Taurovenator

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Taurovenator
Temporal range: Late Cretaceous, (Cenomanian), ~95–93.9  Ma
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Taurovenator violantei.png
Reconstruction of the head of Taurovenator violantei
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Family:† Carcharodontosauridae
Tribe:† Giganotosaurini
Genus:† Taurovenator
Motta et al., 2016
Type species
†Taurovenator violantei
Motta et al., 2016

Taurovenator ("bull hunter") is a genus of large carcharodontosaurid theropod that lived in what is now Argentina during the Cenomanian age of the Late Cretaceous around 95 to 93.9 million years ago. Initially only known from a single postorbital skull bone described from the Huincul Formation, Taurovenator violantei was named in 2016 by Argentine paleontologist Matias Motta and a group of colleagues. Debate came about over the validity of the genus and species, with some paleontologists arguing that Taurovenator was synonymous with the earlier-named carcharodontosaurid Mapusaurus . This was until a 2024 study referred a well-preserved partial skull and incomplete skeleton to Taurovenator.

Contents

Discovery and naming

Initial discovery and description

In 2005, a right postorbital of a theropod dinosaur was unearthed by Argentine paleontologist Matias Motta from a section of sandstone strata deriving from the lower member of the Huincul Formation, dating to the Cenomanian stage of the Late Cretaceous (95-93.9 million years old). [1] The strata located on the Violante Farm, a farm located in Rio Negro Province, Argentina. This fossil was found in the vicinity of the megaraptoran Aoniraptor, several abelisauroids, and a possible unenlagiine paravian. [2] The fossils recovered were then transported to the Museo Provincial "Carlos Ameghino", with the isolated postorbital deposited under catalogue number MPCA-Pv 803. [2] This postorbital remained undescribed until in 2016, when Matias Motta and colleagues described the postorbital as the holotype (name-bearing specimen) of a new genus and species of carcharodontosaurid dinosaur, Taurovenator violantei. Taurovenator was then the second carcharodontosaurid described from the Huincul Formation, with Mapusaurus named in 2006 based on several incomplete skeletons. [3] [4] The generic name Taurovenator means "bull hunter", derived from the Latin roots taurus "bull" and venor "hunter" whereas the specific name is in honor of Violante Farm where the postorbital was unearthed. [2] [1]

Validity debate and additional specimen

Taurovenator went largely unnoticed due to its fragmentary nature, [5] [1] with Coria et al. (2019) suggesting that Taurovenator is synonymous with Mapusaurus, considering both of its original autapomorphies as shared with Mapusaurus and also pointing out that both taxa shared a curved lateral margin of the palpebral. [6] Additionally, the authors considered that there was a high likelihood of them being coeval, [6] however, Taurovenator is actually from the lower unit of the Huincul Formation, while Mapusaurus is from the upper unit of the formation. [1] Researcher Mickey Mortimer also believed that Taurovenator was synonymous with Mapusaurus due to the lack of sufficient material and diagnostic traits for the genus. [7] In 2022, another carcharodontosaurid from the Huincul Formation, Meraxes, was named on the basis of a well-preserved skull and partial skeleton from the same strata as Taurovenator. In their description of Meraxes, the authors stated that Taurovenator lacks sufficient diagnostic characters and may be coeval with Meraxes. [5] [1]

