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(Dinosauria, Theropoda) from the Late Cretaceous of Argentina

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(Dinosauria, Theropoda) from the Late Cretaceous of Argentina Reu. Mus. Argentino Cienc. Nat., n.s. 611): 61-66,2004 Buenos Acres, ISSA' 1514-5158 Unquillosaurus eeibali Powell, a giant maniraptoran (Dinosauria, Theropoda) from the Late Cretaceous of Argentina Fernando E. NOVAS' & Federico L. AGNOLINP 'CONICET. ZMi~seoArgentino de Ciencias Naturales aBernardino 12ivadaviam, Ax A. Galiordo 4.70; C1405DJR Buorios Aires, Argentina. Telpax: 4981 9282, email: [email protected]. Abstract: A reviewed anatomy and phylogenetic relationships of the Late Cretaceous theropod Unquillosauriis eeibali is presented. This taxon is represented hy an isolated pubis Crom 1,os Blanquitos Formation (Nlaastrichtian),from the province of Salta, NW Argentina. The size of the bone (51 cm long) originaliy lead to interpret chis theropad as a member of the large "Carnosauria". However, the pubic anatomy of Unquilloseurus is more eonpuent with that. present in metornithine caelurasaui.ians, a group including Alvarezsauridae, Theririnosauroidea, Oviraotorosauria. Dcinonvchosauria, and Aves. Derived features presont in U~im~illosaurus Moreover, some Laits sharEd with eariy avians (e.2, pubic foot proximodistally tall and craniocaudally short), suggest that this dinosaur may be more ciosoly related to birds than suspected. The set of autapomorpiiic Foatures recognized in the pubis indicates that Unquilloseurus may belong to an endemic lineage of large maniraptaran theropods. Key words: Thoropoda, Man~raptora,Late Cretacoous, Argentina Uuguillosaurus ceibali was described by Jai- derived linenlagia comahuensis (Novas & Puer- me Powell (1979) on the basis of an almost com- ta, 1997). Thisvariety of dinosaurs offers valuable plete left pubis, discovered in Maastrichtian beds anatomical information that invites to reconsider CrornSaltaProvince, hW Argentina (Fig.1).Powell the phylogenetic relationships of Unquillosaurus. (1979) interpreted this dinosaur as a large carnosaur of uncertain reiationships. With the MATERIAL AND METHODDS exception of some brief coruments made by Molnar et al. (1990) and Bonaparte (1996), no further Institutional abbreviations. AMhW, American reierences to the phylogenetic relationships of this Museum of Natural History, New York; HMN MB, theropod are found in the literature. The Humboldt Museum fur Naturkunde, Berlin; fragmentary nature of Unquillosaurus, coupled MACN, Museo Argentino de Ciencias Naturales with the limited knowledge of Gondwanan <<B.Rivadavia., Buenos Aires; MCZ, Museum of theropods available until recently, delayed the Comparative Zoology, Cambridge; PVL, elucidation of the rela,tionships of this dinosaur. Paleontologia do Vertebrados, Fundaci6n ~Miguel In the last years Che fossil record of Argentine Lillom, San Migucl de Tucuman; PVPH, Cretaceous theropods has been favorably PaleontolodaVertebrados, Museo Municipal <<Car- Coelnrosauria (&., Bonaparte, 1991,1996; Novas, WM, Yale Peabody Museum, New Haven. 1997,1998; Coria& Salgado, 1995). Other findings Studied specimens: The following specimens demonstrate that South America was inhabited have been used for this study: Patagonykuspuertai by a diversity of non-avian coelurosaurian (PVPH 37),Albertosaurus libratus (AMNH 5468), theropods, including Santanaraptor (Kellner, Allosaurus fragilis (AMNH 6767), Archaeopteryx 19991, a possible oviraptorosaur (Frankfurt & lithographica (HMN MB 1880181, and casts of Chiappe, 1999), different alvarezsaurids London and Eichstatt specimens), Carnotaurus (Bonaparte, 1991; Novas, 1997), and the highly sastrei (MACN-CH 8941, Ceratosa~lrusnasicornis 62 Reucsla del Museo Argenlcno de Ciencias Naturales, n. s. 6 ilj,2004 (USNM 4735), Deinonychus antirrhopus (AMNII Tble I. Measurements of pub~sof 1Jnquzllosaurus 3015, MCZ 4371, YPM 5205, 5206, 52361, cezbalz Piatni:zkysaurus floresi (MACN-CH 8951, Rahoo#auis ostromi (UA 8656), and Unenlagia Proximodistal length 514 mm comahuensis (PVPH 78). Craniocaudai diameter of proximal end 155 mm % Sysiematic ~zomenclature.In this paper we are Craniocaudal diameter of distal foot 11.5 mm followingthephylogenetic hypothesis andsystem- Greatest transverse diameter of shaft 36 mm atic nomenclature employed by Xu et al. (2002) Lesser transverse diameter of shaft 56 mm for coelurosaurii~ntheropods. SYSTEMATIC PALEONTOLOGY Ceratosauria (e.g., Carnotaurus, Ceratosaurus; Theropoda Marsh, 1881 Gilmore, 1920; Bonaparte et al., 1990) and Trtanurae Gauthier, 1986 Tyrannosauridae ~Alberlosauruslibratus AMNH Coelurusauria Huene, 1920 5468).In contrast, its reduced size resembles more Maniraptora Gauthier, 1986 that of derived coelurosaurians, such as Metorn~thesPerle, Norell, Chiappe & Clark, Oviraptoridae, Alvarezsauridae, Dromaeosauridae 1993 (e.g., Unenlagia, Bambiraptor; Novas & Puerta, 1997; Burnham et al., 2000), and primitive birds Genus Unquillosaurus Powell, 1979 (e.g., Archaeopteryx, Rahonauis; Wellnhofer, 1974; Forster et al., 1998). On the other hand. the ischi- Type species. Unquillosuurus ceibali Powell, 1979 adic facet of the pubis is dorsoventrally low, compa- rable to that oSDmmamsauridae, but different from Sir-atigraphic distribi~tion.Los Blanquitos For- the well develoued one of most non-coelurosaurian mation, Salta Group (Masstrichtian; Salfity & theropods (e.g., Ceratosaurus, Abelisauridae, Marquillas, 1999). Piatnit&saurus, 'fyrannosauridae; Gilmore, 1920; Geographic distributioiz, Arroyo Morterito, Si- Bonaparte el al., 1990; Bonaparte, 1986; Osborn, erra de la Candelaria, Salta Province, hW Argen- 1917). The ohturator fbramen isventrally open, as tina. it occurs in most tetanurans (Gauthier. 1986) (Fig. 2). Unquillosauncs ceibali Powell, 1979 The intersection between the proximal mar- (Fig. 1) gin of the pubis and the longitudinal axis of the pubic shaft describes an angle of approximately Emendeddiugmsis. The pubis of linquillosaums 45 degrees, reflecting an opisthopubic condition is proportionaliy long and slender It is large (61,4 ofthe bone (Fig. 2). An opisthopubic pelvis is diag- cm long; Table 1) in comparison with other basal nostic of Metornithes (Xu et al., 20021, as it is Maniraptora (= Ornitliolestes + Metornithes; Xuet present in Alvarezsauridae (Novas, 1997), al., 20021, such as Patagon.ykus puertui PVPI-I 37 Therizinosauroidea (Barsbold, 19791, and Paraves ('25 cm), Deinonychus antirrhopus MCZ 4371 (38 (=Deinonychosauria + Aves; Sereno, 1997; Clark cm), Urlenlngia comahuensis PVPH 78 (32 cm), et al., 2002; Xu et al., 2002). Rahonauis ostrorni UA 8656 (6 cm). Its proximal The proximal 3/n of the pubic shaft are end presents a thick edge defining a deep groove, craniocaudally compressed, in contrast to most which was originally interpreted by Powell (1979) theropods (e.g.,Allosaurus,Pataponykus;Gilmore, as diagnosticoSUnp.*illosaum,sceibali.However, this 1920; Novas, 1997) in which the pubis is rod-like abnormal morphology (not recorded in other proximal to the pubic symphysis. The condition saurischian dinosaurs) is better explainedas the bro- mentioned above for Unquillosaurus results in a ken pubic pedicle of ilium adhered to the external prominent external, longitudinal ridge (named surface of pubis. In fact, this piece of bone resembles "crcsta lateral" by Powell, 1979; Fig.1 A, Bi. The the pubic pedicle of the ilium of Deinonychw and margins of the pelvic canal of Unquillosaurus are Unenlagia (Ostrom, 1969; Novas & Puerta, 1997) medially convex, differing from most theropods in beingcraniocaudally extended, ventrally concave, (e.g., Allosaurus, Tyrannosaurus, Patagon.ykus, and with the cranial and ventral margins formingan Velociraptor; Gilmore, 1920; Norell & iWakovicky, angle exceeding 90 degrees in side view (Fig. 1A). 1999; Novas, 1997; Osborn, 1917) in which the The proper acetabular surface is represented margins are medially concave (Fig. 3). The by a reduced surface located caudally to the facet craniocaudal compression of the proximal pubis of for the pubic pedicle of the ilium. The acetabular Unquillosaurus contrasts with the craniocaudal surface of 1Jnquillosaurus is more reduced than in expansion of its distal half, thus resultingin alon- Nocas & Agnolcn Unqu~llosauri~s,a large man~raptorantheropod A D E Fig. 1. Unquillosaurus ceibali left pubis. A) lateral, Bj cranial, Cj distal (restored), Dj medial, and E) caudal.riews. AS, acetabular surface; CaE: caudal process of pubic foot; CrP, cranial process of pubicfoot; FIE facet for the iliac pedicie; LR, iateral ridge; PF pubic foot; PPI, Iliacpubic pedicel. Scale bar 10 cm. A B C D E Fig. 2. Pubis of several theropods in lateral view. Not to scale. A) Unquillosaurus, B) Unenlagia, C) Ouiraptor, D) Tyrannosaurus,E)Piatnitzkysaurus. 64 Reuzsta del Museo Argentzno de Czenczas Natarales, n s 6 (I), 2004 A B C D Fig. 3. Reconstructed pubises of several theropods in cranial view. Not to scale. A) Ornithomimus, B) Patugonykus, C) Unenlagia, D) Unquillosaurus (without the pubic pedicle of ilium). gitudinal "twisting" of the bone (Fig. I). In most & Puerta, 1997; Burnham el al., 2000) in which it theropods, instead, the cranial and caudal mar- is well developed. However, in Unquillosaurus the symphysis is absent (Figs. 1, 31, unlike most theropods (e.g., Allosaurus, Carnotaurus, DISCUSSION Ornithominzus, Velociraptoq Unenlagia, Rahomvis, Archaeopteryx; Gilmore, 1920; Bonaparte el al., The pubis of Unquillo.saurus clearly indicates 1990; Osborn, 1917; Norell & Makovicky, 1999; that this genus is not a member of Abeiisauroidea Novas & Puerta, 199'7; Forster et al., 1998; (=Noasauridae + Abelisauridae; Novas, 1992; Wellnhofer, 1974) in which the apronis transversely Carrano
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