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

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

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

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

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

Carrano et al.g..g. 1998). Finally MACN-PV 622 resembles Noasauridae in the low and craniocaudally elongated neural arch (Coria and Salgado. MACN-PV 622 shares with Elaphrosaurus (Holtz. A.. Laevisuchus indicus from the Maastrichtian Lameta Formation of India (Novas et al. 1998. 1996. 1998.. 1991a. prezygapophyses not directed dorsally and diapophysis almost at the same level that the prezygapophyses. Fig. smaller than the latter. the noasaurids Noasaurus leali (Fig. Discussion and conclusions Frankfurt and Chiappe (1999) nested MACN-PV 622 together with oviraptorids. Ceratosaurus and abelisauroids. Novas. wider U-shaped space between the prezygapophyses. the basal Ilokelesia aguadagrandensis (Coria and Salgado. 1990.. the dorsal surface of the neural arches clearly delimited from lateral surface of the diapophyses (Bonaparte. convergently acquired by oviraptorosaurs). Martinelli / Cretaceous Research 28 (2007) 785e790 foramina.. 2002).... La Carpa Formation of Neuque 1991b.. 2C)... while their feature 8 is unknown in noasaurids (i.e.. 1980). 4. Ligabueino andesi from the Neocomian La Amarga Forma´ n (Argentina) (MACN-PV-N tion of Argentina of Neuque 42.g. the surrounding area indicates that it was very reduced) (Novas. Specimen MACN-PV 622 differs from oviraptorosaurs in having no lateral depression on cervical centrum. 2002. 2B). facing ventrocranially. 1996). 1998). 2001. 2003) or even within abelisauroids (Holtz. 1998). and dorsally directed epipophyses. wider neural canal and articular surfaces of the centrum. 2002) the lack of a lateral depression on cervical centrum. 2004). 1990). Bonaparte. Bonaparte and Powell. Noasaurus and MACN-PV 622 differ from Abelisauridae because they lack several diagnostic characters of this group.. Coria and Salgado. and the ventral surface of the centrum antero-posteriorly grooved with lateral ridges (Frankfurt and Chiappe. Carnotausus sastrei. 1990. and zygapophyses displaced laterally (Makovicky. Fig. Bonaparte et al. 1997). 1999). and Microvenator celer from the Cloverly Formation of Montana (Makovicky and Sues. this character state is present in some abelisauroids (e. Wilson et al. and possibly the neural spine located in the anterior half of the neural arch. In this regard. Moreover.. 1996). Bonaparte.. Character 1 is also reported in abelisauroids such as Carnotaurus sastrei (Bonaparte et al. Noasaurus leali and Ligabueino andesi.L. Ilokelesia aguadagrandensis from the Albian-Cenomanian Huin´ n (Argentina) (Coria and Salgado. Carrano et al. 1998). 1994. 2002). 1996).e. Finally. Chirostenotes pergracilis from the Campanian Dinosaur Park and Campanian-Maastrichtian Horseshoe Canyon formations of Alberta (Canada) (Sues. 1990. Holtz.788 F. Fig.. Comparisons MACN-PV 622 was compared to the following taxa: Ceratosaurs: Elaphrosaurus bambergi from the Tithonian Tendaguru Formation of Tanzania (Janensch. Coelurosaurs: Alvarezsaurus calvoi from the Coniacian Bajo ´ n (MUC-PV 54. cul Formation of Neuque 1998). 5. 1980. Noasaurus leali from the Maastrichtian Lecho Formation of Salta (Argentina) (PVL 4061. Abelisauroidea (Ceratosauria) Specimen MACN-PV 622 shares with Ceratosauria the presence of a double pair of pleurocoels (Rowe et al. Bonaparte and Powell. 1990. Also MACN-PV 622 shares with Abelisauroidea the cervical centrum in anterior view more than 20% broader than tall (Holtz. 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. There is also a well developed prespinal depression (i. . and the presence of a deep prespinal depression (Novas. 2001.1. Coria et al. 1992). Oviraptorosauria (coelurosauria) MACN-PV 622 was nested together with oviraptorosaurs because having a peduncular foramina.. Bonaparte et al. Elaphrosaurus bambergi. 1994). Carrano 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. 4. 2004). 1997. Novas. Elaphrosaurus was traditionally considered as an ornithomimosaur (Barsbold et al.. 1992). wide vertebral canal. notch) as also occurs in neoceratosaurs (i. Mononykus olecranus from the Maastrichtian Nemegt Formation of Mongolia (e.e. therizinosaurids and an unnamed coelurosaur from the Morrison Formation based on a cladistic analysis of ten cervical characters among nineteen theropod taxa. all these features are also reminiscent of noasaurid abelisauroids. Carnotaurus sastrei from the Maastrichtian La Colonia Formation of Chubut (Argentina) (MACN-PV-CH 894.. Bonaparte et al. 1994) and Masiakasaurus (Sampson et al. 1999). well beyond the top of the neural spine (e. Bonaparte and Powell. a U-shaped space between both prezygapophyses.g. Agnolin. In that analysis. therizinosaurids. Bonaparte. prezygapophyses not directed dorsally and the U-shaped space between the prezygapophyses. Additionally. 1980). 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. such as the presence of T-shaped epipophyses and triangular diapophyses in lateral view.2. Chirostenotes. a thin and low ridge connecting the postzygapophyseal area with the diapophysis is also present and better developed in Noasaurus leali (Bonaparte and Powell. 1990) but recent phylogenetic analyses nested this taxon inside the neoceratosaur clade (e.. 1998). Noasaurus leali. Chirostenotes. 1992). but is present in the neoceratosaurs Elaphrosaurus and Spinostropheus. 2D).. strongly opistocoelic centrum. Bonaparte et al. 1920). 1997). and Masiakasaurus knopfleri from the Maastrichtian Maevarano Formation of Madagascar (Sampson et al. Bonaparte. 1980. 4.. is placed at the base of the diapophyses. Perle et al. 1999).G.. 2) and Laevisuchus indicus (Novas et al.