Notably in the same site on Violante Farm as the Taurovenator holotype, an associated skeleton (MPCA-Pv 803) including a partial skull and posterior (back portion) mandible, incomplete cervical (neck vertebrae) series, fragments of dorsal (back) vertebrae, several ribs, two partial forelimbs, a femur (thighbone), a partial pes (foot), gastralia, and a caudal vertebra was unearthed along with the Taurovenator holotype in 2005 but were regarded as belonging to an indeterminate carcharodontosaurid in the 2016 description of Taurovenator. Additionally, the material of MPCA-Pv 803 had not been completely collected, prepared, or made available for study, delaying its description. [2] [1] These fossils were also spread out over a sizeable area and intermingled with bones of an indeterminate sauropod. In 2024, this specimen was described and, despite not overlapping in material with the holotype, was referred to Taurovenator. This was done for several reasons: the holotype fits perfectly with MPCA-Pv 803 suggesting they may be from the same individual, the close proximity to the holotype (the specimen was found 800 metres (870 yd) away from the holotype's dig site, and the fossils are congruent in size. In an analysis of the strata from which the holotype was discovered, the 2024 study noted that the Huincul Formation is separated into two distinct sequences; a lower section of thin, multicolored sandstones and an upper section of thick conglomeratic sediments. Mapusaurus derives from the upper sequence of the formation, whereas Meraxes and Taurovenator are exclusive to the lower rock layers. Meraxes, however, was collected in strata close to the Candeleros-Huincul Formation boundary, whereas Taurovenator's specimens were found over 30 meters above the Candeleros-Huincul Formation limit. It is for these reasons that the three carcharodontosaurids found at Huincul were potentially not coeval, supporting the argument for Taurovenator's validity. Additionally, the holotype preserve features of the Giganotosaurini, further supporting its referral to Taurovenator. A new host of diagnostic traits were found on the bones of MPCA-Pv 803, properly demonstrating its distinctiveness from Mapusaurus and Meraxes. [1]

Description

Taurovenator is a very large carcharodontosaurid. It had an estimated body mass of 5,728 kg (12,628 lb), 5.728 metric tons (6.314 short tons), based on a formula that utilizes the circumference of the femur to predict body mass. For reference, this is smaller than Giganotosaurus 6,349 kilograms (13,997 lb), but larger than Meraxes and Mapusaurus,4,263 kilograms (9,398 lb) and 4,343 kilograms (9,575 lb), respectively). [1] [8] This puts Taurovenator among the largest theropod dinosaurs known and therefore one of the biggest terrestrial carnivores known to science, though still smaller than genera like Tyrannosaurus and Spinosaurus . [9] [10] [11] [12]

Taurovenator life reconstruction, compared to a person. Restauracion en vida de Taurovenator violantei.png
Taurovenator life reconstruction, compared to a person.

Skull

The skull of Taurovenator is known from the holotype postorbital in addition to the cranial and mandibular elements of MPCA-Pv 803 which include: both jugals (cheekbones), left quadrate (skull bone articulating with the lower jaw), left palatine (main palate bone), right ectopterygoid (smaller palate bone), both surangulars and angulars (back of lower jaw bones), prearticulars (rearmost lower jaw bone), left articular (lower jaw bone articulating with the skull), right ceratohyal (throat bone), and three isolated teeth. The maximum length of the skull of MPCA-Pv 803 was estimated to be around 135 centimetres (4.43 ft) based on the more complete skulls of relatives like Meraxes. The postorbital is strongly rugose and projected out like a horn, markedly different from the orbital bosses of other carcharodontosaurids. Both Motta et al., 2016 and Rolando et al., 2024 consider this a unique trait, [2] [1] but its sister taxon Meraxes too has a postorbital that was described as laterally projecting out like a horn. [5] The 2024 description of MPCA-Pv 803 however argued that this horn-like projection is distinct from that of Meraxes, with the projection measuring 3–4 centimetres (1.2–1.6 in) in height and 2–3 centimetres (0.79–1.18 in) in diameter. In addition, there is a deep fossa (depression in bone) and excavation on the ventral surface of the postorbital. Though the excavations are observable in the postorbitals of Mapusaurus, Meraxes, and other carcharodontosaurids, they all do not preserve the unique fossa found on the Taurovenator holotype. [1] [2] [3]

