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Osteology of Viavenator Exxoni (Abelisauridae; Furileusauria) from the Bajo De La Carpa Formation, NW Patagonia, Argentina

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Osteology of Viavenator Exxoni (Abelisauridae; Furileusauria) from the Bajo De La Carpa Formation, NW Patagonia, Argentina Accepted Manuscript Osteology of Viavenator exxoni (Abelisauridae; Furileusauria) from the Bajo de la Carpa Formation, NW Patagonia, Argentina Leonardo S. Filippi, Ariel H. Méndez, Federico A. Gianechini, Rubén D. Juárez Valieri, Alberto C. Garrido PII: S0195-6671(17)30147-7 DOI: 10.1016/j.cretres.2017.07.019 Reference: YCRES 3669 To appear in: Cretaceous Research Received Date: 27 March 2017 Revised Date: 4 July 2017 Accepted Date: 29 July 2017 Please cite this article as: Filippi, L.S., Méndez, A.H., Gianechini, F.A., Juárez Valieri, Rubé.D., Garrido, A.C., Osteology of Viavenator exxoni (Abelisauridae; Furileusauria) from the Bajo de la Carpa Formation, NW Patagonia, Argentina, Cretaceous Research (2017), doi: 10.1016/j.cretres.2017.07.019. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. ACCEPTED MANUSCRIPT 1 Osteology of Viavenator exxoni (Abelisauridae; Furileusauria) from the Bajo de 2 la Carpa Formation, NW Patagonia, Argentina. 3 4 Leonardo S. Filippi 1, Ariel H. Méndez 2, Federico A. Gianechini 3, Rubén D. Juárez 5 Valieri 4 and Alberto C. Garrido 5 6 1 7 Museo Municipal Argentino Urquiza . Jujuy y Chaco s/n, 8319, Rincón de los Sauces, 8 Neuquén, Argentina. [email protected] 2 9 CONICET. Instituto de Investigación en Paleobiología y Geología, Isidro Lobo y Belgrano, 10 8332, General Roca, Río Negro, Argentina. [email protected] 3 11 CONICET. Área de Zoología, Departamento de Bioquímica y Ciencias Biológicas, 12 Facultad de Química, Bioquímica y Farmacia, UniversMANUSCRIPTidad Nacional de San Luis, 13 Chacabuco 917, 5700, San Luis, Argentina. [email protected] 4 14 Secretaría de Cultura de la Provincia de Río Negro. Perito Moreno 70, 8324, Cipolletti, Río 15 Negro, Argentina. [email protected] 5 16 Museo Provincial de Ciencias Naturales Prof. Dr. Juan Olsacher , Dirección Provincial de 17 Minería. Etcheluz y Ejército Argentino, 8340, Zapala, Neuquén, Argentina. 18 [email protected] 19 ACCEPTED 20 Abstract 21 The complete osteology of the abelisaurid Viavenator exxoni, from the Bajo de la Carpa 22 Formation (Santonian, Upper Cretaceous) is described. Viavenator is characterized by a 23 series of autapomorphies, including: transversely compressed parietal depressions on both ACCEPTED MANUSCRIPT 24 sides of the supraoccipital crest; ventral edges of the paraoccipital processes located above 25 the level of the dorsal edge of the occipital condyle; basioccipital-opisthotic complex about 26 two and a half times the width and almost twice the height of the occipital condyle, in 27 posterior view; well-developed crest below the occipital condyle, diverging ventrally and 28 defining the subcondylar recess; deeply excavated and sub-circular basisphenoidal recess, 29 with its major axis transversely oriented; basipterygoid processes horizontally placed with 30 respect to the cranial roof and located slightly dorsally to the basal tubera; mid and posterior 31 cervical centra with slightly convex lateral and ventral surfaces; hyposphene-hypanthrum 32 articular complex present from dorsal 2 onward; presence of an interspinous accessory 33 articular system in middle and posterior dorsal vertebrae; presence of a pair of pneumatic 34 foramina within the prespinal fossa in anterior caudal vertebrae; distal end of the scapular 35 blade posteriorly curved. Particularly, Viavenator shows plesiomorphic cranial characters, 36 i.e. flat frontals lacking domes or horns, combined with derived postcranial characters, e.g. 37 the interspinous accessory joint system of dorsal vertebrae.This combination between 38 plesiomorphic and derived traits suggests that ViavenatorMANUSCRIPT is a transitional form, which is an 39 idea supported by its intermediate stratigraphic and phylogenetic plscement between the 40 basal and older (e.g. Skorpiovenator, Ilokelesia ) and derived and younger members of the 41 clade (e.g. Aucasaurus, Carnotaurus ). 42 Keywords: Abelisauridae, Furileusauria, Bajo de la Carpa Formation, Late Cretaceous, 43 Neuquén Basin. 