Notocolossus , one of the largest land animal that lived in the Earth Knowledge of titanosaurian pedal structure is critical to  under...

News: the last paper of Mendozasaurus, the huge saurtopod from Mendoza

Osteology of the Late Cretaceous Argentinean sauropod dinosaur Mendozasaurus neguyelap: implications for basal titanosaur relationships

No hay texto alternativo automático disponible.

Mendozasaurus (artwork Bernardo Gonzalez Riga)


The titanosaurian sauropod dinosaur Mendozasaurus neguyelap is represented by several partial skeletons from a single locality within the Coniacian (lower Upper Cretaceous) Sierra Barrosa Formation in the south of Mendoza Province, northern Neuquén Basin, Argentina. A detailed revision of Mendozasaurus, including previously undocumented remains from the holotype site, allows us to more firmly establish its position within Titanosauria, as well as enabling an emended diagnosis of this taxon. Autapomorphies include: (1) middle and posterior cervical vertebrae with tall and transversely expanded neural spines that are wider than the centra, formed laterally by spinodiapophyseal laminae that are not connected with the pre- or postzygapophyses; (2) anterior caudal vertebrae (excluding anteriormost) with ventrolateral ridge-like expansion of prezygapophyses; and (3) humerus with divided lateral distal condyle on anterior surface. New remains demonstrate that the presacral vertebrae of Mendozasaurus were not unusually short anteroposteriorly, with this compression instead resulting from taphonomic crushing. Comparative studies of articulated pedes of other taxa allow us to interpret that the pedal formula of Mendozasaurus was 2-2-2-2-0, based on disarticulated bones that form a right hind foot. Mendozasaurus was incorporated into an expanded version of a titanosauriform-focussed phylogenetic data matrix, along with several other contemporaneous South American titanosaurs. The resultant data matrix comprises 84 taxa scored for 423 characters, and our phylogenetic analysis recovers Mendozasaurus as the most basal member of a diverse Lognkosauria, including Futalognkosaurus and the gigantic titanosaurs ArgentinosaurusNotocolossusPatagotitan and Puertasaurus. Lognkosauria forms a clade with Rinconsauria (Muyelensaurus + Rinconsaurus), with Epachthosaurus and Pitekunsaurus recovered at the base of this grouping. A basal lithostrotian position for this South American clade is well supported, contrasting with some analyses that have placed these taxa outside of Lithostrotia or closer to Saltasauridae. The sister clade to this South American group is composed of an array of near-global taxa and supports the hypothesis that most titanosaurian clades were widespread by the Early–middle Cretaceous.

Some sauropods from North America - Part 1

Part 1. Comments on the paper of Tschopp et al.

Tschopp, E., Mateus, O., Benson, R. A. 2015. A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda focused on different taxa and their diagnoses.  PeerJ 3:e857

This paper is a great contribution for understanding Diplodocid sauropods. Here I give you some comments and invite to read this paper.

Diplodocidae sauropods includes some of the most popular dinosaurs with elongated necks and tails.
The clade is historically important, having provided the first published reconstruction of an entire sauropod skeleton (‘Brontosaurus’ excelsus; Marsh, 1883), the first complete sauropod skull to be described (Diplodocus; Marsh, 1884), and the first mounted sauropod specimen (Apatosaurus AMNH 460; Matthew, 1905). 

A recent paper (Tschopp et al., 2015) is focused on different taxa and their diagnoses. The authors outlines that although new taxa continue to be discovered, the vast majority of diplodocid species were described in the late 1800s and early 1900s. The high rate of early descriptions, particularly during the so-called ‘Bone Wars’ of the late 1800s, resulted also in a large number of species that are now considered invalid, questionable, or synonymous.

This work is detailed and therefore helps to clarify the valid species. Many fossil bones found long time ago and do not have precise references (taphonomic maps). Likewise, some species were based on bones that today do not have diagnostic characters. 

Figure 1Apatosaurus louisae (CM 3018), a diplodocid from the Late Jurassic at Carnegie Museum of Natural History, Pittsburgh, United States of America.  This species was 21–22.8 m long and had a mass of 16.4–22.4 t (16.1–22.0 long tons; 18.1–24.7 short tons). However, a recent paper (Bates et al. 2015) estimates for this specimen at 21.8–38.2 t (21.5–37.6 long tons; 24.0–42.1 short tons), similar in mass to the South American titanosaur Dreadnoughtus (photograph of B. Gonzalez Riga).

Figure 2. Diplodocus carnegeii close to Apatosaurus louisae at Carnegie Museum of Natural History. (photograph of B. Gonzalez Riga).

Figure 3. Barosaurus lentus at American Museum of Natural History, New York. (photograph of B. Gonzalez Riga).

Selected information of Tschopp et al. 2015:

Apatosaurus Marsh, 1877a.
Type species: Apatosaurus ajax Marsh, 1877a.
Referred species: Apatosaurus louisae Holland, 1915a.
Invalid proposed species: Apatosaurus grandis Marsh, 1877a (= Camarasaurus
grandis), A. laticollis Marsh, 1879 (nomen dubium; = A. louisae), A. minimus Mook,
1917 (non-diplodocoid neosauropod), A. alenquerensis Lapparent & Zbyszewski, 1957
(= Lourinhasaurus alenquerensis), A. yahnahpin Filla & Redman, 1994 (= Brontosaurus

Barosaurus (Marsh, 1890).
Type and only species: Barosaurus lentus (Marsh, 1890).
Invalid proposed species: Barosaurus affinis (Marsh, 1899) (nomen dubium),
Barosaurus gracilis (Russell, B´eland & McIntosh, 1980) (nomen nudum).

Brontosaurus Marsh, 1879.
Syn.: Elosaurus Peterson & Gilmore, 1902, Eobrontosaurus Bakker, 1998.
Type species: Brontosaurus excelsus Marsh, 1879.
Referred species: Brontosaurus parvus (Peterson & Gilmore, 1902), Brontosaurus
yahnahpin (Filla & Redman, 1994).
Invalid proposed species: Brontosaurus amplus Marsh, 1881 (=Brontosaurus excelsus).

Diplodocus Marsh, 1878.
Syn. Seismosaurus Gillette, 1991
Type species: Diplodocus carnegii Hatcher, 1901 (suppressing the D. longus Marsh, 1878,
see above).
Referred species: Diplodocus hallorum (Gillette, 1991).
Invalid proposed species: Diplodocus longus Marsh, 1878 (nomen dubium, previous
type species, case to ICZN in preparation to propose D. carnegii as substitute), D.
lacustris Marsh, 1884 (nomen dubium), D. hayi Holland, 1924 (=Galeamopus hayi).

Supersaurus Jensen, 1985.
Syn. Dystylosaurus Jensen, 1985; Ultrasauros Olshevsky, 1991; Dinheirosaurus Bonaparte
& amp;Mateus, 1999.
Type species: Supersaurus vivianae Jensen, 1985.
Referred species: Supersaurus lourinhanensis (Bonaparte&Mateus, 1999).

The caudofemoral musculature on the titanosaurian tail skeleton

Comments about the paper titled:

The influence of caudofemoral musculature on the titanosaurian (Saurischia: Sauropoda) tail skeleton: morphological and phylogenetic implications

Authors: Lucio M. Ibiricu, Matthew C. Lamanna & Kenneth J. Lacovara

Reference: Lucio M. Ibiricu, Matthew C. Lamanna & Kenneth J. Lacovara (2014) The influence of caudofemoral musculature on the titanosaurian (Saurischia: Sauropoda) tail skeleton: morphological and phylogenetic implications, Historical Biology, 26:4, 454-471, DOI: 10.1080/08912963.2013.787069

This is an interesting and smart paper about the anatomy of caudal vertebrae of titanosaurs. The authors reconstruct the morphology and interpret the implications of selected soft-tissues associated with the titanosaurian caudal skeleton. These tissues, especially the M. caudofemoralis longus (CFL), exerted a considerable influence on the anatomy of the caudal vertebrae and haemal arches. 