The jugal is long and sub-triangular in shape with a slightly posteromedially (backwards) pointed postorbital ramus . Its lateral (exposed side) surface is rugose with a thick shelf, a condition similarily observed in South American carcharodontosaurids like Tyrannotitan and Meraxes but absent from the jugals of other allosauroids. The anterior (front) process of the jugal is compressed and bears an ovular pneumatic foramen (pits in bone for air sac systems) on its articulating (contacting) surface with the lacrimal . The quadrate of MPCA-PV 803 is distinctly stout, triangular in lateral view, bearing a large, anteriorly projected pterygoid flange as in related genera. The pterygoid fossa is wide, subdivided by a crest, and lacks pneumaticity as in contemporary carcharodontosaurids of its time. In contrast the palatine of Taurovenator is tetraradiate with an anteriorly expanded vomeropterygoid process (expansion), a short and sub-triangular pterygoid process, a narrow jugal process, and an elongated maxillary process. The ectopterygoid is C-shaped from dorsoventral (top and bottom) views with a pneumatic recess on its medial surface, a trait in other tetanurans. A distinguishing feature of Taurovenator's pterygoid is the thin, long jugal process which is longer than those of Meraxes and Giganotosaurus. As for the mandible, only the posterior portion is known from Taurovenator. It preserves a robust and dorsoventrally (top-down) high morphology, akin to the mandibles of Acrocanthosaurus, Tyrannosaurus, and other tyrannosauroids. Two teeth are preserved from Taurovenator, both of which are large, around 4 centimetres (1.6 in) in length. The margins of these teeth are convex and straight, with chisel-like denticles which number around 8.5 per 5 millimetres (0.20 in) at the mid-crown and 11 per 5 millimetres (0.20 in) at the base. Their blood grooves are well developed and their enamel wrinkles are notably deep, [1] a distinct characteristic of carcharodontosaurid teeth. [13] [14]

Postcrania

Distinctively, Taurovenator's neck vertebrae bore prominent neural spines with flange-like dorsal tips. As a result, the neural spines of cervical vertebrae C3-C6 are "imbricated", ie interlocking with each other. The authors describing this morphology termed this unusual structure a "cervical complex", and likened them to overlapping roof tiles. A similar, though less extreme condition is also known in the C3-C5 of the more basal carcharodontosaurid Acrocanthosaurus . [1] [15] Available information of Giganotosaurus and Mapusaurus further suggests that this "cervical complex" is a unique synapomorphy of the group. The presence of the cervical complex would have likely restricted the range of movement of the cervical vertebrae. On the other hand, the skull of Taurovenator and other carcharodontosaurids had a ball-shaped occipital condyle similar to that seen in the skulls of ceratopsian dinosaurs. This could have allowed a large range of rotational movement between the skull and the first cervical vertebra. Furthermore, the cervical complex of Taurovenator could have similar functional implications to those of the syncervical vertebrae (ie fused C1-C3 vertebrae) of ceratopsians, strengthening the anterior region of the neck, and increasing the surface area for epaxial cervico-cranial muscles. [1]

Only remains of two dorsal vertebrae are known. One is composed of a centrum, but the other is composed of a very tall, 52 cm high neural arch. [1]

Taurovenator had proportionately the smallest arms of all known allosauroids. The nearly completely preserved arms were reduced to a greater degree than even in other carcharodontosaurids, being proportionately smaller than that of taxa such as Meraxes , particularly where the forearm is concerned. Despite such limb reduction, the forearms were robust, and the digits had a great degree of flexibility. Nonetheless like other giganotosaurines, the forelimbs were likely incapable of a wide range of movement. [1]

Taurovenator also shares with Meraxes an enlarged ungual claw on the second toe, approximately 20% longer than the equivalent phalanx of the third toe and more laterally compressed. [1]

