Quilmesaurus

Quilmesaurus; Temporal range: ; Late Cretaceous, 75–66 Ma PreꞒ Ꞓ O S D C P T J K Pg N
	The tibia of Quilmesaurus (A-B) compared to that of other abelisaurids
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Dinosauria
Clade:	Saurischia
Clade:	Theropoda
Family:	† Abelisauridae
Clade:	† Furileusauria
Tribe:	† Carnotaurini
Genus:	† Quilmesaurus ; Coria, 2001
Type species
	†Quilmesaurus curriei; Coria, 2001

Last updated August 15, 2023

Quilmesaurus is a genus of carnivorous abelisaurid theropod dinosaur from the Patagonian Upper Cretaceous (Campanian stage) of Argentina. It was a member of Abelisauridae, closely related to genera such as Carnotaurus.^[1] The only known remains of this genus are leg bones which share certain similarities to a variety of abelisaurids. However, these bones lack unique features, which may render Quilmesaurus a nomen vanum (more commonly known as a nomen dubium , or "dubious name").^[2]

Discovery and naming

During the late 1980s, a field crew from the Universidad Nacional Tucumán, led by Jaime Powell, uncovered forty kilometres south of Roca City, in Río Negro province, southern Argentina, the remains of a theropod near the Salitral Ojo de Agua. In 2001, Rodolfo Aníbal Coria named and described the type species Quilmesaurus curriei. The genus name is derived from the Quilme, a Native American people, and the specific name honours Dr. Philip John Currie, a Canadian theropod specialist.^[3]

The holotype and currently only specimen was designated the collection number MPCA-PV-100, in the Museo Provincial "Carlos Ameghino". It consists of the distal (lower or outermost) half of the right femur (thighbone), and a complete right tibia (inner shinbone), collected from the Allen Formation of the Malargüe Group in the Neuquén Basin. These deposits date from the Campanian to Maastrichtian. The specimen came from the fluvial sandstones at the bottom of the Allen Formation. The taxon is notable as it represents one of the youngest records of a non-avian theropod from Patagonia.^[3]

Description

The preserved portion of femur is robust and boxy in shape. The rear face of the tip of the bone possessed prominent condyles (joint bumps) for connecting to the tibia (on the inner face of the leg) and fibula (on the outer face of the leg). The lateral condyle (which connected to the fibula) is slightly lower from front-to-back compared to the medial condyle (which connected to the tibia), but it is also wider from side-to-side. An additional finger-like bone spur (an epicondyle) would have also been present on the lateral condyle, although this spur is broken off in the only known Quilmesaurus femur. Just above the medial condyle is a low yet noticeable ridge which juts away from the rest of the bone, towards the midline of the animal's body. This ridge is known as a mesiodistal crest. The area immediately above the condyles possesses a shallow yet wide lowered area known as an extensor groove. Overall the femur is almost identical to that of other abelisaurids.^[2]

The proximal (upper or innermost) part of the tibia possesses a myriad of complex features. A large and hatchet-shaped structure known as a cnemial crest points forwards at the proximal portion of the tibia. The tip of the cnemial crest is hooked due to the presence of a downward pointing spur, known as a ventral process. Although Coria (2001) considered a hooked cnemial crest to be unique to Quilmesaurus,^[3] Valieri et al. (2007) noted that this structure was also possessed by Aucasaurus and Majungasaurus , as well as the ambiguous abelisaurid Genusaurus . The distal part of the tibia possesses its own projections for connecting to ankle bones, known as malleoli. This part has the form of an asymmetrical triangle when seen from the front, with the massive lateral malleolus projecting further distally than the smaller medial malleolus. This combination of distal tibia features was also once presumed to have been unique to Quilmesaurus. However, Valieri et al. (2007) note that the distal tibia of Rajasaurus was very similar to that of Quilmesaurus.^[2]

In 2016, Quilmesaurus was estimated to have measured 5.3 metres (17 ft) in length. This would have made it among the smallest derived abelisaurids, although its legs were proportionally robust like those of Pycnonemosaurus , one of the largest members of the family.^[4]