The authors propose the existence of three general evolutionary states in titanosaurian proximal caudal vertebrae: (1) a protuberance on the lateral aspect of the vertebrae that persists until at least caudal 20 (considered state 0), (2) a primary lateral surface that becomes dorsoventrally narrow distally and is replaced by the secondary lateral surface at approximately caudal 9–12 (considered state 1) and (3) a rim that migrates across the lateral aspect of the centrum, becoming situated on its ventrolateral corner at about caudal 8 (considered state 2). These three states may easily be rephrased as a character statement for use in phylogenetic analysis, as follows: ‘morphology of lateral aspect of proximal and middle caudal vertebrae: transverse process or rudimentary transverse process persists until at least caudal 20 (0); primary lateral surface replaced by secondary lateral surface at approximately caudal 9–12 (1); low rim migrates ventrally across lateral surface of centrum,  terminating on ventrolateral corner at approximately caudal 8 (2).’


The caudal vertebrae of titanosaurs are the most abundant skeletal elements in the fossil record (González Roga, 2011). In the last years, papers on osteology and miology give us new phylogenetic and functional information.  

Reference: Gonzalez Riga BJ. 2011. Paleontologıa y dinosaurios desde America Latina: Proceedings del III Congreso Latinoamericano de Paleontologıa. In: Calvo J, Porfiri J, Gonzalez Riga BJ, Dos Santos D, editors. Paleobiology of South American titanosaurs. Mendoza: Universidad Nacional de Cuyo. p. 125–141.


In this paper, a new and gigantic titanosaur, Dreadnoughtus schrani, from Upper Cretaceous sediments in southern Patagonia (Argentina) is described. Represented by approximately 70% of the postcranial skeleton, plus craniodental remains, Dreadnoughtus is one of the most complete giant titanosaur yet discovered, and provides new insight into the morphology and evolutionary history of these colossal animals. 

Scientific Paper

Lacovara et al. 2014. A gigantic, exceptionally complete titanosaurian sauropod dinosaur from Sourthern Patagonia, Argentina. Scientific Reports 4, article number 6196.

Additional information:


Titanosaur sauropods in Mendoza (Argentina), 71 million year ago. These criatures of 13 m long  produced the Titanopodus footprints (Credit, illustration of B. Gonzalez Riga)

New findings of dinosaur ichnites from Agua del Choique section (Mendoza Province, Argentina) provides ichnological and anatomical information about the Cretaceous sauropods and theropods. Around 330 tracks distributed in six footprint levels were identified in this area, one of most important of South America. Two ichnocenoses are located in different paleoenvironmental contexts. In the Anacleto Formation (early Campanian) around 20 titanosaurian tracks were found in floodplain and ephemeral channel deposits. Herein, one pes track shows three claw impressions and this is congruent to two new titanosaur specimens recently discovered in Mendoza Province that have articulated and complete pedes. In this context, for the first time to titanosaurs, ichnological evidences are supported by skeletal elements. In the Loncoche Formation (late Campanian-early Maastrichtian) titanosaurian tracks of Titanopodus mendozensisare abundant (around 310 tracks) and were produced by titanosaurs that walked in a very wet substrate of tidally dominated deltas related with the first Atlantic transgression for northern Patagonia. In this facies association, three different trydactl tracks indicate the presence of small theropods (1–2 m long), expanding the knowledge about the faunistic components that lived in these marine marginal environments.

The scientific publications are:

González Riga, B.J. and Calvo, J. 2009. A new wide-gauge Sauropod track site from the Late Cretaceous of Mendoza, Neuquén Basin, Argentina. Palaeontology 52(3): 631-640.

Gonzalez Riga, B.J. 2011. Speeds and stance of titanosaur sauropods: analysis of Titanopodus tracks from the Late Cretaceous of Mendoza, Argentina. Anais da Academia Brasileira de Ciências, 83(1): 279-290. 

González Riga, B.J., Ortiz David, L., Tomaselli, B., Candeiro, R., Coria, J.P., and Pramparo, M. 2015. Sauropod and theropod dinosaur tracks from the Upper Cretaceous of Mendoza (Argentina): Trackmakers and anatomical aspects.  Journal of South American Earth Sciences 61: 134-141.


Notocolossus, one of the largest land animal that lived in the Earth

Knowledge of titanosaurian pedal structure is critical to understanding the stance and locomotion of these enormous herbivores and, by extension, gigantic terrestrial vertebrates as a whole. However, completely preserved pedes are extremely rare among Titanosauria, especially as regards the truly giant members of the group. 
With a powerfully-constructed humerus 1.76 m in length, Notocolossus gonzalezparejasi is one of the largest known dinosaurs. Furthermore, the complete pes of the new taxon exhibits a strikingly compact, homogeneous metatarsus—seemingly adapted for bearing extraordinary weight—and truncated unguals, morphologies that are otherwise unknown in Sauropoda. The pes underwent a near-progressive reduction in the number of phalanges along the line to derived titanosaurs, eventually resulting in the reduced hind foot of these sauropods.

Scientific Reference: 
González Riga, B.J., Lamanna, M., Ortiz David, L., Calvo, J., Coria, J. 2016. A gigantic new titanosaurian dinosaur from Argentina and the evolution of the sauropod hind foot. Scientific Reports (Nature) 6: 19165.

Figure 1Researcher team members Dr. Bernardo Gonzalez Riga (Universidad Nacional de Cuyo, CONICET, Mendoza, Argentina, center), Dr. Matthew Lamanna (Carnegie Museum of Natural History, Pittsburgh, USA, right), and Leonardo ortiz David (Universidad Nacional de Cuyo, CONICET, Mendoza, Argentina, left) with the right humerus (upper arm bone) of the gigantic Notocolossus. Credit: Bernardo Gonzalez Riga.

Comments: Researchers have discovered a gigantic new species of dinosaur that is among the largest yet known to science. Named Notocolossus gonzalezparejasi, the new creature provides key information about the hind foot of giant titanosaurs, which are widely regarded as the most massive land animals that have ever existed. Notocolossus was described from fossil bones belonging to the back, tail, forelimb, and pelvis, plus a complete ankle and foot. The paper describing the discovery appears today in Scientific Reports, a freely-accessible journal from the publishers of NatureThe two described fossil skeletons of Notocolossus were unearthed in southern Mendoza Province, Argentina, from rocks laid down late in the Cretaceous Period, roughly 86 million years ago. Both specimens were discovered by the study leader and project director, Argentine paleontologist Dr. Bernardo González Riga of CONICET (the Consejo Nacional de Investigaciones Científicas y Técnicas) and the Laboratorio de Dinosaurios of the Facultad de Ciencias Exactas y Naturales of the Universidad Nacional de Cuyo (UNCUYO) in Mendoza Province. Other members of the research team include Dr. Matt Lamanna of Carnegie Museum of Natural History in Pittsburgh, USA, and three other Argentine paleontologists: Leonardo Ortiz David and Juan Pedro Coria of CONICET-IANIGLA and the Laboratorio de Dinosaurios (UNCUYO), and Dr. Jorge Calvo of the Centro Paleontológico Lago Barreales of the Universidad Nacional del Comahue in Neuquén Province.

Figure 2. Life reconstruction of the new titanosaur Notocolossus with 1.7 m (5 ft 7 in) human silhouette for approximate scale. Credit: author, Bernardo Gonzalez Riga, Universidad Nacional de Cuyo and CONICET.

Figure 3. Humerus (upper arm bone) of Notocolossus. In the picture, paleontologist Bernardo Gonzalez Riga. 

According to Dr. González Riga, “Giant titanosaurs were the heaviest terrestrial creatures that ever lived. But the hind feet of these dinosaurs—which are critical for understanding how they stood and moved—were not completely known until now. Now we have new evidence that helps to solve this mystery.” 

