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Buitreraptor Gonzalezorum[I] A peer-reviewed version of this preprint was published in PeerJ on 26 March 2018. View the peer-reviewed version (peerj.com/articles/4558), which is the preferred citable publication unless you specifically need to cite this preprint. Gianechini FA, Makovicky PJ, Apesteguía S, Cerda I. 2018. Postcranial skeletal anatomy of the holotype and referred specimens of Buitreraptor gonzalezorum Makovicky, Apesteguía and Agnolín 2005 (Theropoda, Dromaeosauridae), from the Late Cretaceous of Patagonia. PeerJ 6:e4558 https://doi.org/10.7717/peerj.4558 Postcranial skeletal anatomy of the holotype and referred specimens of Buitreraptor gonzalezorum Makovicky, Apesteguía and Agnolín 2005 (Theropoda, Dromaeosauridae), from the Late Cretaceous of Patagonia Federico A Gianechini Corresp., 1 , Peter J Makovicky 2 , Sebastián Apesteguía 3 , Ignacio Cerda 4 1 Instituto Multidisciplinario de Investigaciones Biológicas (IMIBIO-SL), CONICET-Universidad Nacional de San Luis., San Luis, Argentina 2 Field Museum of Natural History, Chicago, Illinois, United States 3 CONICET. Fundación de Historia Natural ‘Félix de Azara’, CEBBAD, Universidad Maimónides., Buenos Aires, Argentina 4 CONICET. Instituto de Investigación en Paleobiología y Geología, Universidad Nacional de Río Negro, General Roca, Río Negro, Argentina Corresponding Author: Federico A Gianechini Email address: [email protected] Here we provide a detailed description of the postcranial skeleton of the holotype and referred specimens of Buitreraptor gonzalezorum. This taxon was recovered as a unenlagiine dromaeosaurid in several recent phylogenetic studies and is the best represented Gondwanan dromaeosaurid discovered to date. It was preliminarily described in a brief article, but a detailed account of its osteology is becoming known for recent work. Hitherto, the holotype is the most complete specimen found, so its exhaustive description provides much valuable anatomical information. The holotype and referred specimens preserve the axial skeleton, pectoral and pelvic girdles, and both fore- and hindlimbs. Diagnostic postcranial characters of this taxon include: anterior cervical centra exceeding the posterior limit of neural arch; antepenultimate and penultimate cervical vertebral centra with lateroventral tubercles; pneumatic foramina only in anteriormost dorsals; medial and distal caudal centra with a complex of ridges on lateral surfaces; pneumatic furcula with two pneumatic foramina on the ventral surface; scapular blade transversely expanded at mid-length; well-projected flexor process on distal humerus; dorsal rim of the ilium laterally everted; and concave dorsal rim of the postacetabular iliac blade. A paleohistological study of limb bones shows that the holotype represents a younger ontogenetic stage than one of the referred specimens (MPCA 238), which correlates with the fusion of the last sacral vertebra to the rest of the sacrum in MPCA 238. A revised phylogenetic analysis recovered Buitreraptor as a unenlagiine dromaeosaurid, in agreement with previous works. The phylogenetic implications of the unenlagiine synapomorphies and other characters, such as the specialized pedal digit II and the distal ginglymus on metatarsal II, are discussed within the evolutionary framework of Paraves. PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.26450v1 | CC BY 4.0 Open Access | rec: 22 Jan 2018, publ: 22 Jan 2018 1 Postcranial skeletal anatomy of the holotype and referred 2 specimens of Buitreraptor gonzalezorum Makovicky, 3 Apesteguía and Agnolín 2005 (Theropoda, Dromaeosauridae), 4 from the Late Cretaceous of Patagonia 5 Federico A. Gianechini1, Peter J. Makovicky2, Sebastián Apesteguía3 and Ignacio Cerda4 6 1 Instituto Multidisciplinario de Investigaciones Biológicas (IMIBIO-SL), CONICET-Universidad Nacional de 7 San Luis. Área de Zoología, Facultad de Química, Bioquímica y Farmacia, UNSL, Ciudad de San Luis, San 8 Luis, Argentina. 9 2 Field Museum of Natural History, Chicago, Illinois, U.S.A. 10 3 CONICET. Fundación de Historia Natural ‘Félix de Azara’, CEBBAD, Universidad Maimónides, Buenos 11 Aires, Argentina. 12 4 Instituto de Investigación en Paleobiología y Geología, CONICET-Universidad Nacional de Río Negro, 13 General Roca, Río Negro, Argentina. 