ( 1). character was coded as (0,l) for Or- Craniocaudal extent of the posterior nithomimidae by Holtz (2001) in his trochanter of femur: short (0); long (I ). phylogenetic analysis. This character is Pleurocoels absent on sacral vertebrae coded here as (0, I), in agreement with (0); or present on anterior sacrals only Holtz, instead of being coded as (1) as in (I); or present on all sacrals (2). A pleu- Maryanska et al. (2002). rocoel may be present on the first sacral in 173. Metatarsals I1 and IV: not in contact on Alxasaul-us elesitaiensis, although this the plantar surface (0); contacting distally area is badly crushed (Russell and Dong, (1). 1993) (Hwang et al.. 2004). 174. Metatarsal I length: more than 50% of Anterior and posterior trochanters: well metatarsal I1 length (0); less than 50% of separated (0); contacting (1 ); fused (2). metatarsal I1 length (1 ); metatarsal I ab- Dorsal extremity of the anterior trochanter sent (2). of femur: well below the posterior Metatarsal IV length relative to metatarsal trochanter (0); about level with the poste- I1 length: about equal (0); longer ( I). rior trochanter ( I). Epipterygoid present (O), absent (1) Fourth trochanter of femur: well devel- (new). An epipterygoid is known in al- oped (0); weakly developed or absent ( 1 ). losaurids (Madsen. 1976). ornithomimids Adductor fossa and associated craniome- (Barsbold 198 1). tyrannosaurids (Molnar. dial crest on the distal femur: weak or ab- 1973), DI-omueosaurus alhertrnsis (Cur- sent (0); well developed ( 1 ). rie. 1995), the derived oviraptorid Citipati Strong distal projection of the fibular osmolskue (Clark et al.. 2002) and kmeg- condyle on the femur: absent (0); present tia harsholdi. It is not clear in Inci- ( 1). si\~osaurus gauthieri, and it is absent in Medial surface of the fibular head: flat or "Ovil-aptor*" mongoliensis and birds. shallowly concave (0); with a deep fossa Metatarsus length to femur length ratio: ( I ). 0.4-0.6 (0); about 0.3 (1 ); 0.7-0.8 (2). Contacts of the distal end of the fibula Thoracic vertebral cohnt: 13-14 (0): with tarsus: present (0); absent (1). In 11-12 (1); fewer than i1 (2) (Chiappe, most theropod dinosaurs, and H+anniu 2002). huangi, the distal end of the fibula con- Ossified uncinate processes: absent (0); tacts the tarsus, but in Ingeniu ,vanshini. present ( I). this contact is absent. similar to the case in Coracoid shape: short (0);elongated with most birds. trapezoidal profile ( I ); strut like (2). Ascending process of the astragalus: as The proximal end of metacarpal Ill: tall as wide across the base (0); taller than contacts the distal carpals (0); does not wide ( 1 ). contact (I) (new). Distal tarsals: not fused with metatarsals Metacarpals I and 11 fused proximally: (0); fused with metatarsals ( I ). absent (0); present (I) (new). Proximal coossification of metatarsals Ectopterygoid with constricted opening I l -I V: absent (0); present ( 1). into fossa (0): or with open ventral fossa Arctometatarsus: absent (0); present in the main body of the element (1) New Oviraptorid Dinosaur. Mongolia 127