Titanosaurs are an important but puzzling group of dinosaurs. They are a type of sauropod, the huge, long-necked, long-tailed plant-eaters that many people think of when they hear the word “dinosaur.” Comprising more than 60 named species, titanosaurs lived on every continent and ranged in size from the weight of a cow to at least the weight of the heaviest humpback whales. They were the most common large herbivores in the Gondwanan (i.e., Southern Hemisphere) continents during the Cretaceous Period, the third and final time period of the Mesozoic Era, or Age of Dinosaurs. Despite their extraordinary species richness and diversity in body size, many aspects of titanosaur anatomy, evolution, behavior, and ecology are not well understood. This is due largely to the fact that most of these dinosaurs are known from woefully incomplete fossils, a situation that—with only a few exceptions, such as the still-unnamed species unveiled last week at New York’s American Museum of Natural History—is particularly pronounced in giant titanosaurs. Says Dr. Lamanna, “Most of the very biggest titanosaurs are known from just a few bones, which has made it really hard for paleontologists to learn much about them.” Notocolossus is no exception in this regard; nevertheless, evidence suggests that it was among the largest titanosaurs, and therefore one of the heaviest land animals, yet discovered. Although the incompleteness of the skeleton of the new sauropod has prevented scientists from making precise estimates of its size, its humerus, or upper arm bone, is 1.76 m (5 ft 9 in) in length, which is longer than that of any other titanosaur for which this bone is known, including other giants such as Dreadnoughtus, Futalognkosaurus, and Paralititan. It is also the biggest humerus known for any other Cretaceous dinosaur. If, as is likely, the body proportions of Notocolossus were comparable to those of better preserved titanosaurs, the new beast was more than 30 meters in length and may have weighed between 40,000 and 60,000 kg (44–66 short tons, as much as roughly 9–13 zoo elephants put together). The gargantuan size and Argentinean location of the new titanosaur were the inspiration for its genus name, Notocolossus, which translates to “southern giant.” The species name is in honor of Jorge González Parejas, a Mendoza-based lawyer who has made significant contributions to the protection of that region’s paleontological heritage.

The enormous sizes attained by gigantic sauropods have generated a great deal of interest in the biology of these dinosaurs. Notocolossus is the first truly giant titanosaur for which the hind foot skeleton is known in its entirety. Interestingly, its foot shows anatomical peculiarities relative to those of other titanosaurs—such as an exceptionally short and robust, uniform construction—that may well be adaptations for supporting its extraordinary bulk. Moreover, the foot of the new creature contains a small number of bones, underscoring the fact that titanosaurs had the most reduced toes of all sauropods. Why these huge creatures apparently shrunk their toes remains a puzzle, but it stands in stark contrast to the evolutionary trend observed in another group of colossal land animals – the Proboscidea, which includes elephants and their close relatives. Rather than decreasing their number of toe bones, proboscideans actually increased the number of bones in their feet over the course of their evolution. The hind feet of elephants and sauropods show that these creatures evolved different skeletal strategies for supporting their massive bodies. “Now that we have the whole foot of a giant titanosaur, we can learn more about how these dinosaurs were able to carry more weight around than any other land animal in the history of life,” notes Dr. González Riga. “Argentina was truly the land of giants during the Cretaceous – and Notocolossus gives us new evidence on how these giants got so big.”

Figure 4. Prof. Juan Coria working in a caudal vertebra. 

Figure 5. Life reconstruction of the gigantic new titanosaurian dinosaur species Notocolossus gonzalezparejasi in its ~86 million-year-old habitat in southern Mendoza Province, Argentina. Notocolossus is shown threatening a pair of much smaller, carnivorous abelisaurid dinosaurs, which are also known from fossils from the same area. Credit: Taylor Maggiacomo, Carnegie Museum of Natural History and Carnegie Mellon University.


Some news about Notocolossus in the world

Página 12 (Buenos Aires, Argentina): El gigante de las 60 toneladas
La Nación (Buenos Aires, Argentina): Descubren en Mendoza un dinosaurio que sería el más grande del mundo.
Clarín (Buenos Aires, Argentina): Descubren en Mendoza a uno de los dinosaurios más grandes del mundo 
Radio Rivadavia (Buenos Aires, Argentina): En Mendoza presentaron el fósil del dinosaurio Notocolossus un titán de 60 toneladas.

The Spokesman-Review: Fossil discoveries uncover massive dinosaurs
Pittsburgh Post-Gazette (Pittsburgh, USA): Carnegie Museum scientist helps unearth new dinosaur 
El Nuevo Herald (New york, USA): Hallan en Argentina restos de un dinosaurio gigante desconocido
El tiempo (Colombia): Hallan nueva especie de titanosaurio al sur de Argentina
 Panamá América (Panamá): Enorme descubrimiento
20 minutos (España): Descubren en Argentina a uno de los mayores dinosaurios del mundo
Diario Crónica (México): Descubren en Argentina uno de los mayores dinosaurios del mundo.

La Nación (Chile): Hallan en Argentina restos de un dinosaurio gigante desconocido.


Comentarios en español: 

Investigadores han descubierto una nueva especie que se encuentra entre los dinosaurios más grandes conocidos por la ciencia. Nombrado Notocolossus gonzalezparejasi, la nueva criatura proporciona una información “clave” sobre la anatomía de la extremidad trasera de los titanosaurios gigantes, los cuales son considerados generalmente como los animales terrestres más grandes que han existido. Notocolossus fue descrito a partir de huesos fósiles pertenecientes a la espalda, cola, pata delantera y pelvis, sumando además, un pie posterior completo. El artículo que describe este descubrimiento aparece hoy en Scientific Reports, una revista de libre acceso de los editores de Nature

Los huesos fósiles de Notocolossus fueron hallados en el sur de la provincia de Mendoza (Argentina) en rocas de fines del Período Cretácico, cuya antigüedad se estima en 86 millones años. Los ejemplares fósiles fueron descubiertos por el líder del estudio y director del proyecto, el paleontólogo argentino Dr. Bernardo González Riga del CONICET (Consejo Nacional de Investigaciones Científicas y Técnicas), IANIGLA, y Laboratorio de Dinosaurios de la Facultad de Ciencias Exactas y Naturales de la Universidad Nacional de Cuyo (UNCUYO) en la provincia de Mendoza. Otros miembros del equipo de investigación son el paleontólogo norteamericano Dr. Matt Lamanna del Museo Carnegie de Historia Natural de Pittsburgh, Estados Unidos de América, y otros tres paleontólogos argentinos: Leonardo Ortiz David y Juan Coria del CONICET-IANIGLA y el Laboratorio de Dinosaurios de la UNCUYO, y el Dr. Jorge Calvo del Centro Paleontológico Lago Barreales de la Universidad Nacional del Comahue, en la provincia de Neuquén.
Según el Dr. González Riga, “los titanosaurios gigantes fueron las criaturas terrestres más pesadas ​​que han existido. Las extremidades traseras de estos dinosaurios, fundamentales para conocer su tipo de locomoción y modo de soportar el peso, no eran completamente conocidas. Ahora tenemos nuevas evidencias que ayudan a resolver parte de este misterio”.
Los titanosaurios son un grupo numeroso y ciertamente enigmático. Son saurópodos, es decir enormes herbívoros con largo cuello y cola. Representan lo que mucha gente piensa cuando oye la palabra “dinosaurio”. Comprenden más de 60 especies y vivían en todos los continentes. Su peso variaba entre el de una vaca hasta el de una ballena jorobada. Los titanosaurios fueron los herbívoros más abundantes de Gondwana (continentes del Hemisferio Sur) durante el Período Cretácico, el tercer y último período de la Era Mesozoica, o “era de los dinosaurios”. A pesar de su extraordinaria riqueza de especies y la diversidad en el tamaño del cuerpo, no se conocen bien muchos aspectos de su anatomía, evolución, comportamiento y ecología. Esto se debe principalmente al hecho de que la mayoría de estos dinosaurios se conocen a partir de esqueletos incompletos, una situación que, con sólo unas pocas excepciones, es particularmente pronunciada en los titanosaurios gigantes. Dice el Dr. Lamanna, “La mayoría de los grandes titanosaurios son conocidos mediante unos pocos huesos, lo que limita el conocimiento que pueden generan los paleontólogos”.
Notocolossus no es una excepción en este sentido; sin embargo, la evidencia sugiere que fue uno de los animales más pesados ​​que haya sido descubierto en la Tierra. Aunque el carácter incompleto de su esqueleto impide realizar estimaciones precisas de su tamaño, su húmero (hueso del brazo), tiene 1,76 m de longitud, siendo más largo que el de cualquier otro titanosaurios conocido, incluyendo los gigantes Dreadnoughtus, Futalognkosaurus y Paralititan. Si, como es probable, las proporciones corporales de Notocolossus fueron comparables a los de los titanosaurios mejor preservados, la nueva bestia tenía alrededor de 25–28 m de largo y pesaba entre 40 y 60 toneladas, es decir 9 a 13 elefantes juntos. El tamaño descomunal y la localización argentina inspiraron el nombre de este género, Notocolossus, que se traduce como “gigante del sur”. La especie se refiere a Jorge González Parejas, un abogado con sede en Mendoza que ha hecho contribuciones significativas para la protección del patrimonio paleontológico.
El enorme tamaño alcanzado por los saurópodos gigantes ha generado un gran interés para la biología. Notocolossus es el primer titanosaurio verdaderamente gigante que ha preservado uno de sus pies en forma completa. Curiosamente, sus pies muestran características anatómicas diferentes a la de otras especies, tales como metatarsos y falanges excepcionalmente cortas y robustas. Por otra parte, sus pies presentan un reducido número de falanges, menor que la de otros saurópodos. ¿Por qué estas enormes criaturas –aparentemente– encogieron sus pies? Por ahora esto es un enigma, pero está en marcado contraste con la tendencia evolutiva observada en otro grupo de animales terrestres colosales, los elefantes y sus parientes cercanos. Ellos, en lugar de disminuir su número de huesos en los dedos del pie, los aumentaron a lo largo de su evolución. En suma, las patas traseras de elefantes y saurópodos muestran estrategias evolutivas diferentes para sostener sus cuerpos. “Ahora que tenemos todo el pie de un titanosaurio gigante, podemos aprender más acerca de cómo estos dinosaurios fueron capaces de soportar más peso que cualquier otro animal terrestre en la historia de la vida”, señala el Dr. González Riga. “Argentina era verdaderamente la tierra de los gigantes durante el Cretácico y Notocolossus nos da nuevas evidencias de cómo estos gigantes alcanzaron tamaños colosales”.

