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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: email@example.com (F.L. Agnolin), agustin_ firstname.lastname@example.org (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
Maastrichtian (Bonaparte et al.. narrow. The centrum is low. Microvenator. The neural spine is not preserved. Frankfurt and Chiappe. lacking only the postzygapophyseal and epipophyseal regions. Makovicky and Sues. Chiappe. Additionally. Neuque PVL: Palaeontological Collection of the Instituto Miguel ´ n Province. anterior (B). 1990). MACN-PV 622.786 F. Systematic palaeontology Suborder: Theropoda Marsh (1881) Infraorder: Ceratosauria Marsh (1884) Superfamily: Abelisauroidea Bonaparte (1991a) Family: Noasauridae Bonaparte and Powell (1980) Gen et sp. northwestern Argentina) previously described by Frankfurt and Chiappe (1999). Argentina.g. Description The vertebra is nearly complete. 1998) and many tetanurans (e. and the presence of unambiguous trait are recorded in several theropod families. Ligabueino andesi. Martinelli / Cretaceous Research 28 (2007) 785e790 and Australian specimens because the fragmentary condition of the specimens. Locality and horizon. 1998). 1999). northwestern Argentina.g.g. and ventral (D) views. The aim of this contribution is to review and discuss the afﬁnities of the isolated vertebra (MACN-PV 622. 1) from the Lecho Formation (Maastrichtian) (Salta Province. The centrum is as high as broad and about ﬁve times long than high. Tucuma 2. Institutional Abbreviations MACN-PV: Vertebrate Palaeontological Collection (CH. Agnolin. Lecho Formation. Salta Group. 1993.. ´ n Collection) of the Museo ArChubut Collection. 1980. Lillo. MACN-PV 622: almost complete isolated cervical vertebra. Neuque gentino de Ciencias Naturales ‘‘Bernardino Rivadavia’’. 1.L. Fig. Salta 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. Upper Cretaceous.. dorsal (C). Buenos Aires Province. . indet. Bonaparte.G. 2C). dad Nacional del Comahue. but it seems to be reduced as in abelisauroids (e. Fig.. The neural arch is lower than the centrum and is cranially broad. A. MUC-PV: Palaeontological Collection of the Museo de Ciencias Naturales de la Universi´ n Province. oviraptorids (e. 3. the lack of synapomorphies. Bonaparte and Powell. 1977. ornithomimosaurs. Anterior cervical vertebra of a possible Noasauridae in lateral (A). Referred specimen. Argentina. and elongate bearing a ventral longitudinal groove as is also present in Elaphrosaurus Fig. N. Estancia El Brete. 1996. Argentina. Barsbold and ´ lska.
posterior (left below). p. and therizinosaurids (Britt. and dorsal (right below) views. 1999.102) are not preserved. A. there are two peduncular foramina on the neural arch. 2C). 2D). The diapophyses are well developed and are highly extended laterally as in Elaphrosaurus. In dorsal view. 1997). and D. 1999. oviraptorosaurs (e. but there is a shallow ridge that runs from the base of the epipophyses to the base of the diapophyses. (1990)) in lateral. Sues.. 1996. A third pair of pneumatic . anterior. 1999 e they interpreted it as platycoelous) with the anterior surface. 1997. In anterior view. B. and the centrum is slightly ophistocoelous (contra Frankfurt and Chiappe.g... 1990. and in a lesser degree. anterior (right above). Fig. Spinostropheus and oviraptorosaurs. Not to scale.L. C. located below the level of the prezigapophyses similar to abelisauroids (e. Another pair of pleurocoels is present caudodorsally to the diapophyses as is diagnostic of Ceratosauria (Rowe et al. 1998). the space between the prezygapophyses is U-shaped as occurs in Elaphrosaurus (Janensch. 1997). Agnolin. oviraptorosaurs (Frankfurt and Chiappe. Salta specimen in lateral.G. anterior.F. 1990. The diapophyses are connected with the prezygapophyses by a strong ridge. convex dorsoventrally and the posterior articular surface clearly concave. although eroded slightly. The epipophyses (contra Frankfurt and Chiappe. Noasaurus leali (after Bonaparte and Powell (1980)) in lateral (left above). Ligabueino (Bonaparte. 1993). Both cranial and caudal articular surfaces are subquadrangular. Fig. (Janensch. and dorsal views. Bonaparte et al... 1998). 1920). Makovicky. Holtz. Carnotaurus sastrei (after Bonaparte et al. Comparison of cervical vertebrae among: A. Carnotaurus (Bonaparte et al. stout and their articular facets are oval. Martinelli / Cretaceous Research 28 (2007) 785e790 787 Fig. Ligabueino andesi (after Bonaparte (1996)) in dorsal view. and dorsal views. The prezygapophyses are small. 2004). in the basal ceratosaur Spinostropheus (Sereno et al. Coria and Salgado.. 2.g. 1920).