14 15 Corresponding Author: 16 Federico A. Gianechini1 17 Ejército de los Andes 950, Ciudad de San Luis, San Luis, D5700HHV, Argentina 18 Email address: [email protected] 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 Abstract 38 39 Here we provide a detailed description of the postcranial skeleton of the holotype and referred 40 specimens of Buitreraptor gonzalezorum. This taxon was recovered as a unenlagiine 41 dromaeosaurid in several recent phylogenetic studies and is the best represented Gondwanan 42 dromaeosaurid discovered to date. It was preliminarily described in a brief article, but a detailed 43 account of its osteology is becoming known for recent work. Hitherto, the holotype is the most 44 complete specimen found, so its exhaustive description provides much valuable anatomical 45 information. The holotype and referred specimens preserve the axial skeleton, pectoral and 46 pelvic girdles, and both fore- and hindlimbs. Diagnostic postcranial characters of this taxon 47 include: anterior cervical centra exceeding the posterior limit of neural arch; antepenultimate and 48 penultimate cervical vertebral centra with lateroventral tubercles; pneumatic foramina only in 49 anteriormost dorsals; medial and distal caudal centra with a complex of ridges on lateral 50 surfaces; pneumatic furcula with two pneumatic foramina on the ventral surface; scapular blade 51 transversely expanded at mid-length; well-projected flexor process on distal humerus; dorsal rim 52 of the ilium laterally everted; and concave dorsal rim of the postacetabular iliac blade. A 53 paleohistological study of limb bones shows that the holotype represents a younger ontogenetic 54 stage than one of the referred specimens (MPCA 238), which correlates with the fusion of the 55 last sacral vertebra to the rest of the sacrum in MPCA 238. A revised phylogenetic analysis 56 recovered Buitreraptor as a unenlagiine dromaeosaurid, in agreement with previous works. The 57 phylogenetic implications of the unenlagiine synapomorphies and other characters, such as the 58 specialized pedal digit II and the distal ginglymus on metatarsal II, are discussed within the 59 evolutionary framework of Paraves. 60 1. Introduction 61 62 Buitreraptor gonzalezorum is a paravian theropod whose remains were found in Late Cretaceous 63 outcrops of the Candeleros Formation in the ‘La Buitrera’ fossiliferous area, in Patagonia, 64 Argentina. The paleontological significance of the many transformative discoveries made in this 65 site were already enumerated in our previous papers (e.g., Apesteguía & Zaher, 2006; 66 Gianechini, Makovicky & Apesteguía, 2011; Rougier, Apesteguía & Gaetano, 2011; Makovicky, 67 Apesteguía & Gianechini, 2012; Gianechini & de Valais, 2016). 68 Most phylogenetic analyses performed on coelurosaurian theropods recover Buitreraptor 69 as member of a monophyletic Dromaeosauridae, within the subfamily Unenlagiinae, although 70 some recent analyses recover Buitreraptor and other unenlagiines as stem avialans (Agnolín & 71 Novas, 2011, 2013). Dromaeosauridae has experienced a remarkable increase in diversity since 72 the 2000’s, including species with both small and large body sizes, some of them represented by 73 almost complete skeletons preserving plumage (e.g., Currie & Varricchio 2004; Norell et al., 74 2006; Turner, Hwang & Norell, 2007; Turner et al., 2007; Longrich & Currie, 2009; Xu et al., 75 2010; Zheng et al., 2010; Evans, Larson & Currie, 2013; Han et al., 2014; DePalma et al., 2015; 76 Lü & Brusatte, 2015). Unfortunately, the Gondwanan record of these theropods remains sparse. 77 Hitherto, the most significant Gondwanan specimens were found in Patagonia and Madagascar. 78 The Patagonian record of unenlagiines currently includes five taxa, three of which preserve only 79 post-cranial remains: Unenlagia comahuensis, Unenlagia paynemili and Neuquenraptor 80 argentinus and two species with cranial remains, i.e., Buitreraptor gonzalezorum and 81 Austroraptor cabazai. Recently, a fragmentary coelurosaurian represented only by hindlimb 82 remains, but with possible deinonychosaurian affinities was described as Pamparaptor micros 83 (Porfiri, Calvo & Dos Santos, 2011). Its potential relationships with unenlagiines have not been 84 thoroughly evaluated yet. 85 Buitreraptor is the best represented unenlagiine to date. Its holotype consists of an 86 almost complete, semi-articulated, and very well-preserved skeleton and at least five referred 87 specimens are known. This taxon was named by Makovicky, Apesteguía & Agnolín (2005), who 88 concluded that it is the earliest dromaeosaurid found so far in Gondwana. The anatomical 89 information provided by this taxon was significant for uniting Gondwanan dromaeosaurids 90 within their own monophyletic clade, and for the understanding of the character distributions and 91 morphological trends in paravian phylogeny. However, some traits of Buitreraptor, and also of 92 other unenlagiines, are similar to the anatomy of basal avialans, lending support to the alternate 93 phylogenetic hypothesis proposed by Agnolín & Novas (2011, 2013) mentioned above. 94 Buitreraptor was only briefly described by Makovicky, Apesteguía & Agnolín (2005), 95 and a detailed osteology has been wanting. We recently provided
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