South American titanosaurs

The sauropod clade Titanosauria is one of the most diverse and interesting dinosaur groups. Their fossils show an amazing size range, and some of them, were the largest animals ever to walk the Earth. Titanosaurs like NotocolossusArgentinosaurus, Futalognkosaurus, and Puertasaurus surpassed lengths of 30 m and masses of 40 tons.

Discovery of an articulated pes (titanosaur sauropod) in the Argentinean Patagonia.
 / Descubrimiento de un pie articulado (saurópodo titanosaurio) en la Patagonia argentina.

Titanosaurs exhibit a worldwide distribution useful for paleobiogeographic studies. They were important terrestrial herbivores during the Jurassic and specially the Cretaceous periods, and their remains have been recovered from all continents. Moreover, their fossil record is important for studying the composition of dinosaur faunas near the mass extinction at the K-T boundary. Another interesting aspect of titanosaurs is their extraordinary diversity, similar to that of the Late Cretaceous North American hadrosaurids. At the time of this writing, more than 48 titanosaur species have been discovered, around one third of all known sauropods. In this context, phylogenetic studies of titanosaurs are important as a basis for further evolutionary and paleobiogeographic interpretations. Likewise, locomotion and growth strategies are topics of great interest.

Cervical vertebra of Mendozasaurus,
a huge titanosaur from Argentina (González Riga, 2005).

Unfortunately, these research topics have been limited by the fossil record, since most titanosaur species, like other sauropods, are represented by incomplete and mostly disarticulated skeletal elements, such as dorsal and caudal vertebrae, and appendicular bones. However, this situation is changing with the discovery of well preserved specimens, such as the titanosaur Rapetosaurus collected in Madagascar (Curry Rogers and Forster, 2001). Several new genera, such as Gobititan (You et al. 2003) and Sonidosaurus (Xu et al., 2006), were discovered in Asia. In Europe, the titanosaurs Aragosaurus (Sanz et al., 1987; Canudo et al., 2001) and Lirainosaurus (Sanz et al., 1999) and the basal somphospondilyan Tastavinsaurus (Canudo et al., 2008) were described. Likewise, during the last decade, news titanosaur genera were discovered in South America; in Argentina and Brazil the record now reaches 40 genera (see Table 1).
Moreover, in Patagonia, extraordinary embryos and eggs (Chiappe et al., 1998, 2001; Salgado et al., 2005a) and exceptionally articulated specimens (Martínez et al., 2004; Calvo et al., 2007a; González Riga et al., 2008) were found.These discoveries show the importance of the South American record for understanding both the phylogeny and paleobiology (ontogenetic stages, behavior and locomotion) of titanosaurs.

Phylogenetic relationships of some titanosaur sauropods, after González Riga et al. (2009).

Cladogram showing the phylogenetic relationships of some titanosaurs (Gonzalez Riga and Ortiz David, 2014)

 Phyogenetic relationships of Eusauropoda showing the new titanosaur Notocolossus (Gonzalez Riga et al. 2016).


Table 1. South American titanosaur species

Unidad litoestratigráfica del hallazgo

Adamantisaurus mezzalirai
Santucci y Bertini 2006
Formación Adamantina

Aeolosaurus rionegrinus
Powell, 1987
Formación Angostura Colorada
 tardío- Maastrichtiano

Aeolosaurus colhuehapensis
Casal, Martínez, Luna, Sciutto y Lamanna 2007
Formación Bajo Barreal
Cenomaniano tardío-Turoniano temprano

Aeolosaurus maximus
Santucci y Campos 2011
Formación Adamantina

Agustinia ligabuei
Bonaparte 1999
Formación Lohan Cura
Apitano tardío-Albiano

Amargatitanis macni
Apesteguía 2007
Formación La Amarga
Barremiano-Aptiano temprano

Andesaurus delgadoi
Calvo y Bonaparte 1991
Formación Cyeleros
Cenomaniano temprano
Antarctosaurus wichmanianus
Huene 1929
Formación Anacleto?
Campaniano temprano

Argentinosaurus huinculensis
Bonaparte y Coria 1993
Formación Huincul
Cenomaniano temprano

Argyrosaurus superbus
Lydekker 1893
Formación Bajo Barreal
Cenomaniano tardío-Turoniano temprano

Atacamatitan chilensis
Kellner, Rubilar-Rogers, Vargas y Suárez 2011
Formación Tolar

Baurutitan britoi
Kellner, Campos y Trotta 2005
Formación Marília

Barrosasaurus casamiquelai
Salgado y Coria 2009
Formación Anacleto
Campaniano temprano


Bonatitan reigi
Martinelli y Forasiepi 2004
Formación Allen
Campaniano tardío-Maastrichtiano temprano

Bonitasaura salgadoi
Apesteguía 2004
Formación Bajo de la Carpa

Chubutisaurus insignis
Del Corro 1975
Formación Cerro Barcino

Drusilasaura deseadensis
Navarrete, Casal, Martínez 2011
Bajo Barreal
Cenomaniano tardío-
Turoniano temprano

Epachthosaurus sciuttoi
Powell, 1990
Formación Bajo Barreal
Cenomaniano temprano-Turoniano tardío

Futalonkosaurus dukei
Calvo, Porfiri, González Riga y Kellner 2007
Formación Portezuelo
Turoniano tardío-Coniaciano temprano

Gondwanatitan faustoi
Kellner y Azevedo 1999
Grupo Bauru

Laplatasaurus araukanicus
Huene 1929
Formación Anacleto?
Campaniano temprano

Ligabuesaurus leanzai
Bonaparte, González Riga, y Apesteguía 2006
Formación Lohan Cura
Aptiano tardío-Albiano

Malarguesaurus florenciae
González Riga, Previtera, y Pirrone 2009
Formación Portezuelo
Turoniano tardío-Coniaciano temprano

Maxakalisaurus topai
Kellner, Campos, Azevedo, Trotta, Henriques, Craik, y Silva 2006
Formación Adamantina

Mendozasaurus neguyelap
González Riga 2003
Formación Portezuelo-Plottier
Turoniano tardío- Coniaciano

Muyelensaurus pecheni
Calvo, González Riga y Porfiri, 2007
Formación Portezuelo
Turoniano tardío-Coniaciano temprano
Narambuenatitan palomoi
Filippi, García y Garrido 2011
Formación Anacleto
Campaniano medio

Neuquensaurus australis
(Lydekker, 1893) Powell, 1986
Formación Anacleto
Campaniano temprano

Panamericansaurus schroederi
Calvo y Porfiri 2010
Formación Allen
Campaniano tardio-Maastrichtiano

Pellegrinisaurus powelli
Salgado 1996
Formación Anacleto
Campaniano temprano

Petrobrasaurus puestohernandezi
Filippi, Canudo, Salgado, Garrido, García, Cerda y Otero 2011