the noasaurids Noasaurus leali (Fig. 1999).. 1980. 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. Chirostenotes pergracilis from the Campanian Dinosaur Park and Campanian-Maastrichtian Horseshoe Canyon formations of Alberta (Canada) (Sues. and Masiakasaurus knopﬂeri from the Maastrichtian Maevarano Formation of Madagascar (Sampson et al.. 2002)... 1998).. Bonaparte and Powell. 1998). In that analysis. In this regard. smaller than the latter. 4. a U-shaped space between both prezygapophyses. Agnolin. well beyond the top of the neural spine (e. Carnotausus sastrei. Elaphrosaurus was traditionally considered as an ornithomimosaur (Barsbold et al. 2D). all these features are also reminiscent of noasaurid abelisauroids. Elaphrosaurus bambergi... 2003) or even within abelisauroids (Holtz. Moreover. 1990) but recent phylogenetic analyses nested this taxon inside the neoceratosaur clade (e. wide vertebral canal. 1992). Coelurosaurs: Alvarezsaurus calvoi from the Coniacian Bajo ´ n (MUC-PV 54. 1990. Oviraptorosauria (coelurosauria) MACN-PV 622 was nested together with oviraptorosaurs because having a peduncular foramina. 1990). Noasaurus leali and Ligabueino andesi. Carrano et al.g. therizinosaurids and an unnamed coelurosaur from the Morrison Formation based on a cladistic analysis of ten cervical characters among nineteen theropod taxa. Novas. wider U-shaped space between the prezygapophyses. Bonaparte. Mononykus olecranus from the Maastrichtian Nemegt Formation of Mongolia (e. strongly opistocoelic centrum. Ligabueino andesi from the Neocomian La Amarga Forma´ n (Argentina) (MACN-PV-N tion of Argentina of Neuque 42. Bonaparte et al. 1997).. 1980). Bonaparte. and the presence of a deep prespinal depression (Novas. Coria et al. this character state is present in some abelisauroids (e. Bonaparte. wider neural canal and articular surfaces of the centrum. 1990. Also MACN-PV 622 shares with Abelisauroidea the cervical centrum in anterior view more than 20% broader than tall (Holtz. and Microvenator celer from the Cloverly Formation of Montana (Makovicky and Sues. There is also a well developed prespinal depression (i. 1996). Carnotaurus sastrei from the Maastrichtian La Colonia Formation of Chubut (Argentina) (MACN-PV-CH 894.1. 1996). prezygapophyses not directed dorsally and the U-shaped space between the prezygapophyses. Carrano et al. 1999). Finally MACN-PV 622 resembles Noasauridae in the low and craniocaudally elongated neural arch (Coria and Salgado. 1999). Bonaparte. 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...e. . prezygapophyses not directed dorsally and diapophysis almost at the same level that the prezygapophyses. 1997. Bonaparte et al. 4. Bonaparte et al. Wilson et al. but is present in the neoceratosaurs Elaphrosaurus and Spinostropheus.e. 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. La Carpa Formation of Neuque 1991b. Novas. 2004). and the ventral surface of the centrum antero-posteriorly grooved with lateral ridges (Frankfurt and Chiappe. Bonaparte and Powell. Ilokelesia aguadagrandensis from the Albian-Cenomanian Huin´ n (Argentina) (Coria and Salgado. Fig.. 2002) the lack of a lateral depression on cervical centrum... cul Formation of Neuque 1998). 