A dinosaur ilium from the Late Triassic of Brazil with comments on key-character supporting Saturnaliinae

GARCIA, MAURÍCIO S.; PRETTO, FLÁVIO A.; DIAS-DA-SILVA, SÉRGIO; MÜLLER, RODRIGO T.

doi:10.1590/0001-3765201920180614

MAURÍCIO S. GARCIA

Curso de Ciências Biológicas, Centro de Ciências Naturais e Exatas, Universidade Federal de Santa Maria, Av. Roraima, 1000, Bairro Camobi, 97105-900 Santa Maria, RS, Brazil

Centro de Apoio à Pesquisa Paleontológica da Quarta Colônia, Universidade Federal de Santa Maria, Rua Maximiliano Vizzotto, 598, 97230-000 São João do Polêsine, RS, Brazil

http://orcid.org/0000-0001-5317-0140

FLÁVIO A. PRETTO

Centro de Apoio à Pesquisa Paleontológica da Quarta Colônia, Universidade Federal de Santa Maria, Rua Maximiliano Vizzotto, 598, 97230-000 São João do Polêsine, RS, Brazil

SÉRGIO DIAS-DA-SILVA

Laboratório de Paleobiodiversidade Triássica, Departamento de Ecologia e Evolução, Universidade Federal de Santa Maria, Av. Roraima, 1000, Bairro Camobi, 97105-900 Santa Maria, RS, Brazil

RODRIGO T. MÜLLER

Centro de Apoio à Pesquisa Paleontológica da Quarta Colônia, Universidade Federal de Santa Maria, Rua Maximiliano Vizzotto, 598, 97230-000 São João do Polêsine, RS, Brazil

Correspondence to: Maurício Silva Garcia E-mail: mauriciossauro@gmail.com ORCid: https://orcid.org/0000-0001-5317-0140

* Contribution to the centenary of the Brazilian Academy of Sciences.

AUTHOR CONTRIBUTIONS

MSG and RTM designed the study; FAP and RTM performed collection and preparation of the specimen; MSG, FAP, and RTM performed the analyses; MSG, FAP, SDS, and RTM discussed the results; MSG, FAP, SDS, and RTM wrote the article.

Figures (5)
Tables (1)

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Figure 1
Map of the São João do Polêsine area, Rio Grande do Sul state, southern Brazil, showing the distribution of dinosauromorph remains in the nearby fossiliferous sites (modified from Zerfass et al. 2007ZERFASS H, SANDER A and FLORES AF. 2007. Agudo, Folha SH.22-V.C.V, escala 1:100.000, Rio Grande do Sul. Serviço Geológico do Brasil (CPRM), Brasília.). “Marchezan” site (1), where an indeterminate herrerasaurid was collected; “Predebon” site (2), where tridactyl footprints attributed to indeterminate dinosaurs and to the ichnogenus Grallator were found (da Silva et al. 2008DA SILVA RC, CARVALHO IS and FERNANDES ACS. 2008. Pegadas de dinossauros do Triássico (Formação Santa Maria) do Brasil. Ameghiniana 45(4): 783-790.); “Buriol” site (3), where the holotypes and referred specimens of Buriolestes schultzi and Ixalerpeton polesinensis were collected (Cabreira et al. 2016, Müller et al. 2017bMÜLLER RT, PRETTO FA, STEFANELLO M, SILVA-NEVES E and DIAS-DA-SILVA S. 2017b. On a dinosaur axis from one of the oldest dinosaur-bearing sites worldwide. Acta Palaeontol Pol 62(3): 543-548., Müller et al. 2018a); “Piche site” (4), where CAPPA/UFSM 0200 was collected; “Janner” site (5), where the holotypes and referred specimens of Bagualosaurus agudoensis, Pampadromaeus barberenai and other dinosauriform remains were collected (Cabreira et al. 2011, Müller et al. 2014, 2015, 2017b, Pretto et al. 2015, Pretto et al. 2018); “Sacisaurus” site (6); whereas the holotype and referred specimens of Sacisaurus agudoensis and indeterminate sauropodomorphs were collected (Ferigolo and Langer 2007FERIGOLO J and LANGER MC. 2007. A Late Triassic dinosauriform from South Brazil and the origin of the ornithischian predentary bone. Historical Biology: An Inter J Paleo: 1-11.); “Wachholz” site (7), whereas the holotype and paratypes of the sauropodomorph Macrocollum itaquii were collected (Müller et al. 2018cMÜLLER RT, LANGER MC and DIAS-DA-SILVA S. 2018c. An exceptionally preserved association of complete dinosaur skeletons reveals the oldest long-necked sauropodomorphs. Biol Letters 14(11): 1744-9561.); “Linha São Luiz” site (8), whereas referred specimens of Guaibasaurus candelariensis were collected (Langer et al. 2011LANGER MC, BITTENCOURT JS and SCHULTZ CL. 2011. A reassessment of the basal dinosaur Guaibasaurus candelariensis, from the Late Triassic Caturrita Formation of south Brazil. Earth Env Sci T R So 101: 301-332.) and large tridactyl “theropod-like” footprints attributed to the ichnogenus Eubrontes were found (da Silva et al. 2012DA SILVA RC, BARBONI R, DUTRA T, GODOY MM, BINOTTO RB. 2012. Footprints of large theropod dinosaurs and implications on the age of Triassic biotas from southern Brazil. J S Am Earth Sci 39: 16-23.).