Pitekunsaurus macayai
Filippi y Garrido 2008
Formación Anacleto
Campaniano temprano
 Puertasaurus reuili
Novas, Salgado, Calvo, y Agnolin 2005
Formación Pari Aike
Maastrichtiano temprano

Quetecsaurus rusconii
Gonzalez Riga y Ortiz David, 2014
Formacion Anacleto
Campaniano inferior

Rinconsaurus caudamirus
Calvo y González Riga 2003
Formación Portezuelo-Plottier
Turoniano tardío-Coniaciano

Rocasaurus muniozi
Salgado y Azpilicueta 2000
Formación Allen
Campaniano tardío-Maastrichtiano temprano

Saltasaurus loricatus
Bonaparte y Powell 1980
Formación Lecho

Tapuiasaurus macedoi
Zaher, Pol, Carvalho, Nascimento, Riccomini, Larson, Juarez-Valieri, Pires-Domingues, da Silva, Campos 2011
Formación Quiricó

Traukutitan eocaudata
Juárez Valieri y Calvo 2011
Formación Bajo de la Carpa


Trigonosaurus pricei
Campos, Kellner, Bertini, y Santucci 2005
Formación Marília

Uberabatitan ribeiroi
Salgado y Souza Carvalho 2008
Formación Marília

Table 1. South American taxa (González Riga and Ortiz David, 2014)

Rinconsaurus caudamirus
(life restoration of B. González Riga, all right reserved).

References (in working...I am including new cites)

Allain, R., Taquet, P., Battail, B., Dejax, J., Richir, P., Véran, M., Limon-Duparcmeur, F., Vacant, R., Mateus, O., Sayarath, P., Khenthavong, B. and Phouyavong, S. 1999. Un nouveau genre de dinosaure sauropode de la formation des Grès supérieurs (Aptien-Albien) du Laos. Académie des Sciences de Paris 329: 609-616.

Apesteguía, S. 2004. Bonitasaura salgadoi gen. et sp. nov.: a beaked sauropod from the Late Cretaceous of Patagonia. Naturwissenschaften 91: 493-497.

Apesteguía, S. 2007. The sauropod diversity of the La Amarga Formation (Barremian), Neuquén (Argentina). Gondwana Research 12 (4): 533-546.

Benson RBJ, Butler RJ, Alroy J, et al., 2016, Near-Stasis in the Long-Term Diversification of Mesozoic Tetrapods, PLOS Biology, Vol:14, ISSN:1545-7885.
Benson RBJ, Campione NE, Carrano MT, et al., 2014, Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage,PLOS Biology, Vol:12, ISSN:1545-7885.

Bonaparte, J.F. 1999. An armoured sauropod from the Aptian of Northern Patagonia, Argentina. In: Tomida, T.H. Rich and P. Vickers-Rich (eds.), Proceeding of the Second Gondwanan Dinosaur Symposium, National Science Museum Monographs 15: 1-12

Bonaparte, J.F.; Powell, J.E. 1980. A continental assemblage of tetrapods from the Upper Cretaceous beds of El Brete, northwestern Argentina (Sauropoda, Coelurosauria, Carnosauria, Aves). Mémoires de la Société Geólogique de France 139: 19-28.

Bonaparte, J.F.; Coria, R.A. 1993. Un nuevo y gigantesco saurópodo Titanosaurio de la Formación Río Limay (Albiano-Cenomaniano) de la provincia del Neuquén, Argentina. Ameghiniana 30: 271-282.

Bonaparte, J.F., González Riga, B.J.; Apesteguía, S. 2006. Ligabuesaurus leanzai nov. gen. et sp., a new titanosaur from the Aptian of Patagonia, Argentina. Cretaceous Research 27: 364-376.

Bonnan, M.F. 2005. Pes anatomy in sauropod dinosaurs: implications for functional morphology, evolution, and phylogeny. In: Carpenter, K., Tidwell, V. (eds.), Thunder-Lizards: The Sauropodomorph Dinosaurs. Indiana University Press, Bloomington, pp. 346-380.

Borsuk-Bialyncka, M. 1977. A new camarasaurid sauropod Opisthocoelicaudia skarzynskii, gen.n. sp.n. from the Upper Cretaceous of Mongolia. Palaeontologia Polonica 37: 45-64.

Brusatte SL, Butler RJ, Barrett PM, et al., 2015, The extinction of the dinosaurs, Biological Reviews, Vol:90, ISSN:1464-7931, Pages:628-642
Calvo, J.O. 1991. Huellas de dinosaurios en la Formación Río Limay (Albiano-Cenomaniano?), Picún Leufú, Provincia del Neuquén, República Argentina (Ornithischia-Saurischia: Sauropoda-Theropoda). Ameghiniana 28: 241-258.

Calvo, J.O. 1999. Dinosaurs and other vertebrates of the Ezequiel Ramos Mexía área, Neuquén - Patagonia Argentina.. In: Tomida, Y., Rich, T. and Vickers-Rich, P. (eds). Proceedings of the Second Gondwanan Dinosaur Symposium, National Science Museum Monographs 15: 13-15.

Calvo, J.O., Bonaparte, J.F. 1991. Andesaurus delgadoi n.g.n.sp. (Saurischia, Sauropoda) dinosaurio Titanosauridae de la Formación Río Limay (Albiano - Cenomaniano), Neuquén, Argentina. Ameghiniana 28: 303-310.

Calvo, J.O; González Riga, B.J. 2003. Rinconsaurus caudamirus gen. et sp. nov., a new titanosaurid (Dinosauria, Sauropoda) from the Late Cretaceous of Patagonia, Argentina. Revista Geológica de Chile 30 (2): 333-353.

Calvo, J.O.; Kellner, A.W. 2006. Description of a sauropod dinosaur braincase (Titanosauridae) from the Late Cretaceous Rio Colorado Subgroup, Patagonia. Anais da Academia Brasileira de Ciencias 78(1): 175-182

Calvo, J.O.; Mazzetta, G.V. 2004. Nuevos hallazgos de huellas de dinosaurios en la Formación candeleros (albiano-Cenomaniano), Picún leufú, Neuquén, Argentina. Ameghiniana 41: 545-554.

Calvo, J.O., Coria, R.A., Salgado, L. 1997. Uno de los más completos titanosáuridos (Dinosauria, Sauropoda) registrados en el mundo. 13° Jornadas Argentinas de Paleontología de Vertebrados, Ameghiniana 34(4): 534.

Calvo, J.O., Porfiri, J.D., González Riga, B.J., Kellner, A.W.A. 2007. A new Cretaceous terrestrial ecosystem from Gondwana with the description of a new sauropod dinosaur. Anais da Academia Brasileria de Ciencias 79: 529-541.
Calvo, J.O., González Riga, B.J., and Porfiri, J.A. 2007. A new titanosaur sauropod from the Late Cretaceous of Neuquén, Patagonia, Argentina. Arquivos do Museu Nacional 65(4): 485-504.
Calvo, J.O., Porfiri, J.D., González Riga, B.J. and Kellner, A.W.A. 2007. Anatomy of Futalognkosaurus dukei Calvo, Porfiri, Gonzalez Riga & Kellner, 2007 (Dinosauria, Titanosauridae) from the Neuquén Group (Late Cretaceous), Patagonia, Argentina. Arquivos do Museu Nacional 65(4): 511-526.
Campos, D.A., Kellner, A.W.A., Bertini, R.J. and Santucci, R.M. 2005. On a Titanosauridae (Dinosauria, Sauropoda) vertebral column from the Bauru Group, Late Cretaceous of Brazil. Arquivos do Museu Nacional 63: 565–593.

Canudo, J.I., Barco, J.L., Royo-Torres, R., and Ruiz-Omeñaca, J.I. 2001. Precisiones sobre la posición taxonómica de Aragosaurus ischiaticus (Dinosauria, Sauropoda). XVII Jornadas de la Sociedad Española de Paleontología, Resúmenes: 263–270.

Canudo, J.I., Royo-Torres, R. and Cuenca-Bescós, G. 2008. A new sauropod: Tastavinsaurus sanzi gen. et sp. nov. from the Early Cretaceous (Aptian) of Spain. Journal of Vertebrate Paleontology 28 (3): 712–731.

Carabajal, A. and Salgado, L. 2007. El basicráneo de un titanosaurio (Dinosauria, Sauropoda) del Cretácico Superior del norte de Patagonia: descripción y aportes al conocimiento del oído interno de los dinosaurios. Ameghiniana 44: 109–120.

