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The Evolution of Gondwanan Dromaeosaurids a Bizarre Downloaded from rspb.royalsocietypublishing.org on 25 February 2009 A bizarre Cretaceous theropod dinosaur from Patagonia and the evolution of Gondwanan dromaeosaurids Fernando E Novas, Diego Pol, Juan I Canale, Juan D Porfiri and Jorge O Calvo Proc. R. Soc. B 2009 276, 1101-1107 doi: 10.1098/rspb.2008.1554 Supplementary data "Data Supplement" http://rspb.royalsocietypublishing.org/content/suppl/2009/02/20/276.1659.1101.DC1.ht ml References This article cites 24 articles, 2 of which can be accessed free http://rspb.royalsocietypublishing.org/content/276/1659/1101.full.html#ref-list-1 Subject collections Articles on similar topics can be found in the following collections evolution (427 articles) graph theory (1 articles) Email alerting service Receive free email alerts when new articles cite this article - sign up in the box at the top right-hand corner of the article or click here To subscribe to Proc. R. Soc. B go to: http://rspb.royalsocietypublishing.org/subscriptions This journal is © 2009 The Royal Society Downloaded from rspb.royalsocietypublishing.org on 25 February 2009 Proc. R. Soc. B (2009) 276, 1101–1107 doi:10.1098/rspb.2008.1554 Published online 16 December 2008 A bizarre Cretaceous theropod dinosaur from Patagonia and the evolution of Gondwanan dromaeosaurids Fernando E. Novas1,*, Diego Pol2, Juan I. Canale3, Juan D. Porfiri4 and Jorge O. Calvo4 1CONICET, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Avenida A´ ngel Gallardo 470, Buenos Aires 1405, Argentina 2CONICET, Museo Paleontolo´gico Egidio Feruglio, Avenida Fontana 140, Trelew, Chubut 9100, Argentina 3CONICET, Museo Municipal Ernesto Bachmann, Dr. Natali S/N, Villa El Choco´n 8311, Neuque´n, Argentina 4Centro Paleontolo´gico Lago Barreales-Proyecto Dino, Universidad Nacional del Comahue, Avenida Megaraptor 1450, Lago Barreales, Neuque´n CC.122, Argentina Fossils of a predatory dinosaur provide novel information about the evolution of unenlagiines, a poorly known group of dromaeosaurid theropods from Gondwana. The new dinosaur is the largest dromaeosaurid yet discovered in the Southern Hemisphere and depicts bizarre cranial and postcranial features. Its long and low snout bears numerous, small-sized conical teeth, a condition resembling spinosaurid theropods. Its short forearms depart from the characteristically long-armed condition of all dromaeosaurids and their close avian relatives. The new discovery amplifies the range of morphological disparity among unenlagiines, demonstrating that by the end of the Cretaceous this clade included large, short-armed forms alongside crow-sized, long-armed, possibly flying representatives. The new dinosaur is the youngest record of dromaeosaurids from Gondwana and represents a previously unrecognized lineage of large predators in Late Cretaceous dinosaur faunas mainly dominated by abelisaurid theropods. Keywords: Theropoda; Dromaeosauridae; Unenlagiinae; Cretaceous; South America; forelimb Abbreviations: IGM, Geological Institute, Mongolian Academy of Sciences, Ulaan Baatar, Mongolia; MCF PVPH, Museo Municipal Carmen Funes, Plaza Huincul, Neuque´n, Argentina; MCZ, Museum of Comparative Zoology, Cambridge, Massachussets, USA; MML, Museo Municipal de Lamarque, Rı´o Negro, Argentina 1. INTRODUCTION 2. DESCRIPTION AND COMPARATIVE ANATOMY Dromaeosaurids are close avian relatives mainly known (a) Systematic palaeontology from the Northern Hemisphere. These are gracile and slender theropod dinosaurs, most of them measuring Theropoda (Marsh 1884) between50cmand3m inlength(Ostrom 1969; Xu Coelurosauria (Huene 1920) et al. 2000; Senter et al.2004; Turner et al. 2007). A Deinonychosauria (Colbert & Russell 1969) few examples from the Northern Hemisphere, however, Dromaeosauridae (Matthew & Brown 1922) reached up to 5 m long (e.g. Achillobator from Asia and Unenlagiinae (Bonaparte 1999) Utahraptor from North America; Kirkland et al.1993; Austroraptor cabazai, new taxon. Perle et al. 1999). In recent years, a few dromaeosaurid taxa have been discovered in Gondwana (Novas & Puerta 1997; Forster et al. 1998; Calvo et al.2004; (b) Holotype Makovicky et al. 2005; Novas & Pol 2005), but their Specimen number MML-195 consists of right frontal and evolutionary history remains poorly understood. Here postorbital, lacrimals, maxillae and dentaries with teeth, we report a startling new example from Campanian– right surangular and prearticular, cervicals 3, 5, 6, 7 and 8, Maastrichtian beds of Patagonia, which acquired a dorsals 2 and 4, isolated ribs and gastralia, right humerus, large body size convergently with the large Laurasian manual ungual of digit III, left pubic shaft, left femur, and taxa and departed from the characteristic morphology right tibia, astragalus, calcaneum, metatarsal