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Agnolin & Martinelli, 2007

Agnolin & Martinelli, 2007

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Cretaceous Research 28 (2007) 785e790 www.elsevier.


Did oviraptorosaurs (Dinosauria; Theropoda) inhabit Argentina?
´n G. Martinelli b Federico L. Agnolin a,*, Agustı

Museo Argentino de Ciencias Naturales ‘‘Bernardino Rivadavia’’, Laboratorio de Anatomı ´a Comparada y Evolucio ´ n de los Vertebrados. Av. Angel Gallardo 470 (C.P. 1405), Buenos Aires, Argentina b Museo Argentino de Ciencias Naturales ‘‘Bernardino Rivadavia’’, Seccio ´ n Paleontologı ´a de Vertebrados. Av. Angel Gallardo 470 (C.P. 1405), Buenos Aires, Argentina Received 11 January 2006; accepted in revised form 24 October 2006 Available online 1 July 2007

Abstract In this contribution a putative oviraptorosaurian cervical vertebra discovered in the Campanian-Maastrichtian El Brete Formation from Salta Province, NW Argentina is analysed. Based on the resemblances of this vertebra with those of the basal neoceratosaurian Elaphrosaurus and with the noasaurid Noasaurus, the Salta specimen is interpreted to belong to the third or fourth cervical vertebra of a Noasauridae (eventually Noasaurus). Furthermore, it is suggested that the supposed anterior cervical vertebrae of Masiakasaurus, Laevisuchus and Noasaurus possibly correspond to a more posterior position than previously considered. Contrary to abelisaurids, the morphology of the anterior cervical vertebrae of noasaurids indicates that they probably were long neck theropods resembling ornithomimid coelurosaurs. Therefore, the occurrence of oviraptorosaurs in Argentina is rejected. Ó 2007 Elsevier Ltd. All rights reserved.
Keywords: Theropod; Abelisauroid; Noasaurid; Argentina

1. Introduction Theropod records from Cretaceous rocks of Gondwana are dominated by medium sized abelisaurid ceratosaurs (e.g., Bonaparte et al., 1990; Bonaparte, 1991a, 1996; Novas, 1997a; Sampson et al., 1998; Coria et al., 2002; Kellner and Campos, 2002) and gigantic carcharodontosaurid and spinosaur tetanurans (e.g., Coria and Salgado, 1995; Sereno et al., 1996; Coria and Currie, 2006). Coelurosaurian remains are relatively sparse, recently new insights provided a better understanding concerning the compositional theropod faunas in Gondwana (e.g., Novas, 1997b; Novas and Puerta, 1997; Kellner, 1999; Makovicky et al., 2005; Novas and Pol, 2005). Among coelurosaurians, oviraptorosaurs are small theropods with a highly specialized skull, grouped in two families

* Corresponding author. E-mail addresses: fedeagnolin@yahoo.com.ar (F.L. Agnolin), agustin_ martinelli@yahoo.com.ar (A.G. Martinelli). 0195-6671/$ - see front matter Ó 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.cretres.2006.10.006

(Oviraptoridae and Caenagnathidae), and a series of basal line´ lska et al., 2004), ages (e.g. Incisivosaurus, Caudipteryx; Osmo being relatively well known in Cretaceous outcrops of North America, Mongolia, and China (e.g., Barsbold et al., 1990; Norell et al., 2001; Dong and Currie, 1996; Clark et al., 2001; Xu et al., 2002). Additionally, oviraptorosaur remains have also been recognized in at least three different localities from Gondwana. The first mention was made by Frey and Martill (1995) based on an incomplete sacrum from the Aptian Santana Formation in the Araripe Basin (Brazil). Later on, an isolated cervical vertebra from the Maastrichtian Lecho Formation in northwestern Argentina was interpreted as belonging to an indeterminated oviraptorosaur (Frankfurt and Chiappe, 1999). Outside South America, oviraptorosaur remains were also recognized in Lower Cretaceous rocks of Victoria, Australia (Dinosaur Cove East Locality; Rich and Rich, 1989) based on a possible incomplete right surangular and a small dorsal vertebra (Currie et al., 1996). Kellner (1999), and Makovicky and Sues (1998) cast doubt on the interpretations made of Brazilian