Carballido, J.L.; D. Pol; I. Cerda y L. Salgado. 2011. The Osteology of Chubutisaurus insignis Del Corro, 1975 (Dinosauria: Neosauropoda) from the ‘middle’ Cretaceous of central Patagonia, Argentina. Journal of Vertebrate Paleontology 31(1): 93–110.

Casal, G., R. Martínez, M. Luna, J. C. Sciutto and M. Lamanna. 2007. Aeolosaurus colhuehuapensis sp. nov. (Sauropoda, Titanosauria) de la Formación Bajo Barreal, Cretácico Superior de Argentina. Revista Brasileira de Paleontologia 10:53-62.

Cerda , I.A.; A. Paulina Carabajal; L. Salgado; R.A. Coria; M. Reguero; C. Tambussi y  J. Moly. 2011. The first record of a sauropod dinosaur from Antarctica. Naturwissenschaften. pp 1 - 5.

Chiappe, L. M., Coria, R. A., Dingus, L., Jackson, F., Chinsamy, A., and Fox, M. 1998. Sauropod dinosaur embryos from the Late Cretaceous of Patagonia. Nature 396: 258–261.

Chiappe, L.M., Salgado, L. and Coria, R.A. 2001. Embryos skulls of titanosaur sauropod dinosaurs. Science 293: 2444-2446.

Chiappe. L. M., Schmitt, J. G., Jackson, F, Garrido, A. Dingus, L., and Grellet-Tinner, G. 2004. Nest structure for sauropods: sedimentary criteria for recognition of dinosaur nesting traces. Palaios 19: 89–95.

Curry Rogers, K. 2005. Titanosauria. In: Curry Rogers, K.A., Wilson, J.A. (eds.), The Sauropods, evolution and paleobiology. University of California Press, Berkeley, pp. 50-103.

Curry Rogers, K. and Ericsson, G.M. 2005. Sauropod histology. In: Curry Rogers, K.A., Wilson, J.A. (Eds.), The Sauropods, evolution and paleobiology. University of California Press, Berkeley, pp. 303-326.

Curry Rogers, K. and Foster, C.A. 2001. The last of the dinosaur titans: a new sauropod from Madagascar. Nature 412: 530-534.

Curry Rogers, K. and Foster, C.A., 2004. The skull of Rapetosaurus krausei (Sauropoda: Titanosauria) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology 24 (1): 121-144.

Curry-Rogers, K. A. 2005. Titanosauria: a phylogenetic overview. pp. 50-103 in K. Curry Rogers and J. Wilson (eds.), The Sauropods: Evolution and Paleobiology, University of California Press, Berkeley and Los Angeles, California.

De La Fuente, M, Salgado, L., Albino, A., Báez, A., Bonaparte, J., Calvo, J., Chiappe, L.,  Codorniú, L., Coria, R., Gasparini, Z., González Riga, B.J., Novas, F., y Pol, D. 2007. Tetrápodos continentales del Cretácico de la Argentina: una síntesis actualizada. Asociación Paleontológica Argentina, Publicación Especial 11 (ISSN 0328-347X), Ameghiniana 50º Aniversario, pp. 137-153.
Del Corro, G. 1975. Un nuevo saurópodo del Cretácico Superior. Chubutisaurus insignis gen. et. sp. nov. (Saurischia-Chubutisauridae) del Cretácico Superior (Chubutiano) Chubut, Argentina. 1º Congreso Argentino de Palentología y Bioestratigrafía, Actas 2: 229-240.

Dingus, L.;  A. Garrido; G.R. Scott;  L.M. Chiappe; J. Clarke y J.G. Schmitt. 2009. The litho-, bio-, and magnetostratigraphy of titanosaurian nesting sites in the Anacleto Formation at Auca Mahuevo (Campanian, Neuquén Province, Argentina). En: L. Barry Albright III (Ed.): Papers on Geology, Vertebrate Paleontology, and Biostratigraphy in Honor of Michael O. Woodburne. Museum of Northern Arizona Bulletin 65: 237-258.

Filippi, L.S. and Garrido, A.C. 2008. Pitekunsaurus macayai gen. et. sp. nov., nuevo titanosaurio (Saurischia, Sauropoda) del Cretácico Superior de la Cuenca Neuquina, Argentina. Ameghiniana 45: 575–590.

Filippi, L.S.; R.A. García y A. Garrido. 2011. A new titanosaur sauropod dinosaur from the Upper Cretaceous of North Patagonia, Argentina. Acta Palaeontologica Polonica 56 (3): 505–520.

Filippi, L.S.; J.I. Canudo; L. Salgado; A. Garrido; R.A. García, I. Cerda y A. Otero. 2011. A new sauropod titanosaur from the plottier formation (upper cretaceous) of Patagonia (argentina). Geologica Acta, vol. 9, Nº 1 - pp. 1-12.

Franco-Rosas, A.C., Salgado, L., Rosas, C.F. and De Souza Carvalho, I. 2004. Nuevos materiales de titanosaurios (Sauropoda) en el Cretácico Superior de Mato Grosso, Brasil. Revista Brasileira de Paleontología 7(3): 329-336.

Gallina, P.; Otero, A. 2015Reassessment of Laplatasaurus araukanicus (Sauropoda: Titanosauria) from the Upper Cretaceous of Patagonia, Argentina.

González Riga, B.J. 1999. Hallazgo de vertebrados fósiles en la Formación Loncoche, Cretácico Superior de la provincia de Mendoza, Argentina. Ameghiniana 36 (4): 401-410.

González Riga, B.J. 2002. Dinosaurs and environments from the Cretaceous of the Province of Mendoza, Argentina. En: D. Trombotto y R. Villalba (editores),  IANIGLA, 30 años de investigación básica y aplicada en ciencias ambientales,  CONICET, 197-202 pp., 

González Riga, B.J. 2003. A new titanosaur (Dinosauria, Sauropoda) from the Upper Cretaceous of Mendoza Province, Argentina. Ameghiniana 40: 155-172.

González Riga, B.J. 2005. Nuevos restos fósiles de Mendozasaurus neguyelap (Sauropoda: Titanosauria) del Cretácico Tardío de Mendoza, Argentina. Ameghiniana 42 (3): 535-538.
González Riga, B.J.; Curry Rogers, K.A. 2006. Osteohistology of the titanosaur Mendozasaurus (Dinosauria, Sauropoda): preliminary interpretations. 9ª Congreso Argentino de Paleontología y Bioestratigrafía, Academia Nacional de Ciencias, Resúmenes: 84.

González Riga, B.J.; Astini, R. 2007. Fossil preservation of large titanosaur sauropods in overbank fluvial facies: a case study in the Cretaceous of Argentina. Journal of South American Earth Sciences 23: 290-303.

González Riga, B.J.; Calvo, J. 2007. Huellas de dinosaurios saurópodos en el Cretácico de Argentina. 4th European Meeting on the Palaeontology and Stratigraphy of Latin America (Madrid, España). Cuadernos del Museo Geominero (ISBN 978-84-7840-707-1), nº 8: 173-179.

González Riga, B.J. and Calvo, J. 2009. A new wide-gauge Sauropod track site from the Late Cretaceous of Mendoza, Neuquén Basin, Argentina. Palaeontology 52 (3): 631-640.

Gonzalez Riga, B.J. and Ortiz David, L. 2014. A new titanosaur (Dinosauria, Sauropoda) from the Upper Cretaceous (Cerro Lisandro Formation) of Mendoza Province, Argentina. Ameghiniana 51 (1):3-25.

González Riga, B.J., Calvo, J.O. and Porfiri, J. 2008.
An articulated titanosaur from Patagonia (Argentina): new evidences of the pedal evolution. Palaeoworld 17: 33-40.
González Riga, B.J., Previtera, E. and Pirrone, C. 2009a. Malarguesaurus florenciae gen. et sp. nov., a new titanosauriform (Dinosauria, Sauropoda) from the Upper Cretaceous of Mendoza, Argentina. Cretaceous Research 30: 135-148.
González Riga, B.J., Pramparo, M., and Calvo, J. 2009b. A mega track site of Mendoza (Argentina): a case study of wide-gauge sauropod trackway from the Late Cretaceous. 10th Mesozoic Terrestrial Ecosystems (Teruel, España), Abstracts.