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Figure 2
Photographs and drawings of CAPPA/UFSM 0200, a sauropodomorph left ilium. a, lateral view. b, medial view. c, lateral view. d, medial view. Abbreviations: AW, Acetabular wall; BF, Brevis fossa; BS, Brevis shelf; C, Concavity; CV, Caudosacral vertebra; F. Foramen; IB, Iliac blade; IP, Ischiadic peduncle; MB, Medial blade; R, Rugosity; SC, Supraacetabular crest; SV I, Sacral primordial vertebra I; SV II, Sacral primordial vertebra II.

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Figure 3
Selection of dinosauriform ilia and highlight of the muscle scars on the pre and postacetabular alae and on the iliac blade (rugosities and striations). a, CAPPA/UFSM 0200 in lateral view; b, left ilium of Buriolestes schultzi (ULBRA-PVT280) in lateral view; c, left ilium of Buriolestes schultzi (ULBRA-PVT280) in lateral view; d, left ilium of Pampadromaeus barberenai (ULBRAPVT016) in lateral view; e, right ilium of Chromogisaurus novasi (PVSJ 845) in lateral view (reversed), modified from Martínez et al. 2013; f, right ilium of Saturnalia tupiniquim (MCP 3946-PV) in lateral view (reversed), modified from Bittencourt et al. 2012; g, left ilium of Saturnalia tupiniquim (MCP 3945-PV) in lateral view; h, right ilium of Plateosaurus (SMNS 12250) in lateral view (reversed), modified from Bittencourt et al. 2012; i, left ilium of Silesaurus opolensis (ZPAL AbIII404/1) in lateral view, modified from Peecook et al. 2013; j, left ilium of Lutungutali sitwensis (NHCC LB32); k, right ilium of Herrerasaurus ischigualastensis (PVL 2566) in lateral view (reversed), modified from Bittencourt et al. 2012. Specimens not to scale. Muscle scars marked in green are striations and in red are rugosities.

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Figure 4
Phylogenetic affinities of CAPPA/UFSM 0200, reduced strict consensus trees. a, Analysis employing the dataset by Cabreira et al. 2016; b, Analysis employing the dataset by Cabreira et al. 2016 (modified); c, Analysis employing the dataset by Martínez et al. 2013 (modified); d, Analysis employing the dataset by Langer et al. 2017 (modified); e, silhouette presenting a hypothesized hindlimb and pelvic girdle reconstruction to CAPPA/UFSM 0200. Purple box, Saurischia; Orange box, Theropoda; Light Blue box, Sauropodomorpha; Dark Blue box, Saturnaliinae. Numbers represent Bremer support values above 1.

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Figure 5
Hypothetical ontogenetic model to Saturnalia tupiniquim (above) and Buriolestes schultzi (below) based on the iliac muscle scars and morphology. Specimens placed from the “least mature” to the “most mature” from left to right. a, Saturnalia tupiniquim (MCP 3946-PV); b, Saturnalia tupiniquim (MCP 3945-PV); c, hypothetical ontogenetic advanced individual of Saturnalia tupiniquim (CAPPA/UFSM 0200); d, Buriolestes schultzi (putative juvenile specimen ULBRA-PVT056); e, Buriolestes schultzi (ULBRA-PVT280); f, Buriolestes schultzi (CAPPA/UFSM 0035); g, hypothetical ontogenetic advanced individual of Buriolestes schultzi (CAPPA/UFSM 0200). Specimens under scale.

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TABLE I
Specimens and references used in the comparisons. Specimens that were not seen first-hand by any of the authors are marked with an asterisk (*).