the neural spine was probably Osmo located in the anterior half of the neural arch instead of being in the middle as in oviroptorosaurs (Makovicky and Sues.. northwestern Argentina. Barsbold and ´ lska. The neural arch is lower than the centrum and is cranially broad. anterior (B). dorsal (C). Bonaparte and Powell. Argentina. The centrum is low. . MUC-PV: Palaeontological Collection of the Museo de Ciencias Naturales de la Universi´ n Province. ´ n Collection) of the Museo ArChubut Collection. The neural spine is not preserved.G. oviraptorids (e. Frankfurt and Chiappe. Anterior cervical vertebra of a possible Noasauridae in lateral (A).g. Lillo.. Buenos Aires Province. Upper Cretaceous. MACN-PV 622: almost complete isolated cervical vertebra. dad Nacional del Comahue.g. and elongate bearing a ventral longitudinal groove as is also present in Elaphrosaurus Fig. Neuque PVL: Palaeontological Collection of the Instituto Miguel ´ n Province. 1998).786 F. Salta Group. Description The vertebra is nearly complete. A. N. 1999). but it seems to be reduced as in abelisauroids (e. Systematic palaeontology Suborder: Theropoda Marsh (1881) Infraorder: Ceratosauria Marsh (1884) Superfamily: Abelisauroidea Bonaparte (1991a) Family: Noasauridae Bonaparte and Powell (1980) Gen et sp. Ligabueino andesi. 1996. MACN-PV 622. Locality and horizon. Microvenator. 1990). Institutional Abbreviations MACN-PV: Vertebrate Palaeontological Collection (CH. Tucuma 2. Martinelli / Cretaceous Research 28 (2007) 785e790 and Australian specimens because the fragmentary condition of the specimens. Neuque gentino de Ciencias Naturales ‘‘Bernardino Rivadavia’’. Fig. lacking only the postzygapophyseal and epipophyseal regions.. The aim of this contribution is to review and discuss the affinities of the isolated vertebra (MACN-PV 622. northwestern Argentina) previously described by Frankfurt and Chiappe (1999).. Lecho Formation.g. and the presence of unambiguous trait are recorded in several theropod families. Fig. Chiappe. ornithomimosaurs. 1977. narrow. Estancia El Brete. Argentina. Makovicky and Sues. Salta Province. and ventral (D) views. Argentina.L. 1993. Referred specimen. Agnolin. 1980. The centrum is as high as broad and about five times long than high. 3. Bonaparte. 1. Maastrichtian (Bonaparte et al. 1998) and many tetanurans (e. Additionally. indet. 1) from the Lecho Formation (Maastrichtian) (Salta Province. 2C). the lack of synapomorphies.

1997). 1997). A. located below the level of the prezigapophyses similar to abelisauroids (e. and therizinosaurids (Britt. In dorsal view. and the centrum is slightly ophistocoelous (contra Frankfurt and Chiappe. B. stout and their articular facets are oval. (1990)) in lateral.. Noasaurus leali (after Bonaparte and Powell (1980)) in lateral (left above). p.F. and in a lesser degree.. 2. 1920). 1998). anterior. A third pair of pneumatic . Coria and Salgado. and dorsal (right below) views. Makovicky. posterior (left below).. The diapophyses are well developed and are highly extended laterally as in Elaphrosaurus. convex dorsoventrally and the posterior articular surface clearly concave. 1998).102) are not preserved. and dorsal views. Agnolin. but there is a shallow ridge that runs from the base of the epipophyses to the base of the diapophyses. 1999 e they interpreted it as platycoelous) with the anterior surface. The epipophyses (contra Frankfurt and Chiappe. 2C).g.g. oviraptorosaurs (Frankfurt and Chiappe. C. (Janensch. In anterior view. 1990. Comparison of cervical vertebrae among: A. 2004). although eroded slightly. Carnotaurus sastrei (after Bonaparte et al. anterior.G. Both cranial and caudal articular surfaces are subquadrangular. 2D). Fig. there are two peduncular foramina on the neural arch. oviraptorosaurs (e. 1990. Fig. 1997. Ligabueino (Bonaparte.. Holtz. The prezygapophyses are small. anterior (right above).. Not to scale. Ligabueino andesi (after Bonaparte (1996)) in dorsal view. Martinelli / Cretaceous Research 28 (2007) 785e790 787 Fig. Salta specimen in lateral. Sues. Bonaparte et al. 1999. and D. 1920). 1996. the space between the prezygapophyses is U-shaped as occurs in Elaphrosaurus (Janensch. 1999. 1993). in the basal ceratosaur Spinostropheus (Sereno et al. and dorsal views. Carnotaurus (Bonaparte et al. Another pair of pleurocoels is present caudodorsally to the diapophyses as is diagnostic of Ceratosauria (Rowe et al.. Spinostropheus and oviraptorosaurs. The diapophyses are connected with the prezygapophyses by a strong ridge.L.