Classification

When originally described, Coria could not find a more precise placement for Quilmesaurus than Theropoda.^[3] The presence of a notch in the distal articular surface of the tibia was cited by him as evidence of a possible relationship with basal Tetanurae, which would be surprising as Quilmesaurus lived during a time when South American theropod assemblages were dominated by abelisaurids and carcharodontosaurs. Other theropod material has been recovered from within these same strata and has in 2005 also provisionally been referred to the Tetanurae.^[5] However, in a 2004 abstract (and later a 2007 full paper), Rubén Juárez Valieri et al. concluded that Quilmesaurus, in view of the hatchet-shaped cnemial crest, was a member of the Abelisauridae.^[6]^[2]

Unlike members of Megalosauroidea, the tibia of Quilmesaurus does not possess a noticeable anteromedial buttress, and instead it includes a large cnemial crest. Quilmesaurus is also not a coelurosaur due to the distal part of the tibia being asymmetrical in shape as well as having a socket for the astragalus which is lower than that of coelurosaurs. Finally, the shallow and wide (rather than deep and thin) extensor groove excludes Quilmesaurus from Carnosauria, as does the possession of parallel upper and lower edges of the cnemial crest.^[2]

However, some features do support its placement within Ceratosauria. These include a pronounced cnemial crest of the tibia and large mesiodistal crest of the femur. The asymmetrical distal part of the tibia and small socket for the astragalus specifically place it within the family Abelisauridae. The preserved bones share features with various abelisaurid taxa throughout the family, although such similarities are widespread and seemingly pop up at random among the taxa, thus making more specific placement difficult. The hook-like shape of the cnemial crest suggests that Quilmesaurus was a member of the subfamily Carnotaurinae, which Sereno (1998) defined to include all abelisaurids closer to Carnotaurus than to Abelisaurus.^[2]

Size comparison of Quilmesaurus (far right) with other "carnotaurins"

However, the validity of Carnotaurinae has been debated. Although Valieri et al. (2007) considered the subfamily to include taxa such as Majungasaurus, Carnotaurus, Aucasaurus, and Rajasaurus, other studies have found different results. Tortosa et al. (2014) found that Carnotaurinae was an invalid group, as very few abelisaurids could actually apply to the definition set forth by Sereno. According to their analysis, Aucasaurus and Carnotaurus were actually closer to Abelisaurus than they were to Majungasaurus and Rajasaurus, thus forcing the latter two taxa to be excluded from the subfamily. Quilmesaurus was retained as close to Aucasaurus and Carnotaurus, although Sereno's name and definition of Carnotaurinae was completely demolished. In its place the tribe Carnotaurini was used, which includes all abelisaurids descended from the last common ancestor of Aucasaurus and Carnotaurus.^[7] Tortosa et al. (2014)'s result has largely been supported over that of Valieri et al. (2007). Filippi et al. (2016) created a new clade, Furileusauria, to include abelisaurids more closely related to Carnotaurus than to Ilokelesia , Skorpiovenator , or Majungasaurus. They included Quilmesaurus among the furileusaurians.^[1]

Valieri et al. (2007) were unable to establish a single autapomorphy (distinctive or unique trait) of the taxon, concluding that Quilmesaurus were a nomen vanum .^[2]

Paleoecology

The Allen Formation is believed to have been a humid coastal environment which gradually transitioned from a freshwater floodplain to marshy estuaries and then shallow lagoons as sea levels rose. A diverse assemblage of aquatic life inhabited the area, including various fish, frogs, and turtles. More recent intervals of the formation even include a few marine reptiles, such as various plesiosaurs including elasmosaurids and polycotylids.^[8] Plant life includes palm trees and conifers of the family Podocarpeaceae ("plum pines"), which formed dense forests and wetlands.^[9]

Remains of land animals were also common in this formation. An indeterminate rhynchocephalian is known, as well as numerous snake taxa including the madtsoiids Patagoniophis and Alamitophis.^[9] Other non-dinosaur animals in the area include the pterosaur Aerotitan ^[10] and a variety of mammals.^[11]