Figure 1 Map of the São João do Polêsine area, Rio Grande do Sul state, southern Brazil, showing the distribution of dinosauromorph remains in the nearby fossiliferous sites (modified from Zerfass et al. 2007ZERFASS H, SANDER A and FLORES AF. 2007. Agudo, Folha SH.22-V.C.V, escala 1:100.000, Rio Grande do Sul. Serviço Geológico do Brasil (CPRM), Brasília.). “Marchezan” site (1), where an indeterminate herrerasaurid was collected; “Predebon” site (2), where tridactyl footprints attributed to indeterminate dinosaurs and to the ichnogenus Grallator were found (da Silva et al. 2008DA SILVA RC, CARVALHO IS and FERNANDES ACS. 2008. Pegadas de dinossauros do Triássico (Formação Santa Maria) do Brasil. Ameghiniana 45(4): 783-790.); “Buriol” site (3), where the holotypes and referred specimens of Buriolestes schultzi and Ixalerpeton polesinensis were collected (Cabreira et al. 2016, Müller et al. 2017bMÜLLER RT, PRETTO FA, STEFANELLO M, SILVA-NEVES E and DIAS-DA-SILVA S. 2017b. On a dinosaur axis from one of the oldest dinosaur-bearing sites worldwide. Acta Palaeontol Pol 62(3): 543-548., Müller et al. 2018a); “Piche site” (4), where CAPPA/UFSM 0200 was collected; “Janner” site (5), where the holotypes and referred specimens of Bagualosaurus agudoensis, Pampadromaeus barberenai and other dinosauriform remains were collected (Cabreira et al. 2011, Müller et al. 2014, 2015, 2017b, Pretto et al. 2015, Pretto et al. 2018); “Sacisaurus” site (6); whereas the holotype and referred specimens of Sacisaurus agudoensis and indeterminate sauropodomorphs were collected (Ferigolo and Langer 2007FERIGOLO J and LANGER MC. 2007. A Late Triassic dinosauriform from South Brazil and the origin of the ornithischian predentary bone. Historical Biology: An Inter J Paleo: 1-11.); “Wachholz” site (7), whereas the holotype and paratypes of the sauropodomorph Macrocollum itaquii were collected (Müller et al. 2018cMÜLLER RT, LANGER MC and DIAS-DA-SILVA S. 2018c. An exceptionally preserved association of complete dinosaur skeletons reveals the oldest long-necked sauropodomorphs. Biol Letters 14(11): 1744-9561.); “Linha São Luiz” site (8), whereas referred specimens of Guaibasaurus candelariensis were collected (Langer et al. 2011LANGER MC, BITTENCOURT JS and SCHULTZ CL. 2011. A reassessment of the basal dinosaur Guaibasaurus candelariensis, from the Late Triassic Caturrita Formation of south Brazil. Earth Env Sci T R So 101: 301-332.) and large tridactyl “theropod-like” footprints attributed to the ichnogenus Eubrontes were found (da Silva et al. 2012DA SILVA RC, BARBONI R, DUTRA T, GODOY MM, BINOTTO RB. 2012. Footprints of large theropod dinosaurs and implications on the age of Triassic biotas from southern Brazil. J S Am Earth Sci 39: 16-23.).