1994. There is also a well developed prespinal depression (i. Discussion and conclusions Frankfurt and Chiappe (1999) nested MACN-PV 622 together with oviraptorids. Elaphrosaurus was traditionally considered as an ornithomimosaur (Barsbold et al. Carnotausus sastrei. 1980).e. Bonaparte et al.1. wider neural canal and articular surfaces of the centrum. 1990. 2004). smaller than the latter.G. 1996. 1997. and Microvenator celer from the Cloverly Formation of Montana (Makovicky and Sues. 1997). Wilson et al.. Coria et al. In that analysis.. Novas. prezygapophyses not directed dorsally and diapophysis almost at the same level that the prezygapophyses. Carrano et al.. notch) as also occurs in neoceratosaurs (i. Chirostenotes. 2D).. 1998).. 1999). Noasaurus leali from the Maastrichtian Lecho Formation of Salta (Argentina) (PVL 4061. Mononykus olecranus from the Maastrichtian Nemegt Formation of Mongolia (e. convergently acquired by oviraptorosaurs). Bonaparte et al. while their feature 8 is unknown in noasaurids (i. Fig. and Masiakasaurus knopfleri from the Maastrichtian Maevarano Formation of Madagascar (Sampson et al. Specimen MACN-PV 622 differs from oviraptorosaurs in having no lateral depression on cervical centrum. 2003) or even within abelisauroids (Holtz.. 2B). Elaphrosaurus bambergi. Bonaparte and Powell. Moreover. Coria and Salgado. a thin and low ridge connecting the postzygapophyseal area with the diapophysis is also present and better developed in Noasaurus leali (Bonaparte and Powell. Additionally. but is present in the neoceratosaurs Elaphrosaurus and Spinostropheus. 2002) the lack of a lateral depression on cervical centrum. Carrano et al. 1991a. 1998)... this character state is present in some abelisauroids (e.e..788 F. MACN-PV 622 shares with Elaphrosaurus (Holtz. Laevisuchus indicus from the Maastrichtian Lameta Formation of India (Novas et al. 1999). cul Formation of Neuque 1998). Chirostenotes pergracilis from the Campanian Dinosaur Park and Campanian-Maastrichtian Horseshoe Canyon formations of Alberta (Canada) (Sues.. 1990). 1990. 1998).. Noasaurus leali and Ligabueino andesi. and zygapophyses displaced laterally (Makovicky. the basal Ilokelesia aguadagrandensis (Coria and Salgado. such as the presence of T-shaped epipophyses and triangular diapophyses in lateral view. . Bonaparte. 1994). Chirostenotes. Also MACN-PV 622 shares with Abelisauroidea the cervical centrum in anterior view more than 20% broader than tall (Holtz. 4.. Ligabueino andesi from the Neocomian La Amarga Forma´ n (Argentina) (MACN-PV-N tion of Argentina of Neuque 42. Carnotaurus sastrei from the Maastrichtian La Colonia Formation of Chubut (Argentina) (MACN-PV-CH 894.. 1996). The clade formed by these five taxa shared the presence of peduncular foramina (Character 1) and of a ventral sulcus flanked by ventrolaterally directed ridges (Character 8) (Frankfurt and Chiappe. 1998). Oviraptorosauria (coelurosauria) MACN-PV 622 was nested together with oviraptorosaurs because having a peduncular foramina. therizinosaurids and an unnamed coelurosaur from the Morrison Formation based on a cladistic analysis of ten cervical characters among nineteen theropod taxa. 2002. Abelisauroidea (Ceratosauria) Specimen MACN-PV 622 shares with Ceratosauria the presence of a double pair of pleurocoels (Rowe et al.. Novas.. facing ventrocranially. strongly opistocoelic centrum. the surrounding area indicates that it was very reduced) (Novas. A. 1997). Bonaparte and Powell. 5. 2C). 1999). Comparisons MACN-PV 622 was compared to the following taxa: Ceratosaurs: Elaphrosaurus bambergi from the Tithonian Tendaguru Formation of Tanzania (Janensch. 1990. 1920). 1998. Character 1 is also reported in abelisauroids such as Carnotaurus sastrei (Bonaparte et al.g. Finally.. the dorsal surface of the neural arches clearly delimited from lateral surface of the diapophyses (Bonaparte. 1992). Finally MACN-PV 622 resembles Noasauridae in the low and craniocaudally elongated neural arch (Coria and Salgado. Perle et al. Bonaparte.L. 1980).. the noasaurids Noasaurus leali (Fig. all these features are also reminiscent of noasaurid abelisauroids. Fig. 1998). Noasaurus and MACN-PV 622 differ from Abelisauridae because they lack several diagnostic characters of this group.. 1996).g. 2001.. La Carpa Formation of Neuque 1991b. It could be assigned to Neoceratosauria on the basis of the following features: neural spine of cervical vertebrae reduced anteroposteriorly (despite the lack of neural spine in this specimen. and the ventral surface of the centrum antero-posteriorly grooved with lateral ridges (Frankfurt and Chiappe. 2004).2. well beyond the top of the neural spine (e. wide vertebral canal. Ceratosaurus and abelisauroids. Agnolin.. Noasaurus leali.. Carrano et al. 1980. 1990.. Bonaparte et al. and the presence of a deep prespinal depression (Novas. 2002). 2001. 1996). Bonaparte. Fig. and possibly the neural spine located in the anterior half of the neural arch. Bonaparte and Powell. Martinelli / Cretaceous Research 28 (2007) 785e790 foramina. Ilokelesia aguadagrandensis from the Albian-Cenomanian Huin´ n (Argentina) (Coria and Salgado. In this regard. Bonaparte. a U-shaped space between both prezygapophyses. wider U-shaped space between the prezygapophyses. is placed at the base of the diapophyses. 1994) and Masiakasaurus (Sampson et al. oviraptorids and the specimen MACN-PV 622 shared the presence of a U-shaped space between prezygapophyses in dorsal view (Character 10) (Frankfurt and Chiappe. 2002).g. 1980. 1992). Coelurosaurs: Alvarezsaurus calvoi from the Coniacian Bajo ´ n (MUC-PV 54.g. 2) and Laevisuchus indicus (Novas et al. 1990) but recent phylogenetic analyses nested this taxon inside the neoceratosaur clade (e. therizinosaurids. 1998. 4.e. prezygapophyses not directed dorsally and the U-shaped space between the prezygapophyses. Holtz. Bonaparte et al. and dorsally directed epipophyses. 4.. 1992).