1998. the basal Ilokelesia aguadagrandensis (Coria and Salgado..e. MACN-PV 622 shares with Elaphrosaurus (Holtz. 1980. Bonaparte et al.G. Fig. a thin and low ridge connecting the postzygapophyseal area with the diapophysis is also present and better developed in Noasaurus leali (Bonaparte and Powell. Character 1 is also reported in abelisauroids such as Carnotaurus sastrei (Bonaparte et al. Additionally. 5. Ceratosaurus and abelisauroids. 2) and Laevisuchus indicus (Novas et al. 1920). convergently acquired by oviraptorosaurs). 1998. 1998).. notch) as also occurs in neoceratosaurs (i..g.. is placed at the base of the diapophyses.. 1992). Holtz.. A. Discussion and conclusions Frankfurt and Chiappe (1999) nested MACN-PV 622 together with oviraptorids. Laevisuchus indicus from the Maastrichtian Lameta Formation of India (Novas et al.g. the surrounding area indicates that it was very reduced) (Novas. 1992). 1990. 1990. 1994. 1994). Perle et al. Fig. 2B). 1996. 2002).. Chirostenotes. 1994) and Masiakasaurus (Sampson et al. 1998).g. Carrano et al. therizinosaurids. 1998). Martinelli / Cretaceous Research 28 (2007) 785e790 foramina. Noasaurus and MACN-PV 622 differ from Abelisauridae because they lack several diagnostic characters of this group. 2004). the dorsal surface of the neural arches clearly delimited from lateral surface of the diapophyses (Bonaparte.. Chirostenotes. facing ventrocranially. such as the presence of T-shaped epipophyses and triangular diapophyses in lateral view. and zygapophyses displaced laterally (Makovicky..L. Noasaurus leali from the Maastrichtian Lecho Formation of Salta (Argentina) (PVL 4061. 1991a.788 F. 2001. Specimen MACN-PV 622 differs from oviraptorosaurs in having no lateral depression on cervical centrum. 2C). 4. 2002. 1997). 1980). Coria and Salgado. and possibly the neural spine located in the anterior half of the neural arch. and dorsally directed epipophyses. Finally. Bonaparte and Powell. Comparisons MACN-PV 622 was compared to the following taxa: Ceratosaurs: Elaphrosaurus bambergi from the Tithonian Tendaguru Formation of Tanzania (Janensch. 1996). 2001.2. Noasaurus leali. Abelisauroidea (Ceratosauria) Specimen MACN-PV 622 shares with Ceratosauria the presence of a double pair of pleurocoels (Rowe et al. while their feature 8 is unknown in noasaurids (i..
Clark. Historical Biology 5. Thus. 2001. Currie. Kramarz for allowing us to study the collections under his care. lightly built carnosaur from the Middle Cretaceous of Patagonia.g. Bossi. F.H. We thank two anonymous reviewers for their enlightening comments. In: Weishampel.J. Novas. Dingus. and the supposed anterior vertebrae of Masiakasaurus. 510e534.. R.. 1996. Powell. 1920). Oviraptorosauria. Martinelli / Cretaceous Research 28 (2007) 785e790 789 the clade integrated by Chirostenotes. J.g. Acknowledgments We wish to thank F.. J...F. Coria. R. and P. such as general proportions. 17e123. 1e25.. 2006. ´ siles de la Formacio ´ n Rı ´o Colorado. J. A new giant carnivorous dinosaur from the Cretaceous of Patagonia. (Eds. 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. P.. Osmo Press. 1999). A basal Abelisauria Novas. Coria. Ukhaa Tolgod.. F. 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