Gonzalez Riga, B.J., Calvo, J.O. and Previtera, E. 2009c. Análisis tafonómicos de saurópodos titanosaurios en la Cuenca Neuquina (Argentina) y sus implicancaciones en estudios sistemáticos. In: Colectivo Arqueologico y Paleontologico de Salas de los Infantes, IV Jornadas Internacionales sobre Paleontologia de Dinosaurios y su entorno (Burgos, España), Actas: 235-241.

González Riga, B.J., Lamanna, M., Ortiz David, L., Calvo, J., Coria, J. 2016. A gigantic new titanosaurian dinosaur from Argentina and the evolution of the sauropod hind foot. Scientific Reports (Nature) 6: 19165.

Huene, F. 1929. Los Saurisquios y Ornitisquios del Cretácico Argentino. Anales del Museo de La Plata 3 (2), 194 pp.

Jacobs, L.L, Winkler, D.A., Downs W.R. and Gomani, E.M. 1993. New material of an Early Cretaceous titanosaurid sauropod dinosaur from Malawi. Palaeontology 36: 523-534.

Juarez Valieri, R. y J. Calvo. 2011. Revision of MUCPv 204, a Senonian Basal Titanosaur from Northern Patagonia. Paleontología y dinosaurios desde América Latina. EDIUNC. UNC. 143-152.

Kellner, A.W.A. and Azevedo, S.A.K. 1999. A new sauropod dinosaur (Titanosauria) from the Late Cretaceous of Brazil. In: Tomida, Y, Rich, T.H. and Vickers Rich, P. (eds.), Proceedings of the Second Gondwanan Dinosaur Symposium, National Science Museum Monographs 15: 111-142.

Kellner, A.W.A., Campos, D.A. and Trotta, M.N.F. 2005. Description of a titanosaurid caudal series from the Bauru Group, Late Cretaceous of Brazil. Arquivos do Museu Nacional 63: 529–564.

Kellner, A.W.A., Campos, D.A. Azevedo, S.A.K., Trotta, M.N.F., Enriques, D.D.R., Craik, M.M.T. and Silva, H.P. 2006. On a new titanosaur sauropod from the Bauru Group, Late Cretaceous of Brazil. Boletim do Museu Nacional, Geologia 74: 1–31.

Kellner, A.W.A.; D. Rubilar-Rogers; A. Vargas y M. Suárez. 2011. A new titanosaur sauropod from the Atacama Desert, Chile. Anais da Academia Brasileira de Ciências (2011) 83(1): 211-219.

Kurzanov, S.M. and Bannikov, A.F. 1983.
A new sauropod from the Upper Cretaceous of Mongolia. Palaeontological Journal 2: 91–97.

 et al. 2014. A gigantic, exceptionally complete titanosaurian sauropod dinosaur from southern Patagonia, ArgentinaSci. Rep. 4, 6196.

Lockley, M.G., Shulp, A.S., Meyer, C.A., Leonardi, G. and Kerumba Mamani, D. 2002. Titanosaurid trackways from the Upper Cretaceous of Bolivia: evidence for large manus, wide-gauge locomotion and gregarious behaviour. Cretaceous Research 23, 383-400.

Lyddeker, R. 1893. Contributions to the study of the fossil vertebrates of Argentina. I, The dinosaurs of Patagonia. Anales del Museo de La Plata, Paleontología 2: 1-14.

Mannion PD, Benson RBJ, Carrano MT, et al., 2015, Climate constrains the evolutionary history and biodiversity of crocodylians, Nature Communications, Vol:6, ISSN:2041-1723

Mannion PD, Upchurch P, Benson RBJ, 
et al., 2014
, The latitudinal biodiversity gradient through deep time, Trends In Ecology and Evolution, Vol:29, Pages:42-50

Mannion PD, Upchurch P, Barnes RN, et al., 2013, Osteology of the Late Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary history of basal titanosauriforms, Zoological Journal of the Linnean Society, Vol:168, ISSN:0024-4082, Pages:98-206.

Martinelli, A.; C.R. Candeiro;  A.M. Forasiepi; E.I. Vera y A. Carvalho. 2010. Dinosaurios Argentinos y Brasileros: Lista De Especies Válidas. Caminhos de Geografia - Uberlândia. Vol. 11, Nº 33, p. 91 - 119.

Martínez, R. 1998. An articulated skull and neck of Sauropoda (Dinosauria: Saurischia) from the Upper Cretaceous of central Pategonia, Argentina. 58th Annual Meeting of The Society of Vertebrate Paleontology, Abstract: 61A.

Martínez, R., Giménez, O., Rodríguez, J., Luna, M. and Lamanna, M. 2004. An articulated specimen of the basal Titanosaurian (Dinosauria: Saurópoda) Epachthosaurus sciuttoi from the Early Late Cretaceous Bajo Barreal Formation of Chubut Province, Argentina. Journal of Vertebrate Paleontology 24 (1):107-120.

Martínez, R., Vita, J., Lamanna, M. and Ibiricu, L. 2006. A CT scan of a titanosauriform skull (Dinosauria: Sauropoda) from Central Patagonia, Argentina. 66th Annual Meeting of the Society of Vertebrate Paleontology, Abstracts: 96A.

Martinelli, A. and Forasiepi, A.M. 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 6 (2): 257-305

Martin, V. Buffetaut, E. and Suteethorn, V. 1994. A new genus of sauropod dinosaur from the Sao Khua Formation (Late Jurassic to Early Cretaceous) of northeastern Thailand. Académie des Sciences de Paris 319 (II): 1085-1092.

Martin, V. Buffetaut, E. and Suteethorn, V. 1999. Description of the type and referred material of Phuwiangosaurus sirindhornae Martin, Buffetaut and Suteethorn, 1994, a sauropod from the Lower Cretaceous of Thailand. Oryctos 2: 39-91.

Mazzetta, G.V., Christiansen, P., and Farina, R.A. 2004. Giants and bizarres: body size of some southern South American Cretaceous dinosaurs. Historical Biology 2004:1-13.

Mcintosh, J.S. 1990. Sauropoda. In: D. B. Weishampel, P. Dodson, and H. Osmólska (eds.), The Dinosauria, Berkeley, University of California Press, 345–401.
Earth Sciencies Series 11: 59-90. Houston.

Navarrete, C.; G. Casal y R. Martínez. 2011. Drusilasaura deseadensis gen. et sp. nov., un nuevo titanosaurio (Dinosauria-Sauropoda), de la Formación Bajo Barreal, Cretácico superior del norte de Santa Cruz, Argentina. Revista brasilera paleontógica, Sociedade Brasileira de Paleontologia. 14 (1): 1-14.

Novas, F.E., Salgado, L., Calvo, J.O. and Agnolin, F. 2005. Giant titanosaur (Dinosauria, Sauropoda) from de Late Cretaceous of Patagonia. Revista del Museo Argentino de Ciencias Naturales 7 (1): 37-41.

Nowinski, A. 1971. Nemegtosaurus mongoliensis n. gen. n. sp. (Sauropoda) from the uppermost Cretaceous of Mongolia. Palaeontologia Polonica 25: 57-81.

Perez, Leandro M.; Otero, Alejandro; APesteguia, Sebastián  y  Gallina, Pablo A.. Estratigrafía y análisis tafonómico de Bonitasaura salgadoi Apesteguía, en el sitio «La Bonita» (Cretácico superior, Río Negro, Argentina). Rev. Mus. Argent. Cienc. Nat.[online]. 2009, vol.11, n.1 [citado  2016-09-25], pp. 39-48 .

Poropat SF, Mannion PD, Upchurch P, et al., New Australian sauropods shed light on Cretaceous dinosaur palaeobiogeography, Scientific Reports, ISSN:2045-2322

Powell, J.E. 1986. [Revisión de los Titanosáuridos de América del Sur. Tesis Doctoral, Universidad Nacional de Tucumán, República Argentina, 340 pp. Inédita].

Powell, J.E. 1987a. Morfología del esqueleto axial de los dinosaurios titanosáuridos (Saurischia, Sauropoda) del estado de Minas Gerais, Brasil. Anais do 10º Congresso Brasileiro de Paleontología (Río de Janeiro, 1987): 151-171.

Powell, J.E. 1987b. The Late Cretaceous Fauna from Los Alamitos, Patagonia, Argentina. Part. VI. The titanosaurids. Revista Museo Argentino Ciencias Naturales, 3(3):147-153.