Classification

Motta et al. (2016) suggested that Taurovenator occupied a derived position within Carcharodontosauridae, comparing it to Giganotosaurus, Carcharodontosaurus and Mapusaurus in particular. [2] Coria et al. (2019) suggested that Taurovenator is synonymous with Mapusaurus, considering both of its original autapomorphies as shared with Mapusaurus and also pointing out that both taxa shared a curved lateral margin of the palpebral. [6] Additionally, the authors considered that there was a high likelihood of them being coeval, [6] however, Taurovenator is actually from the lower unit of the Huincul Formation, while Mapusaurus is from the upper unit of the formation. [1] Rolando et al. (2024) reaffirmed Taurovenator's validity, considering the autapomorphies preserved on the holotype as more strongly developed in Taurovenator than any other carcharodontosaurid, while also considering the supposedly diagnostic curved margin of the palpebral as a more widely distributed feature in Carcharodontosauridae. [1]

In order to test the systematics of Taurovenator with the information supplemented by the new specimen, the study used the phylogenetic dataset used in the description of Meraxes, with some additional data. The results of their phylogenetic analysis are shown in a cladogram below: [1] [5]

Carcharodontosauridae

Palaeoenvironment

Several dinosaurs from the Huincul Formation (Taurovenator is not figured) Huincul Formation Dinosauria Scale.svg
Several dinosaurs from the Huincul Formation (Taurovenator is not figured)

Taurovenator was discovered in the Argentine Province of Neuquén. It was found in the Huincul Formation, a rock formation bordering the Río Limay Subgroup, the latter of which is a subdivision of the Neuquén Group. This unit is located in the Neuquén Basin in Patagonia. The Huincul Formation is composed of yellowish and greenish sandstones of fine-to-medium grain, some of which are tuffaceous. [16] These deposits were laid down during the Upper Cretaceous, either in the middle Cenomanian to early Turonian stages [2] or the early Turonian to late Santonian. [17] The deposits represent the drainage system of a braided river. [18]

Fossilised pollen indicates a wide variety of plants were present in the Huincul Formation. A study of the El Zampal section of the formation found hornworts, liverworts, ferns, Selaginellales, possible Noeggerathiales, gymnosperms (including gnetophytes and conifers), and angiosperms (flowering plants), in addition to several pollen grains of unknown affinities. [19] The Huincul Formation is among the richest Patagonian vertebrate associations, preserving fish including dipnoans and gar, chelid turtles, squamates, sphenodonts, neosuchian crocodilians, and a wide variety of dinosaurs. [2] [20] Vertebrates are most commonly found in the lower, and therefore older, part of the formation. [21]

In addition to Taurovenator, the theropods of the Huincul Formation are represented by the other giant carcharodontosaurids Meraxes and Mapusaurus, abelisaurids including Skorpiovenator , [22] Ilokelesia , and Tralkasaurus , [23] noasaurids such as Huinculsaurus , [24] paravians such as Overoraptor , [25] and other theropods such as Aoniraptor and Gualicho [26] have also been discovered there. [2] Several iguanodonts are also present in the Huincul Formation. [16] The sauropods of the Huincul Formation are represented by the titanosaurs Argentinosaurus and Choconsaurus , [27] and several rebbachisaurids including Cathartesaura , [28] Limaysaurus , [29] [30] and some unnamed species. [21]