Dinosaur remains recovered from the Allen Formation include a diverse and abundant assortment of titanosaurs ( Saltasaurus, Aeolosaurus, Laplatasaurus, Rocasaurus, etc.) and a hadrosaurid of dubious validity ( Willinakaqe ).^[12] Theropods other than Quilmesaurus were also present; they include the large unenlagiine dromaeosaurid Austroraptor ,^[13] a basal ornithuran bird ( Limenavis ),^[14] and a cimolopterygid bird ( Lamarqueavis ).^[15] A tooth has been referred to the family Carcharodontosauridae; this tooth is one of the most recent carcharodontosaurid fossils found as more well known members of this family ( Giganotosaurus , Mapusaurus ) lived millions of years earlier in the Cretaceous.^[9] Indeterminate nodosaurid remains have also been found at this formation, consisting of vertebrae, osteoderms, a femur, and a tooth.^[16]

The Allen Formation is also notable for the high amount of sauropod eggs discovered there. Nesting grounds have been discovered in the bajo de Santa Rosa area of the upper Allen Formation. Some (but not all) of these eggs were designated as the oogenus Sphaerovum . The structure of their eggshells indicate that they were laid in a very damp environment.^[9]

Related Research Articles

Carnotaurus is a genus of theropod dinosaur that lived in South America during the Late Cretaceous period, probably sometime between 71 and 69 million years ago. The only species is Carnotaurus sastrei. Known from a single well-preserved skeleton, it is one of the best-understood theropods from the Southern Hemisphere. The skeleton, found in 1984, was uncovered in the Chubut Province of Argentina from rocks of the La Colonia Formation. Carnotaurus is a derived member of the Abelisauridae, a group of large theropods that occupied the large predatorial niche in the southern landmasses of Gondwana during the late Cretaceous. Within the Abelisauridae, the genus is often considered a member of the Brachyrostra, a clade of short-snouted forms restricted to South America.

Abelisaurus is a genus of predatory abelisaurid theropod dinosaur alive during the Late Cretaceous Period (Campanian) of what is now South America. It was a bipedal carnivore that probably reached about 7.4 metres in length, although this is uncertain as it is known from only one partial skull.

Abelisauridae is a family of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are found on the modern continents of Africa and South America, as well as on the Indian subcontinent and the island of Madagascar. Isolated teeth were found in the Late Jurassic of Portugal, and the Late Cretaceous genera Tarascosaurus and Arcovenator have been described in France. Abelisaurids first appear in the fossil record of the early middle Jurassic period, and at least three genera survived until the end of the Mesozoic era 66 million years ago.

<i>Aucasaurus</i> Extinct genus of dinosaurs

Aucasaurus is a genus of medium-sized abelisaurid theropod dinosaur from Argentina that lived during the Late Cretaceous of the Anacleto Formation. It was smaller than the related Carnotaurus, although more derived in some ways, such as its extremely reduced arms and almost total lack of fingers. The type skeleton is complete to the thirteenth caudal vertebra, and so is relatively well understood, and is the most complete abelisaurid yet described. However, the skull is damaged, causing some paleontologists to speculate that it was involved in a fight prior to death.

Abelisauroidea is typically regarded as a Cretaceous group, though the earliest abelisaurid remains are known from the Middle Jurassic of Argentina and possibly Madagascar possible abelisauridae remains were also discovered in Late Jurassic Tendaguru Beds in Tanzania. Abelisauroids flourished in the Southern hemisphere during the Cretaceous period, but their origins can be traced back to at least the Middle Jurassic, when they had a more global distribution. By the Cretaceous period, abelisauroids had apparently become extinct in Asia and North America, possibly due to competition from tyrannosauroids. However, advanced abelisauroids of the family Abelisauridae persisted in the southern continents until the Cretaceous–Paleogene extinction event 66 million years ago.

Ilokelesia is an extinct genus of abelisaurid theropod, preserved in the layers of the earliest Late Cretaceous of the Huincul Formation, Neuquén Group, located near Plaza Huincul, Neuquén Province, Argentina. The specimen, consisting of very fragmentary elements of the skull and the axial and appendicular skeleton, was described by Rodolfo Coria and Leonardo Salgado in late 1998.

Ekrixinatosaurus is a genus of abelisaurid theropod which lived approximately 100 to 97 million years ago during the Late Cretaceous period. Its fossils have been found in Argentina. Only one species is currently recognized, Ekrixinatosaurus novasi, from which the specific name honors of Dr. Fernando Novas for his contributions to the study of abelisaurid theropods, while the genus name refers to the dynamiting of the holotype specimen. It was a large abelisaur, measuring between 6.5 and 8 m in length and weighing 800 kg (1,800 lb).