Ornithomimosauria. Coria. Also the resemblance is noted on the proportions of the centrum. F. Therefore. H.F.A. 1977.M. Bonaparte.F. Chiappe... Based on the resemblances with Elaphrosaurus... L. Osmo ´ lska. University of Calgary. J.. 17e123. (Eds. Aves). Carnotaurus. In: Weishampel. 1998. Barsbold.). 1990). Masiakasaurus.A. UniWeishampel. Coria... H. Los vertebrados fo ´ n y cercanı ´as.J. Two new oviraptorids (Theropoda: Oviraptorosauria) Late Cretaceous Djadoktha Formation. 1980).. A new carcharodontosaurid (Dinosauria.D. and both Noasaurus (Bonaparte and Powell. Bonaparte. A new giant carnivorous dinosaur from the Cretaceous of Patagonia. There is a great morphological difference between the anterior cervical vertebrae of Elaphrosaurus and MACN-PV 622 and those of the noasaurid Masiakasaurus (Carrano et al. Thesis. 249e258. P. This feature is also observed in abelisauroids (e. R. 224e226. J. we reject the occurrence of oviraptorosaurs in Argentina and South America (Kellner. Powell.A. 2001. R. C. H. 375 pp. 1993.. Chiarelli for their valuable comments. Chiappe. 1e25.. The gondwanan theropod families Abelisauridae and Noasauridae.. Argentina. Nature 377. 225e244.. P. R.E. Argentina.. In: Barsbold. and the supposed anterior vertebrae of Masiakasaurus. Pneumatic postcranial bones in dinosaurs and other archosaurs. Makovicky and Sues. Historical Biology 5. 1980) and Elaphrosaurus (Janensch. S. We thank two anonymous reviewers for their enlightening comments.. 460e465.T.A. nearly straight plane made by the spike-like pronounced epipophyses. plus the several convergences with oviraptorosaur vertebrae (e.F.L. 89e102. Acknowledgments We wish to thank F. Theropoda) from the Upper Cretaceous of Argentina. described by Bonaparte and Powell (1980) may be the seventh or the eighth. 1e27. northwestern Argentina (Sau´ moires Societe ropoda. R. J..F.. 71e118. 19e28. 1996. in contrast thomimosaurs by Barsbold and Osmo with Ceratosaurus and specially Abelisauridae which were short and robust necked theropods (Novas. R. 209e213.. imo al lı Britt.G. L. 1995.A. P. Australia. University of California Dobson. 1990.g.. Vickers-Rich. 1980. 1996. The Dinosauria.. the horned.. B.. and specimen MACN-PV 622 shared the presence of the caudal border of the diapophyses projected laterally at the midpoint.. Mongolia. Maryanska. Bonaparte.. Gaia 15. G. Carrano. Salgado. 73e130.. Osmo ´ lska.. a small abelisauroid (Dinosauria: Theropoda) from the Late Cretaceous of Madagascar. 1999. pp.F. American Museum Novitates 3083. such as general proportions. Me Geologie France 139. Kramarz for allowing us to study the collections under his care. Oviraptoridae. Thus.. We also thank A. Carnosauria. A. R. J. G. Carnosauria. 1990).. Coria. Journal of Vertebrate Paleontology 22.A.. 1999). 1990. whereas the neural arch of Noasaurus leali. Creta ´ cico Superior. Laevisuchus and Noasaurus are probably placed in a more posterior position than previously suggested (Carrano et al. among others) and Spinostropheus (Sereno et al. Currie. 1992. Journal of Vertebrate Paleontology 22. Bonaparte. J. D.. placement of neural spine and zygapophyses and the deep excavation anterior to the postzygapophyses... Martinelli / Cretaceous Research 28 (2007) 785e790 789 the clade integrated by Chirostenotes. R. Clark. Berkeley and Los Angeles. 