See also

References

  1. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 Rolando, Alexis M. Aranciaga; Motta, Matías J.; Agnolín, Federico L.; Tsuihiji, Takanobu; Miner, Santiago; Brissón-Egli, Federico; Novas, Fernando E. (9 October 2024). "A new carcharodontosaurid specimen sheds light on the anatomy of South American giant predatory dinosaurs". The Science of Nature. 111 (6): 56. Bibcode:2024SciNa.111...56R. doi:10.1007/s00114-024-01942-4. ISSN   1432-1904. PMID   39382666. S2CID   273199114.
  2. 1 2 3 4 5 6 7 8 9 10 Motta, Matías J.; Aranciaga Rolando, Alexis M.; Rozadilla, Sebastián; Agnolín, Federico E.; Chimento, Nicolás R.; Egli, Federico Brissón; Novas, Fernando E. (June 2016). "New theropod fauna from the Upper Cretaceous (Huincul Formation) of northwestern Patagonia, Argentina". New Mexico Museum of Natural History and Science Bulletin. 71: 231–253 – via ResearchGate.
  3. 1 2 Coria, R. A., & Currie, P. J. (2006). A new carcharodontosaurid (Dinosauria, Theropoda) from the Upper Cretaceous of Argentina. Geodiversitas, 28(1), 71-118.
  4. ↑ Canale, Juan Ignacio; Novas, Fernando Emilio; Salgado, Leonardo; Coria, Rodolfo Aníbal (1 December 2015). "Cranial ontogenetic variation in Mapusaurus roseae (Dinosauria: Theropoda) and the probable role of heterochrony in carcharodontosaurid evolution". Paläontologische Zeitschrift. 89 (4): 983–993. Bibcode:2015PalZ...89..983C. doi:10.1007/s12542-014-0251-3. hdl:11336/19258. ISSN   1867-6812.
  5. 1 2 3 4 Canale, Juan I.; Apesteguía, Sebastián; Gallina, Pablo A.; Mitchell, Jonathan; Smith, Nathan D.; Cullen, Thomas M.; Shinya, Akiko; Haluza, Alejandro; Gianechini, Federico A.; Makovicky, Peter J. (July 2022). "New giant carnivorous dinosaur reveals convergent evolutionary trends in theropod arm reduction". Current Biology. 32 (14): 3195–3202.e5. Bibcode:2022CBio...32E3195C. doi: 10.1016/j.cub.2022.05.057 . PMID   35803271. S2CID   250343124.
  6. 1 2 3 4 Coria, Rodolfo A.; Currie, Philip J.; Ortega, Francisco; Baiano, Mattia A. (2019). "An Early Cretaceous, medium-sized carcharodontosaurid theropod (Dinosauria, Saurischia) from the Mulichinco Formation (upper Valanginian), Neuquén Province, Patagonia, Argentina". Cretaceous Research . 111: 104319. doi:10.1016/j.cretres.2019.104319.
  7. ↑ "The Theropod Database". theropoddatabase.github.io. Retrieved 6 March 2025.
  8. ↑ Campione, Nicolás E.; Evans, David C.; Brown, Caleb M.; Carrano, Matthew T. (4 July 2014). Revell, Liam (ed.). "Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions". Methods in Ecology and Evolution. 5 (9): 913–923. Bibcode:2014MEcEv...5..913C. doi:10.1111/2041-210X.12226. ISSN   2041-210X.
  9. ↑ Persons IV, W. Scott; Currie, Philip J.; Erickson, Gregory M. (2020). "An Older and Exceptionally Large Adult Specimen of". The Anatomical Record. 303 (4): 656–672. doi:10.1002/ar.24118. ISSN   1932-8494. PMID   30897281.
  10. ↑ Hutchinson, John R.; Bates, Karl T.; Molnar, Julia; Allen, Vivian; Makovicky, Peter J. (12 October 2011). "A Computational Analysis of Limb and Body Dimensions in Tyrannosaurus rex with Implications for Locomotion, Ontogeny, and Growth". PLOS ONE. 6 (10): e26037. Bibcode:2011PLoSO...626037H. doi: 10.1371/journal.pone.0026037 . ISSN   1932-6203. PMC   3192160 . PMID   22022500.