(Hwang et al., 2004). tor gruc~lis. "Oviruptor" mongoliensis, Flange of pterygoid well developed (0); Nemrgtia bursboldi and Oviruptor philo- or reduced in size or absent (I) (Hwang ceratops. the external nasal openings are et al., 2004). elongate. In majority of theropods, the ex- Symphyseal region of dentary broad ternal nasal openings are oval (elongate), and straight, paralleling lateral margin such as in Herrrrnsaurus ischiguulusten- (0) or medially recurved slightly (I); or sis, Coeloph.vsis huuri. Allosaurtrs fragilis, strongly recurved medially (2) (Hwang ornithomimids, troodontids and dro- et al., 2004). maeosaurids. Mandibular articulation surface as long The maxillary fenestra is relatively as distal end of quadrate (0): or twice smaller or absent (0), or larger than the or more as long as quadrate surface, al- antorbital fenestra (1) (new). The maxil- lowing anteroposterior movement of lary fenestra is usually larger. opens later- mandible (I) (Hwang et al., 2004). ally, and is clearly visible in lateral view Maxilla toothed (0); or edentulous (I) (Rauhut, 2003). It is relatively smaller (Hwang et al., 2004). in Incisivosaurus gauthier-i, Cuudipteryx The lower margin of the external nasal dongi and "Oviruptor" mongoliensis. It is opening is below (0). or close or above relatively larger in lngmiu yunshlni (I) the level of the upper corner of the (GIN 100/80), Citipati osmolskae. and Ne- antorbital fenestra (new). The lower rnegria bursboldi. It was thought that in margin of the external nasal opening is caenagnathid Chirostenotes pergrucilis below the level of the upper corner of the (ROM 43250), there was no maxillary antorbital fenestra in Coe1ophvsi.s hauri, fenestra (Rauhut, 2003). but according to Herrerasaurus ischigtlalustensis, kloci- Clark et ul. (2002. Fig. 12). a relatively ruptor mongoliensis, troodontids, or- larger maxillary fenestra is present in Chi- nithomimids, Tyrannosauridae, Allosaurus rostenotes pc.rgrucilis as in Ovirvptor fi-ugilis. Catrdiptery sp., Oviruptor philo- philocrrutops. cwutops, Incisivosuurus guuthieri, Khaun Large openings absent (0), or present mckennui, and lngrniu ( 100180). It is close (0) on the base of the neural arches of in Citiputi osmolskae (Clark et a/.. 2002). anterior caudal vertebrae (new). On the above in Conchoraptor grucilis (Barsbold anterior caudal vertebrae of Ingrnia -van- er ul., 1990), .Wemegtia harsboldi, and shini and "Oviruptor" mongolirnsis, there "Ovirupror" mongoliensis. are large openings on the lateral surface The quadratojugal and quadrate con- near the base of the neural arches, these tact area is far from (0), or near (1) the openings are larger than the pleurocoels; lateral surface of the quadrate articular In Allosaurus fmgilis, He.vucrnnia huungi, surface (new). Conchorup/or gmcili.~, and Cutrdiptrr:w Nasals and premaxillae do not form a sp.. this kind of opening is absent. crest (0), or form a crest (I) (new). Large openings (fossa) present (0), or External nasal opening round (0), or absent (I) on the neural arches of cervi- oval (1) (new). In Incisivosa~crtrs cal vertebrae (new). Large openings are guurhirri, the external nasal opening is present in the neural arches of cervical round in Cuudipterm sp. and Citiputi os- vertebrae in Coe1ophysi.s huul-i, Ar- molskue, the external nasal opening is chueoptcyw lithogruphica, Aviminlus por- nearly round (slightly elongated). In other tmtosus. Microlvnutor- ccler, Conchorup- derived oviraptorids, such as Concho~vp- tor grucilis, Hevuunniu huungi. They are Lii. Tomida. Amma. Dong and Lee absent