Genusaurus is a genus of abelisauroid dinosaur from the Early Cretaceous. Its fossils were found in France. Genusaurus is believed to have lived during the Albian stage, around 112-100 million years ago.

Pycnonemosaurus is a genus of carnivorous theropod dinosaur that belonged to the family Abelisauridae. It was found in the Upper Cretaceous red conglomerate sandstones of the "Cabembe Unit", Mato Grosso, Brazil, and it lived about 70 million years ago during the Late Cretaceous.

The Bajo de la Carpa Formation is a geologic formation of the Neuquén Basin that crops out in northern Patagonia, in the provinces of Río Negro and Neuquén, Argentina. It is the oldest of two formations belonging to the Río Colorado Subgroup within the Neuquén Group. Formerly, that subgroup was treated as a formation, and the Bajo de la Carpa Formation was known as the Bajo de la Carpa Member.

Carnotaurinae is a subfamily of the theropod dinosaur family Abelisauridae. It includes the dinosaurs Aucasaurus, Carnotaurus. The group was first proposed by American paleontologist Paul Sereno in 1998, defined as a clade containing all abelisaurids more closely related to Carnotaurus than to Majungasaurus.

Carnotaurini is a tribe of the theropod dinosaur family Abelisauridae from the Late Cretaceous period of Patagonia. It includes the dinosaurs Carnotaurus sastrei; the type species, Aucasaurus garridoi, and Abelisaurus comahuensis. This group was first proposed by paleontologists Rodolfo Coria, Luis Chiappe, and Lowell Dingus in 2002, being defined as a clade containing "Carnotaurus sastrei, Aucasaurus garridoi, their most recent common ancestor, and all of its descendants."

Skorpiovenator is a genus of abelisaurid theropod dinosaur from the Late Cretaceous Huincul Formation of Argentina. It is one of the most complete and informative abelisaurids yet known, described from a nearly complete and articulated skeleton.

Bonapartenykus is a monospecific genus of alvarezsauroid dinosaur from Argentina that lived during the Late Cretaceous (Campanian-Maastrichtian) in what is now the upper Allen Formation of the Río Negro Province. The type and only species, Bonapartenykus ultimus, is known from a nearly articulated but partial skeleton that was found in close association to two incomplete eggs and several clusters of eggshells belonging to the oogenus Arriagadoolithus. Bonapartenykus was named in 2012 by Federico L. Agnolin, Jaime E. Powell, Fernando E. Novas and Martin Kundrát. Bonapartenykus has an estimated length of 2.5 m (8.2 ft) and weight of 72 kg (159 lb), making it the largest member of the clade Alvarezsauroidea.

Arcovenator is an extinct genus of abelisaurid theropod dinosaurs hailing from the Late Cretaceous of France. The type and only described species is Arcovenator escotae.

Majungasaurinae is a subfamily of large carnivorous theropods from the Upper Cretaceous, found in Madagascar, India, and France. It is a subgroup within the theropod family Abelisauridae, a Gondwanan clade known for their thick and often horned skulls and vestigial arms. The two subfamilies of Abelisauridae are Carnotaurinae, best known from the South American Carnotaurus, and Majungasaurinae, consisting of Madagascar’s Majungasaurus and its closest relatives. Their ancestors emerged in the Middle Jurassic, and the clade lasted until the Upper Cretaceous.

<i>Viavenator</i> Extinct genus of dinosaurs

Viavenator is a genus of carnivorous abelisaurid theropod dinosaur discovered in Argentina's Bajo de la Carpa Formation. It coexisted with the megaraptoran Tratayenia rosalesi.

Bonapartesaurus is an extinct genus of herbivorous ornithopod dinosaur belonging to Hadrosauridae, which lived in the area of modern Argentina during the Campanian and Maastrichtian stages of the Late Cretaceous.

