A new theropod dinosaur from a peculiar Late Triassic assemblage of southern Brazil

https://doi.org/10.1016/j.jsames.2020.103026Get rights and content

Highlights

  • A new theropod dinosaur from the Late Triassic of Brazil is described.

  • It is one of the oldest known theropods worldwide.

  • This sheds light on some of the earliest theropod features.

  • The new theropod comes from a peculiar fauna dominated by traversodontid cynodonts.

  • The new theropod represents the first carnivorous dinosaur from this assemblage.

Abstract

The early evolution of theropod dinosaurs is poorly understood. Whereas sauropodomorphs are well-known from the oldest dinosaur bearing outcrops, the record of theropods is fragmentary and ambiguous. The Triassic deposits from Brazil yielded some of the oldest dinosaurs worldwide. These dinosaurs came from two distinct Assemblage Zones (AZ): the Hyperodapedon AZ (Carnian) and the Riograndia AZ (early Norian). Here, a specimen previously assigned to cf. Dinosauromorpha is reassessed. CAPPA/UFSM 0157 comes from an enigmatic assemblage with predominance of the traversodotind cynodont Siriusgnathus. This assemblage has been tentatively assigned to the Riograndia AZ. However, the absence of index fossils still hampers a reliable assignation. The specimen, which comprises a proximal portion of a left femur, belongs to a new theropod taxon erected here as Erythrovenator jacuiensis gen. et sp. nov. The new dinosaur differs from all other known Triassic dinosaurs based on the absence of a raised dorsolateral trochanter of the femur. Erythrovenator jacuiensis gen. et sp. nov. is regarded as a theropod on the basis of the pyramidal shape of the anterior trochanter in anterior view. The results of a phylogenetic analysis corroborate this assignation. Therefore, the new dinosaur represents one of the oldest theropod dinosaurs worldwide, shedding lights on some of the earliest theropod features. Finally, the new specimen also represents the first carnivorous dinosaur from the assemblage dominated by the traversodontid cynodont Siriusgnathus, increasing our knowledge of the faunal content of this enigmatic assemblage.

Introduction

The Upper Triassic beds from Brazil provide evidences of the origin and initial radiation of dinosaurs (Colbert 1970; Langer et al., 1999; Bonaparte et al., 1999; Cabreira et al., 2016; Marsola et al., 2018a; Pacheco et al., 2019). Distinct Assemblage Zones (AZs) mark different evolutionary stages of the group (Schultz et al., 2020). For instance, the Hyperodapedon AZ encompasses some of the oldest dinosaurs worldwide (Langer et al., 1999; Cabreira et al. 2011, 2016; Pacheco et al., 2019), whereas the Riograndia AZ yielded evidences from the moment when dinosaurs become bigger, more abundant, and geographically widespread (Bonaparte et al., 1999; Leal et al., 2004; Müller et al., 2018). This framework relies on radioisotopic data from typical localities of both AZs (Langer et al., 2018). However, some fossiliferous localities lack index fossils. This is the situation of the peculiar Niemeyer Site (Pavanatto et al., 2018). This site yielded several specimens of the traversodontid cynodont Siriusgnathus niemeyerorum (Pavanatto et al., 2018). In addition, were also reported probainognathian cynodonts and archosauriform remains (Pavanatto et al., 2018). This fossiliferous content was tentatively assigned to the Riograndia AZ based on the presence of Siriusgnathus niemeyerorum (Miron et al., 2020; see Marsola et al. 2018b). Among the archosauriform remains exhumed from the Niemeyer site, one particular specimen draws attention. CAPPA/UFSM 0157 comprises a proximal portion of a left femur of a dinosauromorph. This specimen was previously described by Pavanatto et al. (2018) and was regarded as having dinosauromorph affinities. Despite its fragmentary condition, CAPPA/UFSM 0157 is particularly interesting because it combines a set of unusual traits among Late Triassic dinosauromorphs. However, the phylogenetic affinities of this specimens have not been hypothesized with computational analysis. Furthermore, the unique anatomy of CAPPA/UFSM 0157 suggests that it belongs to an undescribed taxon. Hence, here the specimen is ascribed to a new taxon and its phylogenetic affinities analysed. Moreover, the former description by Pavanatto et al. (2018) is expanded based on new observations and comparisons.

AMNH, American Museum of Natural History, New York, New York, USA; CAPPA/UFSM, Centro de Apoio à Pesquisa Paleontológica da Quarta Colônia/Universidade Federal de Santa Maria, São João do Polêsine, Rio Grande do Sul, Brazil; MCN, Museu de Ciências Naturais, Fundação Zoobotânica do Rio Grande do Sul, Porto Alegre, Brazil; MCP, Museu de Ciências e Tecnologia Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre, Brazil; MCZ, Museum of Comparative Zoology, Harvard, Cambridge, USA; MMACR, Museu Municipal Aristides Carlos Rodrigues, Candelária, Rio Grande do Sul, Brazil; MNA, Museum of Northern Arizona, Flagstaff, Arizona, USA; NHMUK, Natural History Museum, London, UK; NMT, National Museum of Tanzania, Dar es Salaam, Tanzania; PVL, Paleontología de Vertebrados, Instituto Miguel Lillo, Tucumán, Argentina; SAM-PK, Iziko South African Museum, Cape Town, South Africa; UCMP, University of California Museum of Paleontology, Berkeley, California, USA; ULBRA, Universidade Luterana do Brasil, Coleção de Paleovertebrados, Canoas, Brazil; ZPAL, Institute of Paleobiology of the Polish Academy of Sciences in Warsaw, Poland.