  11. ↑ Sereno, Paul C; Myhrvold, Nathan; Henderson, Donald M; Fish, Frank E; Vidal, Daniel; Baumgart, Stephanie L; Keillor, Tyler M; Formoso, Kiersten K; Conroy, Lauren L (30 November 2022). Zhu, Min; Rutz, Christian; Zhu, Min; Holtz, Thomas R; Hone, David (eds.). "Spinosaurus is not an aquatic dinosaur". eLife. 11: e80092. doi: 10.7554/eLife.80092 . ISSN   2050-084X. PMC   9711522 . PMID   36448670.
  12. ↑ Ibrahim, Nizar; Sereno, Paul C.; Dal Sasso, Cristiano; Maganuco, Simone; Fabbri, Matteo; Martill, David M.; Zouhri, Samir; Myhrvold, Nathan; Iurino, Dawid A. (26 September 2014). "Semiaquatic adaptations in a giant predatory dinosaur". Science. 345 (6204): 1613–1616. Bibcode:2014Sci...345.1613I. doi:10.1126/science.1258750. PMID   25213375.
  13. ↑ Farlow, J. O., Brinkman, D. L., Abler, W. L., & Currie, P. J. (1991). Size, shape, and serration density of theropod dinosaur lateral teeth. Modern Geology, 16(1-2), 161-198.
  14. ↑ Brusatte, Stephen L.; Benson, Roger B. J.; Carr, Thomas D.; Williamson, Thomas E.; Sereno, Paul C. (12 December 2007). "The systematic utility of theropod enamel wrinkles". Journal of Vertebrate Paleontology. 27 (4): 1052. doi:10.1671/0272-4634(2007)27[1052:tsuote]2.0.co;2.
  15. ↑ Harrid, Jerald David (1998). A reanalysis of Acrocanthosaurus atokensis, its phylogenetic status, and paleobiogeographic implications, based on a new specimen from Texas. New Mexico Museum of Natural History and Science (published 1 January 1998).
  16. 1 2 Leanza, H.A; Apesteguı́a, S.; Novas, F.E; de la Fuente, M.S. (1 February 2004). "Cretaceous terrestrial beds from the Neuquén Basin (Argentina) and their tetrapod assemblages". Cretaceous Research. 25 (1): 61–87. Bibcode:2004CrRes..25...61L. doi:10.1016/j.cretres.2003.10.005. ISSN   0195-6671.
  17. ↑ Corbella, H.; Novas, F.E.; Apesteguía, S.; Leanza, H. (2004). "First fission-track age for the dinosaur-bearing Neuquén Group (Upper Cretaceous), Neuquén Basin, Argentina". Revista del Museo Argentino de Ciencias Naturales. Nueva Serie. 6 (21): 227=232. doi: 10.22179/REVMACN.6.84 .
  18. ↑ Rainoldi, A.L.; Franchini, Marta; Beaufort, D.; Mozley, P.; Giusiano, A.; Nora, C.; Patrier, P.; Impiccini, A.; Pons, J. (2015). "Mineral reactions associated with hydrocarbon paleomigration in the Huincul High, Neuquén Basin, Argentina". GSA Bulletin. 127 (11–12): 1711–1729. Bibcode:2015GSAB..127.1711R. doi:10.1130/B31201.1. hdl: 11336/36686 .
  19. ↑ Vallati, P. (2001). "Middle cretaceous microflora from the Huincul Formation ("Dinosaurian Beds") in the Neuquén Basin, Patagonia, Argentina". Palynology. 25 (1): 179–197. Bibcode:2001Paly...25..179V. doi:10.2113/0250179.
  20. ↑ Motta, M.J.; Brissón Egli, F.; Aranciaga Rolando, A.M.; Rozadilla, S.; Gentil, A. R.; Lio, G.; Cerroni, M.; Garcia Marsà, J.; Agnolín, F. L.; D'Angelo, J. S.; Álvarez-Herrera, G. P.; Alsina, C.H.; Novas, F.E. (2019). "New vertebrate remains from the Huincul Formation (Cenomanian–Turonian;Upper Cretaceous) in Río Negro, Argentina". Publicación Electrónica de la Asociación Paleontológica Argentina. 19 (1): R26. doi: 10.5710/PEAPA.15.04.2019.295 . hdl: 11336/161858 . S2CID   127726069. Archived from the original on 14 December 2019. Retrieved 14 December 2019.