<span class="mw-page-title-main">Furileusauria</span> Clade of abelisaurid theropod dinosaurs

Furileusauria is an extinct clade of derived abelisaurid dinosaurs only known from South American fossil remains. They represent some of the largest members of the Abelisauridae, with an average length of 7.1 ± 2.1 m (23.3 ± 6.9 ft). The clade is defined as the most inclusive clade containing Carnotaurus sastrei but not Ilokelesia aguadagrandensis, Skorpiovenator bustingorryi, or Majungasaurus crenatissimus.

Tralkasaurus is a genus of abelisaurid dinosaur from the Huincul Formation from Río Negro Province in Argentina. The type and only species is Tralkasaurus cuyi, named in 2020 by Mauricio Cerroni and colleagues based on an incomplete skeleton. A medium-sized abelisaurid, Tralkasaurus exhibits a conflicting blend of characteristics found among the early-diverging abelisauroids with others that characterize the highly specialized clade Brachyrostra, and thus its position within the clade is poorly-resolved.

References

1 2 Filippi, Leonardo S.; Méndez, Ariel H.; Juárez Valieri, Rubén D; C.Garrido, Alberto (2016-06-01). "A new brachyrostran with hypertrophied axial structures reveals an unexpected radiation of latest Cretaceous abelisaurids". Cretaceous Research. 61: 209–219. doi:10.1016/j.cretres.2015.12.018. ISSN 0195-6671.
1 2 3 4 5 6 7 Juárez Valieri, R.D.; Fiorelli, L.E.; Cruz, L.E. (2007). "Quilmesaurus curriei Coria, 2001. Su validez taxonómica y relaciones filogenéticas". Revista del Museo Argentino de Ciencias Naturales "Bernardino Rivadavia" – Paleontología. 9 (1): 59–66. doi: 10.22179/revmacn.9.367 .
1 2 3 4 Coria, R.A. (2001). "A new theropod from the Late Cretaceous of Patagonia". In Tanke, Darren H.; Carpenter, Kenneth (eds.). Mesozoic Vertebrate Life. Life of the Past. Indiana University Press. pp. 3–9. ISBN 978-0-253-33907-2.
↑ Grillo, O. N.; Delcourt, R. (2016). "Allometry and body length of abelisauroid theropods: Pycnonemosaurus nevesi is the new king". Cretaceous Research. 69: 71–89. doi:10.1016/j.cretres.2016.09.001.
↑ Coria, R.A. & Salgado, L. 2005. "Last Patagonian theropods". In: Carpenter, K. 2005. The Carnivorous Dinosaurs, Indiana University Press, pp 153-160
↑ Juárez Valieri R.D, Fiorelli L.E. and Cruz, L.E. 2004. "Quilmesaurus curriei Coria, 2001. Su validez taxonómica y relaciones filogenéticas". XX Jornadas Argentinas de Paleontología de Vertebrados (La Plata), Resúmenes, p. 36-37
↑ Tortosa, Thierry; Buffetaut, Eric; Vialle, Nicolas; Dutour, Yves; Turini, Eric; Cheylan, Gilles (2014-01-01). "A new abelisaurid dinosaur from the Late Cretaceous of southern France: Palaeobiogeographical implications". Annales de Paléontologie. 100 (1): 63–86. doi:10.1016/j.annpal.2013.10.003. ISSN 0753-3969.
↑ O'gorman, José Patricio; Salgado, Leonardo; Gasparini, Zulma (2011). "Plesiosaurios de la Formación Allen (Campaniano-Maastrichtiano) en el Área del Salitral de Santa Rosa (Provincia de Río Negro, Argentina)". Ameghiniana. 48 (1): 129–135. doi:10.5710/AMGH.v48i1(308). S2CID 130603286.
1 2 3 4 Martinelli, Agustín; Forasiepi, Analía (2004). "Late Cretaceous vertebrates from bajo de Santa Rosa (Allen Formation), Río Negro province, Argentina, with the description of a new sauropod dinosaur (Titanosauridae)". Revista del Museo Argentino de Ciencias Naturales. Nueva Serie. 6 (2): 257–305. doi: 10.22179/revmacn.6.88 . ISSN 1853-0400.
↑ Novas, Fernando E.; Kundrat, Martin; Agnolín, Federico L.; Ezcurra, Martín D.; Ahlberg, Per Erik; Isasi, Marcelo P.; Arriagada, Alberto; Chafrat, Pablo (November 2012). "A new large pterosaur from the Late Cretaceous of Patagonia". Journal of Vertebrate Paleontology. 32 (6): 1447–1452. doi:10.1080/02724634.2012.703979. ISSN 0272-4634. S2CID 84340520.
↑ Rougier, Guillermo W.; Chornogubsky, Laura; Casadio, Silvio; Paéz Arango, Natalia; Giallombardo, Andres (2009-02-01). "Mammals from the Allen Formation, Late Cretaceous, Argentina". Cretaceous Research. 30 (1): 223–238. doi:10.1016/j.cretres.2008.07.006. hdl: 11336/103413 . ISSN 0195-6671.
↑ Rubén D. Juárez Valieri, José A. Haro, Lucas E. Fiorelli and Jorge O. Calvo (2010). "A new hadrosauroid (Dinosauria: Ornithopoda) from the Allen Formation (Late Cretaceous) of Patagonia, Argentina" (PDF). Revista del Museo Argentino de Ciencias Naturales. New Series. 11 (2): 217–231.{{cite journal}}: CS1 maint: multiple names: authors list (link)
↑ Novas, Fernando E.; Pol, Diego; Canale, Juan I.; Porfiri, Juan D.; Calvo, Jorge O. (2009-03-22). "A bizarre Cretaceous theropod dinosaur from Patagonia and the evolution of Gondwanan dromaeosaurids". Proceedings of the Royal Society of London B: Biological Sciences. 276 (1659): 1101–1107. doi:10.1098/rspb.2008.1554. ISSN 0962-8452. PMC 2679073 . PMID 19129109.
↑ Clarke, Julia A.; Chiappe, Luis M. (27 February 2001). "A new carinate bird from the late Cretaceous of Patagonia (Argentina)" (PDF). American Museum Novitates (3323): 1–23. doi:10.1206/0003-0082(2001)323<0001:ANCBFT>2.0.CO;2. hdl:2246/2940. S2CID 55064310.
↑ Agnolin, Federico L. (29 May 2010). "Un coracoides de ave del Cretácico Superior de Patagonia, Argentina". Studia Geologica Salmanticensia. 46 (2): 99–119. ISSN 0211-8327.
↑ Salgado, Leonardo; Coria, Rodolfo A. (January 1996). "First evidence of an ankylosaur (Dinosauria, Ornithischia) in South America". Ameghiniana. 33 (4): 367–371.