pp. Elaphrosaurus. pp. Ukhaa Tolgod. The Dinosauria.E.. J. 1e27. L. described by Bonaparte and Powell (1980) may be the seventh or the eighth. UniWeishampel.. Thus. 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. R. J. R.L. Mu ¨ nchner Geowissenschaftliche Abhandlungen (A) 30. In our view. a striking similarity is noted with the last cervical of Elaphrosaurus. Dobson.F. Oviraptorosauria. Bonaparte et al. The elongated anterior cervical vertebra of MACN-PV 622. specimen MACN-PV 622 probably belongs to the third or fourth cervical vertebra (was suggested as fourth or fifth by Frankfurt and Chiappe. 1990). 1992 (TheropodaCeratosauria) from the Cretaceous of Patagonia.. Currie. among others) and Spinostropheus (Sereno et al. Salgado. In: Arratia. Hallazgos de dino´ cicas en la Formacio ´ n Lecho de El Brete (Salta). nearly straight plane made by the spike-like pronounced epipophyses.. Contribution in Science.. 1999). A continental assemblage of tetrapods from the Upper Cretaceous beds of El Brete.. Carnotaurus sastrei Bonaparte. 1998). and the supposed anterior vertebrae of Masiakasaurus.A. M. P. 249e258. Dinosauria) specimens from the Early Cretaceous Otway Group of Dinosaur Cove. ´ siles de la Formacio ´ n Rı ´o Colorado. the horned.). Bossi.. Novas. Comparing the vertebra described as anterior cervical by Carrano et al. (Eds.. Osmo ´ lska. Based on the resemblances with Elaphrosaurus. Me Geologie France 139. such as general proportions. Berkeley and Los Angeles. Barsbold. Bonaparte. Makovicky and Sues. 224e226.. Los vertebrados fo ´ n y cercanı ´as... L.. 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. Laevisuchus and Noasaurus are probably placed in a more posterior position than previously suggested (Carrano et al. 225e244. A new close relative of Carnotaurus sastrei Bonaparte 1985 (Theropoda: Abelisauridae) from the Late Cretaceous of Patagonia. MACN-PV 622 shares many features with abelisauroids.. Coria. Australia. Osmo versity of California Press. Contribution of Sourthern South America to Vertebrate Paleontology. Journal of Vertebrate Paleontology 21. Journal of Vertebrate Paleontology 22. 1990..F. 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. Rede la Ciudad de Neuque vista del Museo Argentino de Ciencias Naturales 4. our assignment of MACN-PV 622 to the noasaurid family indicates the presence of several convergences acquired in abelisauroids and oviraptorosaurians. P. 510e534. 1990)...).. 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