44 1. Introduction 45 Abelisauridae represents the most abundant and best known group of Cretaceous theropod 46 dinosaurs from ACCEPTEDGondwana, and apparently also Europe. Several taxa have been definitely 47 referred to Abelisauridae since its recognition as a monophyletic, mainly in South America 48 (Bonaparte and Novas, 1985; Bonaparte, 1985; Coria and Salgado1998; Coria, 2001; Coria 49 et al., 2002; Kellner and Campos, 2002; Calvo et al., 2004; Canale et al., 2009), Africa 50 (Depéret, 1896; Sereno et al., 2004; Sereno and Brusatte, 2008), the Indian subcontinent 51 (Wilson et al., 2003; Novas et al., 2010,) and Europe (Le Loeuff and Buffetaut, 1991; ACCEPTED MANUSCRIPT 52 Tortosa et al., 2013). The taxonomic diversity of Abelisauridae notably increases the 53 possibilities to interpret and discuss intragroup phylogenetic relationships. This lead to the 54 recognition of new sublineages, such as the “western Gondwanan” South American 55 brachyrostrans, and the “eastern Gondwanan” majungasaurines, which certainly can be 56 considered as a result of both vicariant and dispersal events (Canale et al., 2009; Novas et 57 al., 2013; Tortosa et al., 2013). A new abelisaurid, Viavenator exxoni , was described recently 58 based on a incomplete, partially articulated skeleton comprising the posterior area of the 59 skull, most of the axial sequence and some appendicular elements (Filippi et al., 2016). This 60 abelisaurid comes from the Santonian deposits of the Bajo de la Carpa Formation, Upper 61 Cretaceous, Río Colorado Subgroup, Neuquén Group (Filippi et al., 2016) and it exhibits 62 characters suggesting that it represents a transitional form between the “middle” Cretaceous 63 basal brachyrostrans and the highly-derived latest Cretaceous forms. Viavenator has 64 primitive cranial characters combined with some derived features of the postcranial 65 skeleton. This condition thas enabled the polarization between cranial and postcranial 66 transitional elements, which lead to a more preciseMANUSCRIPT phylogenetic placement of others taxa, 67 as for example Abelisaurus comahuensis. 68 In the present work we provide a detailed and comparative anatomical description of the 69 skeletal elements of Viavenator exxoni . The information supplied is essential for a better 70 comprehension of the anatomical features of this taxon and abelisaurids as a whole, and will 71 be a valuable data source for future studies about this clade of theropod dinosaurs. 72 73 2. Materials and methods 74 2.1. AnatomicalACCEPTED abbreviations 75 a, acromion; ape, anterior projection of epipophysis; aptp , anterior projection of 76 transverse process; ati, atlantal intercentrum; bit, biceps tubercle; Bo, basioccipital; Bs , 77 basisphenoid; bpt, basipterygoid process; bt , basal tuber; caaf, caudal articular facet of 78 chevron; cdl, centrodiapophyseal lamina; cf, coracoid foramen; craf, cranial articular facet of ACCEPTED MANUSCRIPT 79 chevron; crpr, cranial process of chevron; cup, cultriform process; dex, distal expansion of 80 chevron; dp, diapophysis; Eo; exoccipital; ep , epipophyses; F, frontal; gc, glenoid cavity; hc , 81 haemal canal; hp, hyposphene; hy, hypantrum; iaas, interspinous accessory articulation 82 system; ipof , infrapostzygapophyseal fossa; lpf, laminopeduncular foramen; n, 83 neurapophyses; na, neural arch; nc, neural canal; nct, nuchal crest; ns, neural spine; oa , 84 odontoid articulation; oc, occipital condyle; oca, occipital condyle articulation; Op, 85 opisthotic; Os, orbitosphenoid; P, parietal; pcdl, posterior centrodiapophyseal lamina; pcr, 86 paracondylar recess; pe, parietal eminence; pf, pneumatic foramen; Po , postorbital; po, 87 postzygapophysis; pop, paraoccipital process; posf, postspinal fossa; Pro, prootic; pp, 88 parapophysis; pr , prezygapophysis; prel, prezygoepipopiphyseal lamina; pvp, posteroventral 89 process; rbs, basisphenoid recess; rtl, lateral tympanic recess ; sag, sagittal bar; scr, 90 subcondilar recess; spgf ; supraglenoid fossa; Sph, sphenethmoid; Sq , squamosal; So, 91 supraoccipital; sot, supraoccipital tuberosity; stf, supratemporal fenestra; stfo , supratemporal 92 fossa; ssr, subselar recess; tbs, scar for triceps MANUSCRIPTbrachii scapularis ; tp, transverse process; 93 vcd, foramen vena capitis dorsalis; vp, ventral processes. 94 95 2.2. Institutional abbreviations 96 MAU-Pv-LI, Museo Municipal “Argentino Urquiza” , Rincón de los Sauces, Neuquén 97 province, Argentina; MUCPv, Museo de la Universidad del Comahue, Neuquén province, 98 Argentina; MACN, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos 99 Aires; Argentina; MCF, Museo “Carmen Funes”, Plaza Huincul, Neuquén province, 100 Argentina; MPM ACCEPTED, Museo Padre Molina, Río Gallegos, Santa Cruz, Argentina; MPCA , Museo 101 Carlos Ameghino, Cipolletti, Río Negro province, Argentina; FMNH, Field Museum of Natural 102 History, Chicago, USA. 103 104 ACCEPTED MANUSCRIPT 105 3. Systematic Paleontology 106 Theropoda Marsh, 1881 107 Ceratosauria Marsh 1884 108 Abelisauridae Bonaparte & Novas, 1985 109 Brachyrostra Canale et al., 2009 110 Furileusauria Filippi et
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