Section snippets

Material and methods

In order to test its phylogenetic affinities, the new taxon was scored in the data matrix of Müller and Garcia (2020). The former data matrix includes 277 characters and 62 terminal taxa. The data matrix was subjected to an equally weighted parsimony analysis in TNT v. 1.1 (Goloboff et al., 2008). The characters treated by Müller and Garcia (2020) as additive (4, 13, 18, 25, 63, 82, 84, 87, 89, 109, 142, 166, 174, 175, 184, 186, 190, 201, 203, 205, 209, 212, 225, 235, 236, 239, 250 and 256)

Geological settings

The Niemeyer Site is located in municipality of Agudo (29°40′25″ S; 53°14′4.20″ W), Rio Grande do Sul, Brazil (Pavanatto et al., 2018, Fig. 1). Site strata belong to the Candelária Sequence (Horn et al., 2014), which is part of the Santa Maria Suspersequence of the Paraná Basin (Zerfass et al., 2003). The stratigraphic column of the Niemeyer Site comprises a 23m thick lithology of reddish mudstones, sandstones, and intraformational conglomerates, with carbonatic concretions and hydromorphic

Systematic paleontology

Dinosauriformes Novas, 1992.

Dinosauria Owen, 1842.

Theropoda Marsh, 1881.

Erythrovenator gen. nov.

Type species. Erythrovenator jacuiensis gen et sp. nov., monotypic, see below.

Etymology. ‘Erythro’ comes from the Greek erythrós, meaning “red” or “reddish”, referring to the red color of the holotype fossil; ‘venator’ is the Latin word for hunter.

Diagnosis. Same as for the only known species.

Erythrovenator jacuiensis sp. nov. (Fig. 2).

Etymology. The species name ‘jacuiensis’ is in reference to ‘Rio

Phylogenetic analysis

The phylogenetic analysis recovered 36 most parsimonious trees (MPTs) of 986 steps each, with a consistency index of 0.319, a retention index of 0.665, and a rescaled consistency index of 0.212. In all the MPTs, Erythrovenator jacuiensis nests as the basalmost member of Theropoda (Fig. 4). The node is supported by an anterior trochanter separated from the femoral shaft by a marked cleft [212 (1 → 2)]. The same character state seems independently acquired in some ‘silesaurids’ (e.g. Silesaurus

Discussion and conclusions

Erythrovenator jacuiensis is here regarded as a theropod dinosaur based on the pyramidal shape of the anterior trochanter (Fig. 2A), a trait typically present in neotheropods (Tykoski and Rowe 2004). Conversely, in sauropodomorphs, such as Saturnalia tupiniquim (MCP 3844-PV), Pampadromaeus barberenai (ULBRA-PVT016), and Guaibasaurus candelariensis (MCN PV2355), the proximal tip of the anterior trochanter does not tappers to a point. Erythrovenator jacuiensis also differs from these dinosaurs

Funding

This research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors.

Author statement

Rodrigo Temp Müller: Conceptualization, Methodology, Analysis, Writing, Visualization.

Declaration of competing interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Acknowledgements

I thank Federico Agnolin, Mateusz Tałanda, and an anonymous reviewer for comments and suggestions that improved the quality of this manuscript. Additionally, I thank the Willi Henning Society, for the gratuity of TNT software.

References (59)

  • A.B. Arcucci et al.

    A new Triassic carnivorous dinosaur from Argentina

    Ameghiniana

    (2003)
  • M.G. Baron et al.

    Postcranial anatomy of Lesothosaurus diagnosticus (Dinosauria: Ornithischia) from the Lower Jurassic of southern Africa: implications for basalornithischian taxonomy and systematics

    Zool. J. Linn. Soc.

    (2017)
  • M.G. Baron et al.

    A novel hypothesis of dinosaur relationships and early dinosaur evolution

    Nature

    (2017)
  • J.D.S. Bittencourt et al.

    The anatomy and phylogenetic position of the Triassic dinosaur Staurikosaurus pricei Colbert, 1970

    Zootaxa

    (2009)
  • J.F. Bonaparte et al.

    A new early Late Triassic saurischian dinosaur from Rio Grande do Sul state, Brazil. Proceedings of the second gondwanan dinosaur Symposium

    Natl. Sci. Mus. Monogr.