This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.

[:0-1] 1 2 Filippi, Leonardo S.; Méndez, Ariel H.; Juárez Valieri, Rubén D; C.Garrido, Alberto (2016-06-01). "A new brachyrostran with hypertrophied axial structures reveals an unexpected radiation of latest Cretaceous abelisaurids". Cretaceous Research. 61: 209–219. doi:10.1016/j.cretres.2015.12.018. ISSN 0195-6671.

[:1-2] 1 2 3 4 5 6 7 Juárez Valieri, R.D.; Fiorelli, L.E.; Cruz, L.E. (2007). "Quilmesaurus curriei Coria, 2001. Su validez taxonómica y relaciones filogenéticas". Revista del Museo Argentino de Ciencias Naturales "Bernardino Rivadavia" – Paleontología. 9 (1): 59–66. doi: 10.22179/revmacn.9.367 .

[coria2001-3] 1 2 3 4 Coria, R.A. (2001). "A new theropod from the Late Cretaceous of Patagonia". In Tanke, Darren H.; Carpenter, Kenneth (eds.). Mesozoic Vertebrate Life. Life of the Past. Indiana University Press. pp. 3–9. ISBN 978-0-253-33907-2.

[4] Grillo, O. N.; Delcourt, R. (2016). "Allometry and body length of abelisauroid theropods: Pycnonemosaurus nevesi is the new king". Cretaceous Research. 69: 71–89. doi:10.1016/j.cretres.2016.09.001.