III and pedal of dromaeosaurids. phalanges I-2, II-2, III-4 and IV-2 (figure 1). (c) Etymology Austroraptor, from austral in reference to southern South * Author for correspondence ([email protected]). America, and raptor, thief; and cabazai, in honour to Electronic supplementary material is available at http://dx.doi.org/10. the late He´ctor Cabaza, founder of the Museo Municipal 1098/rspb.2008.1554 or via http://journals.royalsociety.org. de Lamarque. Received 27 October 2008 Accepted 14 November 2008 1101 This journal is q 2008 The Royal Society Downloaded from rspb.royalsocietypublishing.org on 25 February 2009 1102 F. E. Novas et al. Evolution of Gondwanan dromaeosaurids (a) 1m lac fr fsp mx mxf (b) f rpmx af o pan po (c) fsp 10 cm g d st (d)(e) ns ( f )(ns prz g)(h) ns prz poz dp pp 10 cm pp cp (i)(j)(k)(l) h h 10 cm Figure 1. Austroraptor cabazai MML 195, holotype. (a) Skeletal reconstruction and body shape, indicating discovered bones. Body length was estimated following equations described by Turner et al. (2007).(b) Skull reconstruction in left lateral view. (c) Left lacrimal in dorsal aspect. (d ) Cervical 3? in left lateral view. (e,f ) Cervical 8? in (e) left lateral and ( f ) cranial views. (g,h) Dorsal 4? in (g) left lateral and (h) cranial views. (i,k) Pedal phalanx II-2 in (i) left lateral and (k) dorsal views. ( j,l ) Pedal phalanx IV-2 in ( j ) left lateral and (l) dorsal views. af, antorbital fenestra; cp, carotid process; d, dentary; dp, diapophysis; f, foramina; fr, frontal; fsp, fan-shaped process of lacrimal; g, groove; h, caudoventral heel; lac, lacrimal; mx, maxilla; mxf, maxillary fenestra; ns, neural spine; o, orbit; pan, postantral wall; po, postorbital; poz, postzygapophysis; pp, parapophysis; prz, prezygapophysis; rpmx, rostral process of maxilla; st, spine table. (d) Locality and horizon group (autapomorphies marked with an asterisk): lacrimal Allen Formation (Campanian–Maastrichtian; Martinelli & highly pneumatized, with descending process strongly Foriasepi 2004), Bajo de Santa Rosa (408030 2800 S, 668480 curved rostralyÃ, and caudal process flaring out hori- 0300 W), approximately 90 km southwest of Lamarque zontally above orbità (differing from Laurasian dromaeo- town, Rı´o Negro Province, Argentina. Theropod dino- saurids, but unknown for other unenlagiines); postorbital saurs collected from the Allen Formation include the lacking dorsomedial process for articulation with the abelisaurid Quilmesaurus curriei (see Jua´rez Valieri et al. frontalÃ, and with squamosal process extremely reduced 2007), a yet unnamed alvarezsaurid (Agnolı´n et al.2006), (differing from Laurasian dromaeosaurids, but unknown and indeterminate tetanurans (Coria & Salgado 2005). for other unenlagiines); maxillary and dentary teeth small, conical-shaped, devoid of serrations and flutted (as in (e) Diagnosis Buitreraptor); humerus short, representing slightly less A large dromaeosaurid with the following combination of than 50 per cent of femur length (a smaller ratio than in characters that distinguish it from other members of this other dromaeosaurids and paravians); pedal phalanx II-2 Proc. R. Soc. B (2009) Downloaded from rspb.royalsocietypublishing.org on 25 February 2009 Evolution of Gondwanan dromaeosaurids F. E. Novas et al. 1103 Table 1. Comparison of cranial and postcranial elements of selected dromaeosaurid taxa. Velociraptor mongoliensis (Barsbold & Unenlagia Buitreraptor Osmo´lska Achillobator Utahraptor Austroraptor comahuensis gonzalezorum 1999; Norell & Deinonychus giganticus ostrommaysi cabazai MCF PVPH (Makovicky Makovicky antirrhopus (Perle et al. (Kirkland MML-195 78 et al. 2005) 1997, 1999) MCZ 4371 1999) et al. 1993) maxilia length (cm) 39a —10b 10.9c —39— heightd (cm) 8.5 — 1.8 3.5c — 11.6 — height/length 21% — 18% 34%c — 29% — ratio (%) tooth count 24 — — 11c —11— humerus length (cm) 26.2 26.5 b 13.5 — 25.4 — — least mid-shaft 2.2 1.7 0.6 — — — — diameter (cm) femur length (cm) 56 36.8 14.5 23.8e 34.4 50.5 — least mid-shaft 7.1 2.2 0.7 1.67 e — 6.7 — diameter (cm) tibia length (cm) 56.5 41.8 a O14.9 25.5e,f 36 49 50.5 distal transverse 14 6.3 a — 3.8 e — 11.3 14.5 width (cm) pedal phalanx 2.II length (cm) 5.7 — 1.8 2.4g — 7.6 — distal transverse 1.7 — — 0.6g ——— width (cm) pedal phalanx 2.IV length (cm) 4.8 — — 2.1g ——— distal transverse 4.7 — — 0.8g ——— width (cm) estimated body 494.3 323.6 135 208e 302 445 — length (cm)h estimated body 368 95 6 23.4e 77 264 — mass (kg)i aBroken. bEstimated. cV.mongoliensis IGM 100/25. dHeight at level of rostral margin of preorbital fenestra. eV.mongoliensis IGM 100/986. fTibiotarsus. gV. mongoliensis IGM 100/985. hCalculated on the basis of body length regression for paravians by Turner et al. (2007). iCalculated with the equation of Christiansen & Farin˜a (2004)
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