in Allosaw-us fragilis, Her- (Makovicky and Sues, 1998), lngenia (GI r~erasuurtrs ischigualustensis, lngeniu 1 OO/3O), Chirostenotes per.gr.ucilis and (GIN IOO/32-02), Caudipreyx sp., Ne- Avinzimzis po~~eti/osus.Although in Avim- megtiu barsboldi, and "Oviraptor-" mon- imus portentosus, a second pair of open- goliensis. ing is present in some vertebrae. it is Constriction between articulated pre- coded here as (0), in contrast to Rauhut maxillae and maxillae: absent (O), pre- (2003). One pair is present in Allosaur-us sent (I). (Rauhut, 2003). In most of fragi1i.s. "Oviraptor" mongoliensis and theropod dinosaurs and a basal ovirap- Hql~uunniuhuangi. torosaur, Incisivo.suun(s gauthieri, there is 199. Pleurocoels developed as: deep depres- no pronounced constriction, but in derived sions (0); foramina (1) on cervical verte- forms such as Oviraptor philoceratops, brae (Rauhut, 2003). Pleurocoels devel- Conchoraptor gracilis, Citipati osmolskae, op as deep depressions in Coe1oph.v.si.s "Oviraptor" mongoliensis, and Nemegtia hauri. Microvenator celer, "Oviraptor" hursholdi. this constriction is distinct. mongoliensis. lngenia ,vanshini; as foram i- Shape of nasals: expanding posteriorly na in Archaeopteryx lithographica, Avinl- (0); of subequal width throughout their innrs portentosus, Conchoraptor gracilis, length (I). (Rauhut, 2003). The nasals Hevuannia huungi. Nemegtia harsholdi, expand posteriorly in Incisivosatirus gau- and Allosazrrus,fi.agilis. thieri. and they are very broad and widen 200. Nasal fusion: absent, nasals separate posteriorly in Conchor*aptor. But the (0); present, nasals fused together (1) nasals are of subequal width in Oviraptor (Holtz, 2000). philoceratops, Citipati osmolskae, "Ovi- raptor" mongoliensis, and Nemegtia burs- Appendix 3 boldi. Characters and data matrix: Character state 0 Postorbital part of the skull roof: as is plesiomorphic; character state 1-3 are apomor- high as the orbital region (0); deflected phic. "-" is not comparable character state; "?" ventrally in adult individuals (1). is either not preserved or unknown. [ ] means (Holtz, 1994). In most dinosaurs, the pari- two character states included. etals and the medial parts of the squamos- als are approximately level with the Allosauroidea frontals above the orbits, and their sur- oooo[ol]ooooolool[ol]loololoo[ol]l l[Ol]OOO faces face dorsally. In Avimimus portento- 0 1 [02]000000 1 100000000000000000000[01 I000 sus. Archaeoptecvx lithographica, Inci- 0 l0000000[0 l]OOO[O I] 1000000000000002[0I] I sivosatirus gauthieri, derived ovirap- 0010000001000001[01]00l0l002120010l02000 torosaurs, ornithomimosaurs and - 10 1000002000[0I ]1 [O 1 ][O 1 ]00[02][O 1 I00 100000 tids, the postorbital part of the skull roof 0010100000010000'?000000000001101000110 deflected ventrally, showing very devel- oped braincase. Ornithomimidae Number of pleurocoels in cervicals: ab- 0000000 100000200000[011 [O 1 I000 I I 10000 1 1 10 sent (O), two, arranged horizontally (I); 10020[01]0[01]010[01]0[01]0010000000????0l one (2) (modified from Rauhut, 2003). 0020 1 I OOO[O I ]000[01] 10000000002[0I ][O I ][O I ] There are no pleurocoels in Herrerusaurus 00l00l00[0l]011[0l]l0000010l0002001002001 ischigualastensis and Caudipteryx sp. Two 001000[02][01][02]1001 l200l0111?001001000 pairs of pneumatic openings are present in 001 l l I I01 10[01][01]2101002000000[01]000lI Coelophysis bauri. Microvenator celer 010'?10-0 New Oviraptorid Dinosaur. Mongolia 129