[5] Coria, R.A. & Salgado, L. 2005. "Last Patagonian theropods". In: Carpenter, K. 2005. The Carnivorous Dinosaurs, Indiana University Press, pp 153-160

[6] Juárez Valieri R.D, Fiorelli L.E. and Cruz, L.E. 2004. "Quilmesaurus curriei Coria, 2001. Su validez taxonómica y relaciones filogenéticas". XX Jornadas Argentinas de Paleontología de Vertebrados (La Plata), Resúmenes, p. 36-37

[7] Tortosa, Thierry; Buffetaut, Eric; Vialle, Nicolas; Dutour, Yves; Turini, Eric; Cheylan, Gilles (2014-01-01). "A new abelisaurid dinosaur from the Late Cretaceous of southern France: Palaeobiogeographical implications". Annales de Paléontologie. 100 (1): 63–86. doi:10.1016/j.annpal.2013.10.003. ISSN 0753-3969.

[8] O'gorman, José Patricio; Salgado, Leonardo; Gasparini, Zulma (2011). "Plesiosaurios de la Formación Allen (Campaniano-Maastrichtiano) en el Área del Salitral de Santa Rosa (Provincia de Río Negro, Argentina)". Ameghiniana. 48 (1): 129–135. doi:10.5710/AMGH.v48i1(308). S2CID 130603286.

[:2-9] 1 2 3 4 Martinelli, Agustín; Forasiepi, Analía (2004). "Late Cretaceous vertebrates from bajo de Santa Rosa (Allen Formation), Río Negro province, Argentina, with the description of a new sauropod dinosaur (Titanosauridae)". Revista del Museo Argentino de Ciencias Naturales. Nueva Serie. 6 (2): 257–305. doi: 10.22179/revmacn.6.88 . ISSN 1853-0400.

[10] Novas, Fernando E.; Kundrat, Martin; Agnolín, Federico L.; Ezcurra, Martín D.; Ahlberg, Per Erik; Isasi, Marcelo P.; Arriagada, Alberto; Chafrat, Pablo (November 2012). "A new large pterosaur from the Late Cretaceous of Patagonia". Journal of Vertebrate Paleontology. 32 (6): 1447–1452. doi:10.1080/02724634.2012.703979. ISSN 0272-4634. S2CID 84340520.

[11] Rougier, Guillermo W.; Chornogubsky, Laura; Casadio, Silvio; Paéz Arango, Natalia; Giallombardo, Andres (2009-02-01). "Mammals from the Allen Formation, Late Cretaceous, Argentina". Cretaceous Research. 30 (1): 223–238. doi:10.1016/j.cretres.2008.07.006. hdl: 11336/103413 . ISSN 0195-6671.

[Willinakaqe-12] Rubén D. Juárez Valieri, José A. Haro, Lucas E. Fiorelli and Jorge O. Calvo (2010). "A new hadrosauroid (Dinosauria: Ornithopoda) from the Allen Formation (Late Cretaceous) of Patagonia, Argentina" (PDF). Revista del Museo Argentino de Ciencias Naturales. New Series. 11 (2): 217–231.{{cite journal}}: CS1 maint: multiple names: authors list (link)

[13] Novas, Fernando E.; Pol, Diego; Canale, Juan I.; Porfiri, Juan D.; Calvo, Jorge O. (2009-03-22). "A bizarre Cretaceous theropod dinosaur from Patagonia and the evolution of Gondwanan dromaeosaurids". Proceedings of the Royal Society of London B: Biological Sciences. 276 (1659): 1101–1107. doi:10.1098/rspb.2008.1554. ISSN 0962-8452. PMC 2679073 . PMID 19129109.

[14] Clarke, Julia A.; Chiappe, Luis M. (27 February 2001). "A new carinate bird from the late Cretaceous of Patagonia (Argentina)" (PDF). American Museum Novitates (3323): 1–23. doi:10.1206/0003-0082(2001)323<0001:ANCBFT>2.0.CO;2. hdl:2246/2940. S2CID 55064310.

[15] Agnolin, Federico L. (29 May 2010). "Un coracoides de ave del Cretácico Superior de Patagonia, Argentina". Studia Geologica Salmanticensia. 46 (2): 99–119. ISSN 0211-8327.

[16] Salgado, Leonardo; Coria, Rodolfo A. (January 1996). "First evidence of an ankylosaur (Dinosauria, Ornithischia) in South America". Ameghiniana. 33 (4): 367–371.

[1]

[2]

[3]

[4]

[5]

[6]

[7]

[8]

[9]

[10]

[11]

[12]

[13]

[14]

[15]

[16]