    (1999)
  • R.J. Butler

    The anatomy of the basal ornithischian dinosaur Eocursor parvus from the lower Elliot Formation (Late Triassic) of South Africa

    Zool. J. Linn. Soc.

    (2010)
  • S.F. Cabreira et al.

    New stem-sauropodomorph (Dinosauria, Saurischia) from the Triassic of Brazil

    Naturwissenschaften

    (2011)
  • E.H. Colbert

    A saurischian dinosaur from the Triassic of Brazil

    Am. Mus. Novit.

    (1970)
  • M.D. Ezcurra et al.

    A revision of the early neotheropod genus Sarcosaurus from the early Jurassic (Hettangian–Sinemurian) of central England

    Zool. J. Linn. Soc.

    (2020)
  • M.D. Ezcurra

    A new early dinosaur (Saurischia: Sauropodomorpha) from the Late Triassic of Argentina: a reassessment of dinosaur origin and phylogeny

    J. Syst. Palaeontol.

    (2010)
  • M.D. Ezcurra

    A new early coelophysoid neotheropod from the Late Triassic of northwestern Argentina

    Ameghiniana

    (2017)
  • M.D. Ezcurra et al.

    Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NW Argentina

    Hist. Biol.

    (2007)
  • P.M. Galton

    Notes on the postcranial anatomy of the heterodontosaurid dinosaur Heterodontosaurus tucki, a basal ornithischian from the Lower Jurassic of South Africa

    Rev. Paléobiol.

    (2014)
  • M.S. Garcia et al.

    On the taxonomic status of Teyuwasu barberenai Kischlat, 1999 (Archosauria: Dinosauriformes), a challenging taxon from the Upper Triassic of southern Brazil

    Zootaxa

    (2019)
  • P.A. Goloboff et al.

    TNT: a free program for phylogenetic analysis

    Cladistics

    (2008)
  • C.T. Griffin et al.

    The femoral ontogeny and long bone histology of the Middle Triassic (? late Anisian) dinosauriform Asilisaurus kongwe and implications for the growth of early dinosaurs

    J. Vertebr. Paleontol.

    (2016)
  • F. von Huene

    Ein neuer Coelurosaurier in der Thüringischen Trias

    Palaeontol. Z.

    (1934)
  • R.B. Irmis et al.

    A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs

    Science

    (2007)
  • M.C. Langer

    The origins of Dinosauria: much ado about nothing

    Palaeontology

    (2014)
  • Cited by (9)

    • Multiproxy taphonomic analysis in the Los Colorados Formation (Upper Triassic), Ischigualasto-Villa Unión Basin, Argentina: A case study through sauropodomorph remains

      2022, Journal of South American Earth Sciences
      Citation Excerpt :

      It is interesting to note that whereas carnian sauropodomorphs, such as Buriolestes, Pampadromaeus and Eoraptor were faunivorous/carnivorous based on the presence of recurved and finely serrated teeth and resembling the ancestral dinosaurian condition (Bronzati et al., 2017; Müller, 2020; Pol et al., 2021), Norian forms modified the dentition to an omnivorous/herbivorous diet, based on the presence of lancelotale teeth (Galton and Upchurch, 2004; McPhee et al., 2020; Müller, 2020; Pol et al., 2021). Accepting the hypothesis of herrerasaurids as non-theropod saurischians, and in contrast to the sauropodomorph record, theropod dinosaurs are so far scarce in Carnian strata (Langer et al., 2010; Müller, 2021). Conversely, theropods start to be abundant as from Norian beds (Bonaparte, 1999; Arcucci and Coria, 2003; Nesbitt and Ezcurra, 2015; Ezcurra, 2017; Martínez and Apaldetti, 2017).

    • Review of the fossil record of early dinosaurs from South America, and its phylogenetic implications

      2021, Journal of South American Earth Sciences
      Citation Excerpt :

      This part of the Santa Maria Formation and the lower part of the Caturrita Formation are enclosed in the Candelária Sequence (Horn et al., 2014), bearing two distinctive AZs: the older Hyperodapedon AZ and the younger Riograndia AZ. The Hyperodapedon AZ yields the dinosaurs Staurikosaurus pricei, Gnathovorax cabrerai, Saturnalia tupiniquim, Buriolestes schultzi, Pampadromaeus barberenai, Bagualosaurus agudoensis, and Nhandumirim waldsangae (e.g., Langer et al., 1999; Cabreira et al., 2016; Pacheco et al., 2019), whereas the Riograndia AZ has yielded Macrocollum itaquii, Unaysaurus tolentinoi, unnamed sauropodomorph specimens, Erythrovenator jacuiensis and Guaibasaurus candelariensis (Bonaparte et al., 1999; Leal et al., 2004; Langer et al., 2010; Müller et al., 2017, 2018a; Müller, 2020a) (Fig. 2). Radioisotopic age data was obtained for these two AZs (Langer et al., 2018): the maximum deposition ages obtained were ~233 Ma for the base of the Hyperodapedon AZ and ~225 Ma for the Riograndia AZ.

    View all citing articles on Scopus
    View full text