Taxon	Specimen
Asilisaurus kongwe	NMT RB176* - Peecook et al. 2013PEECOOK BR, SIDOR CA, NESBITT SJ, SMITH RMH, STEYER JS and ANGIELCZYK KD. 2013. A new silesaurid from the upper Ntawere Formation of Zambia (Middle Triassic) demonstrates the rapid diversification of Silesauridae (Avemetatarsalia, Dinosauriformes). J Vert Paleontol 33(5): 1127. (Nesbitt et al. 2010NESBITT SJ, SIDOR CA, IRMIS RB, ANGIELCZYK KD, SMITH RMH and TSUJI LA. 2010. Ecologically distinct dinosaurian sister group shows early diversification of Ornithodira. Nature 464: 95-98.)
Bagualosaurus agudoensis	UFRGS-PV-1099-T - Pretto et al. 2018
Buriolestes schultzi	ULBRA-PVT056 (a putative juvenile individual)ULBRA-PVT280 - Cabreira et al. 2016CAPPA/UFSM 0035 - Müller et al. 2018a
Caseosaurus crosbyensis	NMMNH P-35995* - Hunt et al. 1998, Baron and Williams 2018UMMP 8870* - Hunt et al. 1998HUNT AP, LUCAS SG, HECKERT AB, SULLIVA R.M AND LOCKLEY MG. 1998. Late Triassic dinosaurs from the western United States. Géobios 31: 511-531., Baron and Williams 2018BARON MG and WILLIAMS ME. 2018. A re-evaluation of the enigmatic dinosauriform Caseosaurus crosbyensis from the Late Triassic of Texas, USA and its implications for early dinosaur evolution. Acta Palaeontol Pol 63(1): 129-145.
Chromogisaurus novasi	PVSJ 845 - Ezcurra 2010 (Martínez et al. 2012)
Efraasia minor	SMNS 12354* - Yates 2003bYATES AM. 2003b. The species taxonomy of the sauropodomorph dinosaurs from the Löwenstein Formation (Norian, Late Triassic) of Germany. Palaeontology 46(2): 317-337. (Bittencourt et al. 2012BITTENCOURT JS, LEAL LA, LANGER MC and AZEVEDO SAK. 2012. An additional basal sauropodomorph specimen from the Upper Triassic Caturrita Formation, southern Brazil, with comments on the biogeography of plateosaurids. Alcheringa 36(2): 269-278.)
Eoraptor lunensis	PVSJ 512 - Sereno et al. 2013SERENO PC, MARTÍNEZ RN and ALCOBER OA. 2013. Osteology of Eoraptor lunensis (Dinosauria, Sauropodomorpha). J Vert Paleontol 32: 83-179.
Guaibasaurus candelariensis	MCN PV2355 - Bonaparte Ferigolo and Ribeiro 1999BONAPARTE JF, FERIGOLO J and RIBEIRO AM. 1999. A new early late Triassic saurischian dinosaur from Rio Grande do Sul state, Brazil. National Science Museum Monographs 15: 89-109. UFRGS PV 0725T - Langer Bittencourt and Schultz 2011
Herrerasaurus ischigualastensis	PVL 2566 - Novas 1994 (Bittencourt et al. 2012, Baron and Williams 2018)
Ignotosaurus fragilis	PVSJ 883* - Martínez et al. 2013
Ixalerpeton polesinensis	ULBRA-PVT059 - Cabreira et al. 2016
Lessemsaurus sauropoides	PVL 4822* - Pol and Powell 2007POL D and POWELL JE. 2007. New information on Lessemsaurus sauropoides (Dinosauria: Sauropodomorpha) from the Upper Triassic of Argentina. Special Papers in Palaeontology 7: 223-243.
Lutungutali sitwensis	NHCC LB32* - Peecook et al. 2013
Marasuchus lilloensis	PVL 3870 - Sereno and Arcucci 1994
Melanorosaurus	NMQR 1551* - McPhee and Choiniere 2016MCPHEE BW and CHOINIERE JN. 2016. A hyper-robust sauropodomorph dinosaur ilium from the Upper Triassic-Lower Jurassic Elliot Formation of South Africa: Implications for the functional diversity of basal Sauropodomorpha. J Afr Earth Sci 123: 177e184.
Nhandumirim waldsangae	LPRP/USP 0651 - Marsola et al. 2019
Pampadromaeus barberenai	ULBRA-PVT016 - Cabreira et al. 2011
Panphagia protos	PVSJ 874 - Martínez and Alcober 2009MARTÍNEZ RN and ALCOBER OA. 2009. A basal sauropodomorph (Dinosauria: Saurischia) from the Ischigualasto Formation (Triassic, Carnian) and the early evolution of Sauropodomorpha. PLoS One 4 (2): e4397.
Pantydraco caducus	BMNH P77/1* - Yates 2003aYATES AM. 2003a. A new species of the primitive dinosaur Thecodontosaurus (Saurischia: Sauropodomorpha) and its implications for the systematics of early dinosaurs. J Syst Palaeontol 1: 1-42.
Plateosaurus	SMNS 12250* - Moser 2003MOSER M. 2003. Plateosaurus engelhardti Meyer, 1837 (Dinosauria: Sauropodomorpha) from the Feuerletten (Mittelkeuper, Upper Triassic) of Bavaria. Zitteliana 24: 3-186.
Riojasaurus incertus	PVL 3808* - Langer Bittencourt and Schultz 2011 PVL 3809* - Novas 1996PVL 3815* - Novas 1996
Saturnalia tupiniquim	MCP 3944-PV - Langer 2003MCP 3945-PV (Langer 2003, Marsola et al. 2019)MCP 3946-PV (Bittencourt et al. 2012)
Silesaurus opolensis	ZPAL AbIII404/1* - Peecoock et al. 2013 (Dzik 2003)ZPAL AbIII 907/6* - Langer Bittencourt and Schultz 2011 (Dzik 2003)
Staurikosaurus pricei	MCZ-1669* - Bittencourt and Kellner 2009BITTENCOURT JS and KELLNER AWA. 2009. The anatomy and phylogenetic position of the Triassic dinosaur Staurikosaurus pricei Colbert, 1970. Zootaxa 2079: 1-56. (Colbert 1970)
Thecodontosaurus antiquus	BMNH R1539* - Benton et al. 2000

[1] Correspondence to: Maurício Silva Garcia E-mail: mauriciossauro@gmail.com ORCid: https://orcid.org/0000-0001-5317-0140