Dromaeosauridae OO[O I ]0000000[0 11002 100 1 10 I00 lo I 10000 1 100 0001001010000000000000000000010[01]00000 0010001 100000000000000101 l[OI]I 1111O[OI]l 0000[01]111102[12]1[01]2021021[01]12011 I I2 OOOOl l[Ol]l l1001002101[01]10[01]0l1[01]000 001000110001100000000?010010100-0

Troodontidae 002 1 1002000001000 100100 I'? I1??O?O 1 O'? 1 O? 1 0 020[0 1 ] 10000 1 1 00000'?00??'??'???10000000 l ?OO 1 1000?'?0'?0'?'?0?0'??1 [O 1 ]O'?O'?1 1000 100 1 1 10 1 lo '?12'?1'?0?1121 1111 101 1011001 110'?1000'?03'?01 101 101 11???11'?01111'?10010000[01]'?00?0llo? "Oviruptor" mongoliensis 00 10-0 112'?0111110112121111?111121010112111111 12'?11122111?011111112111101?111II2120120 Archueoprey lithogr-uphica 01111111112111l11001111011'?010011021111 OO? 1 1 OO?OO 10 1 10 1 1000 1000 1??01 1 1 O? 1 ?00020 21 120100??'???'??1?'?0ll0l121110001I???????] ? 1 ??OOO??O'?O'?OOOOOO l '? 1 '??'?'??OOOO1 123000 1 1- 11 1111 1?????'????1?1'?'???11211111101'?111101 00000000200001??010 1 1 00'?1?0 I I 1000 I I222 10 00?l1211111110120110101111'?100021-?-ll?? GIN 1 00142 ?0100'?0'?110'?1011001100100000010000'?101? 112001101101101211111111121010112111111 121111221110011111112111101011111212012 Avimimus portenrosus 001111111112111111011111011111?01?02110 ??'?'??1 ???O?????'???'?1 ? I?1 ?3? 1 1 12 1 3??OO- 1 ?2 12012011120111111110100I211I00001111111 1 1 1 ? 1 ?? 1 1 0 1 ???'??????????OO???1 1 ??? 1 ??'??? 10 01111111110?100001010?11??11211?????????? OOO???1 O? l1000 1 1 0 1 1 1 1 O'? 1 02????2'??0 1 200 1 1 ??? 1 1 12? 1 '???'????00000?01 1 O?OO 1?0 1 1 O?? 1 1 100 10 0 1 1 0 1 1 1 1 1 20? 1 '?'??O1 ??2 1 ???O??OO?? 1 1 I? Conchoraptor gracilis 102?011?11011012111101111210101121111 Caudipteryx zoui 1112111122111?011111II21I1101?111112120 101'?100?1??110111111l0001??0101l2000l1?1 120011111111121111110111110111'?10011011 11???? 1 ?0??'?0??00???~????'???0I 1 10002? 1 100 1 0 111201211112001111111010002l01000011111 0 1 ? 10 1 ? 1 0 1 1 100000000?00[0 1 ]0020000 1 00200 1101111111110?100001010??1??1?2111101'?00 00000110-101010100000lll0ll101001111101 110'?0100001000010?121010'?'?2110100001I?? 0-0 Ingenia j)unshini 102?011?11011'?121111111112101011211 Chirostenotes pergradis ??I 11?2?11'???1'??11121????1?111112I20

??I? 1 ???OO 1 0 1 ????1 ???0?????????????'??????2?I 111111112111?110011110110?10011010 o'? 1 2'???0?'?0'???????'????1 12 10 1 1002 1 1 100 1 10 1 1 1 1 1 12?1 1 ???'?O? 1 ? 1 1 1 1 1 1 1 ?????1 ???? 1 ??????? ??????I 1???'?1 I'???! 110'????0111 101??1'??1?1?? ?O? 1 0 1 OOO?? 10???2'?1 ????1 ????? 1?? Heyzmnnia huungi

Nomingiu gohiensis 1 O??? 1 1 ???????????? 1 ?'??I????? 1 O???1 1 1 1 1 1 ?2? 130 Lii. Tomida. Azuma. Dong and Lee

Citiputi osmolsliue Khuun mckennui 112101111101121211111111121010112111100 10210'?1?1?0112121111111012001011211'?1111 121111221'?100111l111211llOllll11121?112 ? 1 '????2????'?1 ????? 1 ??????? 1 l l l 1 12 12 1 1 1 ???? 00 1 l l l l l l 1 1 1 1 1 1 1 ?'???'??'?'?'??'?I?'???'???'??I?12? ? I I 1 ?? I ? I 1 1 I ??O 1 ???????????OO1 O'????'??????? 1 1 1 '?O 1 ?0 1 '? 1 10 1 '???'???'?????'???'??1 1 ?'?'??'?'????'?'? 1 1 ?0'???'?'??'?'?0?000?1 O????'??? 1 ??O 1 ??????????? '?'?'?OO?'?OO'?1 1 O??1 ???1 12 1 1 1 1 10 1 ??1 1 1 ??1 ?????????????1???2 1 10 10 1 O?'???I?? 1