  21. 1 2 Bellardini, F.; Filippi, L.S. (2018). "New evidence of saurischian dinosaurs from the upper member of the Huincul Formation (Cenomanian) of Neuquén Province, Patagonia, Argentina". Reunión de Comunicaciones de la Asociación Paleontológica Argentina: 10.
  22. ↑ Canale, J.I.; Scanferla, C.A.; Agnolin, F.L.; Novas, F.E. (2009). "New carnivorous dinosaur from the Late Cretaceous of NW Patagonia and the evolution of abelisaurid theropods". Naturwissenschaften. 96 (3): 409–14. Bibcode:2009NW.....96..409C. doi:10.1007/s00114-008-0487-4. hdl: 11336/52024 . PMID   19057888. S2CID   23619863.
  23. ↑ Cerroni, M.A.; Motta, M.J.; Agnolín, F.L.; Aranciaga Rolando, A.M.; Brissón Egliab, F.; Novas, F.E. (2020). "A new abelisaurid from the Huincul Formation (Cenomanian-Turonian; Upper Cretaceous) of Río Negro province, Argentina". Journal of South American Earth Sciences. 98: 102445. Bibcode:2020JSAES..9802445C. doi:10.1016/j.jsames.2019.102445. S2CID   213781725.
  24. ↑ Baiano, M.A.; Coria, R.A.; Cau, A. (2020). "A new abelisauroid (Dinosauria: Theropoda) from the Huincul Formation (lower Upper Cretaceous, Neuquén Basin) of Patagonia, Argentina". Cretaceous Research. 110: 104408. Bibcode:2020CrRes.11004408B. doi:10.1016/j.cretres.2020.104408. S2CID   214118853.
  25. ↑ Matías J. Motta; Federico L. Agnolín; Federico Brissón Egli; Fernando E. Novas (2020). "New theropod dinosaur from the Upper Cretaceous of Patagonia sheds light on the paravian radiation in Gondwana". The Science of Nature. 107 (3): Article number 24. Bibcode:2020SciNa.107...24M. doi:10.1007/s00114-020-01682-1. hdl: 11336/135530 . PMID   32468191. S2CID   218913199.
  26. ↑ Apesteguía, S.; Smith, N.D.; Juárez Valieri, R.; Makovicky, P.J. (2016). "An unusual new theropod with a didactyl manus from the Upper Cretaceous of Patagonia, Argentina". PLOS ONE. 11 (7): e0157793. Bibcode:2016PLoSO..1157793A. doi: 10.1371/journal.pone.0157793 . PMC   4943716 . PMID   27410683.
  27. ↑ Simón, E.; Salgado, L.; Calvo, J.O. (2017). "A new titanosaur sauropod from the Upper Cretaceous of Patagonia, Neuquén Province, Argentina". Ameghiniana. 55 (1): 1–29. doi:10.5710/AMGH.01.08.2017.3051. hdl: 11336/89326 . S2CID   134332465.
  28. ↑ de Jesus Faria, C.C.; Riga, B.G.; dos Anjos Candeiro, C.R.; da Silva Marinho, T.; David, L.O.; Simbras, F.M.; Castanho, R.B.; Muniz, F.P.; Gomes da Costa Pereira, P.V.L. (1 August 2015). "Cretaceous sauropod diversity and taxonomic succession in South America". Journal of South American Earth Sciences. 61: 154–163. Bibcode:2015JSAES..61..154D. doi:10.1016/j.jsames.2014.11.008. hdl: 11336/37899 . ISSN   0895-9811.
  29. ↑ Calvo, J.O.; Salgado, L. (1995). "Rebbachisaurus tessonei sp. nov. a new Sauropoda from the Albian-Cenomanian of Argentina; new evidence on the origin of the Diplodocidae". Gaia. 11: 13–33.
  30. ↑ Salgado, L.; Garrido, A.; Cocca, S.E.; Cocca, J.R. (2004). "Lower Cretaceous rebbachisaurid sauropods from Cerro Aguada del León (Lohan Cura Formation), Neuquén Province, northwestern Patagonia, Argentina". Journal of Vertebrate Paleontology. 24 (4): 903–912. doi:10.1671/0272-4634(2004)024[0903:lcrsfc]2.0.co;2. S2CID   129233849.
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