1998). (Ed.J.H. J.B. Because the MACN-PV 622 was found associated with the holotype of Noasaurus leali (Frankfurt and Chiappe. Agnolin. 2002). T. 19e28. A basal Abelisauria Novas. a striking similarity is noted with the last cervical of Elaphrosaurus. 73e79. P. Mu ¨ nchner Geowissenschaftliche Abhandlungen (A) 30. 2006. 510e534. 2004).. 2002). L. pp. 1993.. L. Novas. As seen in the fossil record. R. Bonaparte. F. Barsbold.. Coelurosauria. Osmo Press. are very similar to those of typical noasaurids.. A continental assemblage of tetrapods from the Upper Cretaceous beds of El Brete. 1990. 2002. Currie. J. Rich. The Dinosauria. Cretaceous tetrapods of Argentina. Salfitty. Elaphrosaurus. Enantiornithine (Aves) Tarsometatarsi from the Cretaceous Lecho Formation of Northwestern Argentina.g... Contribution in Science.. Oviraptorosauria. lightly built carnosaur from the Middle Cretaceous of Patagonia. M. Sampson. J.. Powell. our assignment of MACN-PV 622 to the noasaurid family indicates the presence of several convergences acquired in abelisauroids and oviraptorosaurians. M. MACN-PV 622 shares many features with abelisauroids. Forster. pro ´ xsaurios y aves creta ´mite con Tucuma ´ n. and specially noasaurid instead of oviraptorosaur affinities. Acta Geolo ´ gica Lilloana 14. 2002.E. Possible oviraptorosaur (Theropoda. pp. P. J. specimen MACN-PV 622 probably belongs to the third or fourth cervical vertebra (was suggested as fourth or fifth by Frankfurt and Chiappe.. 1992 (TheropodaCeratosauria) from the Cretaceous of Patagonia. H. and the material resembles both in morphology (e.F..D. J. (Eds. Bossi. U-shaped space between the prezygapophyses) suggest that probably some noasaurids were long necked theropods resembling the ornithomimid coelurosaurs (Elaphrosaurus was originally compared with orni´ lska. 1999). The elongated anterior cervical vertebra of MACN-PV 622. and P. Berkeley and Los Angeles. until there are any new findings. T. 1999). Coria.. Salgado.B. Geodiversitas 28. In our view.). Noasaurus. Unpublished Ph. Natural History Museum Los Angeles Country 416. . D.. Dingus. Dinosauria) specimens from the Early Cretaceous Otway Group of Dinosaur Cove. Journal of Vertebrate Paleontology 21. Comparing the vertebra described as anterior cervical by Carrano et al. ´ lska. A new close relative of Carnotaurus sastrei Bonaparte 1985 (Theropoda: Abelisauridae) from the Late Cretaceous of Patagonia.. ´ siles de la Formacio ´ n Rı ´o Colorado.. if we follow this interpretation. (2002) with those of Elaphrosaurus. Rede la Ciudad de Neuque vista del Museo Argentino de Ciencias Naturales 4. Novas.E.. the features considered to include MACN-PV 622 within Oviraptorosauria are widely documented in several theropod species reflecting that these do not necessarily implicate a direct phylogenetic relationship. the similar position of peduncular foramina) and size we suggest that MACN-PV 622 belongs to an anteriormost cervical vertebra of Noasaurus leali (Bonaparte and Powell. Contribution of Sourthern South America to Vertebrate Paleontology.g. 1e42.). because as in Elaphrosaurus it shows in lateral view a dorsal. Osmo versity of California Press. nearly forming a right angle with the centrum in dorsal view (Character 9) (Frankfurt and Chiappe.. 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