Powell, J.E. 1990. Epachtosaurus sciutoi (gen. et sp.nov.) un dinosaurio sauropodo del Cretacico de Patagonia (Provincia de Chubut, Argentina). Actas del 5º Congreso Argentino de Paleontologia y bioestratigrafía : 123-128.

Powell, J.E. 1992. Osteología de Saltasaurus loricatus (Sauropoda-Titanosauridae) del Cretácico Superior del noroeste argentino. In: J.L. Sanz, Buscalioni, A.D. (eds.), Los Dinosaurios y su entorno biótico, Instituto “Juan de Valdés”, Cuenca, pp. 165-230.

Powell, J.E. 2003. Revision of South American titanosaurid dinosaurs: palaeonbiological, palaeobiogeographical and phylogenetic aspects. Records of the Queen Victoria Museum, Launceston 111: 1–173.

Rodríguez, J.F.R. 1993. La depositacion de las areniscas verdes (Formación Bajo Barreal—Cretacico Tardio) y sus implicancias tafonomicas. XII Congreso Geológico Argentino y II Congreso de Exploración de Hidrocarburos, Actas 1:194–199.

Rubilar-Rogers, D. 2003. Registro de dinosaurios en Chile. Boletín del Museo Nacional de Historia Natural 52:137–150.

Rubilar-Rogers, D. 2005. Titanosauriformes remains from Quebrada Cortadera (Tolar Formation, Upper Cretaceous), Atacama Desert, Chile. Libro de resúmenes de las XXI Jornadas Argentinas de Paleontología de Vertebrados, Plaza Huincul, Argentina 38R. 

Rubilar-Rogers, D., Gutstein, C. 2012. Los titanosaurios de Chile y su contexto filogenético y biogeográfico. Bol. Mus. Nac. Hist. Nat. 61: 55-73.

Rubilar-Rogers, D., Otero, R., Yury-Yáñez, R., Vargas, A., Gutstein, C. 2012. An overview of the dinosaurs fossil record from Chile. Journal of South American Earth Sciences 37:242-255. 

Salgado, L. 1996. Pellegrinisaurus powelli nov. gen. et sp. (Sauropoda, Titanosauridae) from the Upper Cretaceous of Lago Pellegrini, northwestern Patagonia, Argentina. Ameghiniana 33: 355-365.

Salgado, L. 2003a. Should we abandon the name Titanosauridae? Some comments on the taxonomy of titanosaurian sauropod (Dinosauria). Revista Española de Paleontología 18: 15-21.

Salgado, L. 2003b. Considerations on the bony plates assigned to titanosaurs (Dinosauria, Sauropoda). Ameghiniana 40: 441-456.

Salgado, L. and Coria, R.A., 1993. El género Aeolosaurus (Sauropoda- Titanosauridae) en la Formación Allen (Campaniano- Maastrichtiano) de la Provincia de Río Negro, Argentina. Ameghiniana 30: 119-128.

Salgado, L. and Calvo, J.O. 1997. Evolution of Titanosaurid Sauropods. II: The cranial evidence. Ameghiniana 34: 33-48.

Salgado, L. and Azpilicueta, C., 2000. Un nuevo saltasaurino (Sauropoda, Titanosauridae) de la provincia de Río Negro (Formación Allen, Cretácico Superior), Patagonia, Argentina. Ameghiniana 37, 259-264.

Salgado L. and Souza Carvalho, I. 2008. Uberabatitan ribeiroi, a new titanosaur from the Marilia Formation (Bauru Group, Upper Cretaceous), Minas Gerais, Brazil. Palaeontology 51 (4): 881–901.

Salgado, L., Coria, R.A. and Calvo, J.O. 1997a. Evolution of Titanosaurid Sauropods. I: Phylogenetic analysis based on the postcraneal evidence. Ameghiniana 34: 3-32.

Salgado, L., Coria, R.A. and Calvo, J.O., 1997b. Presencia del género Aelosaurus (Sauropoda, Titanosauridae) en la Formación Los Alamitos, Cretácico Superior de la Provincia de Río Negro, Argentina. Geociencias 2 (6): 44-49.

Salgado, L., R. Coria, and L. Chiappe. 2005a. Osteology of the sauropod embryos from the Upper Cretaceous of Patagonia. Acta Palaeontologica Polonica 50: 79–92.

Salgado, L., Apesteguía, S. and Heredia, S., 2005b. A new specimen of Neuquensaurus australis, a Late Cretaceous saltasaurine titanosaur from north Patagonia. Journal of Vertebrate Paleontology 25(3): 623-634.

Salgado, L., R.A. García, and J. D. Daza, 2006. Consideraciones sobre las láminas neurales de los dinosaurios saurópodos y su significado morfofuncional. Revista del Museo Argentino de Ciencias Naturales 8(1): 69-79.   

Santucci, R.M. and Bertini, R.J. 2006. A new titanosaur from western Sao Paulo State, Upper Cretaceous Bauru Group, south-east Brazil. Palaeontology 49: 59–66.

Sanz, J.L., Buscalioni, A.D., Casanovas, M.L., and Santafé, J.V. 1987. Dinosaurios del Cretácico Inferior de Galve (Teruel, España). Estudios Geológicos, Volumen Extraordinario Galve-Tremp: 45–64.

Sanz, J.L., Powell, J.E., Le Loeuff, J.L., Martínez, R., and Pereda-Suberbiola, X. 1999. Sauropod remains from the Upper Cretaceous of Laño (Northcentral Spain). Titanosaur phylogenetic relationships. Estudios del Museo de Ciencias Naturales de Alava 14: 235–255.

Sereno, P.C. 1998. A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie Paläontologie Abhandlungen 210: 41-83.

Smith, J.B., Lamanna, M.C., Lacovara, K.J., Dodson, P., Smith, J.R., Poole, J.C., Giegengack, R., and Attia, Y. 2001. A Giant sauropod dinosaur from an Upper Cretaceous mangrove deposit in Egypt. Science 292 (5522): 1704–1706

Soto-Acuña, S. Otero, R., Rubilar-Rogers, D. y Vargas, A. 2015. Arcosaurios no avianos de Chile. En: Vertebrados Fósiles de Chile. D. Rubilar-Rogers, Otero, R., Vargas, y Sallaberry, M. pp: 209-263.

Tidwell, V., Carpenter, K., Brooks, and B. 1999. New sauropod from the Lower Cretaceous of Utah, USA. Oryctos 2: 21-37.

Tidwell, V., Carpenter, K., and Meyer, S. 2001. New Titanosariform (Sauropoda) from the Poison Strip Member of the Cedar Mountain Formation (Lower Cretaceous), Utah. In: D. Tanke, Carpenter, K (Eds.), Mesozoic Vertebrate Life. Indiana University Press, Bloomington, pp. 139-165.

Tidwell, V. and Carpenter, K. 2003. Braincase of an Early Cretaceous Titanosauriform Sauropod from Texas. Journal of Vertebrate Paleontology 23 (1): 176-180.

Upchurch, P. 1999. The phylogenetic relationships of the Nemegtosauridae (Saurischia, Sauropoda). Journal of Vertebrate Paleontology 19: 106-125.

Upchurch, P., Barrett, P.M., and Dodson, P. 2004. Sauropoda. In:Weishampel, D. B., Dodson, P., and Osmolska, H. (eds.): TheDinosauria (second edition). Berkeley: University of California Press, 259–322.

Wedel, M.J. 2003a. Vertebral pneumaticity, air sacs, and the physiology of sauropods dinosaurs. Paleobiology 29(2): 243-255.

Wedel, M.J. 2003b. The evolution of vertebral pneumaticity in sauropods dinosaurs. Journal of Vertebrate Paleontology 23(2): 344-357.

Wedel, M.J. 2005. Postcranial Skeletal Pneumaticity in Sauropods and Its Implications for Mass Estimates. In: Curry Rogers, K.A., Wilson, J.A. (Eds.), The Sauropods, evolution and paleobiology. University of California Press, Berkeley, pp. 303-326.

Weishampel, D.B., Dodson, P., and Osmólska, H. (eds.). 2004. The Dinosauria, 2nd. Edition. University of California Press, Berkeley, California, 861 pp.

Wilson, J.A. 2002. Sauropod dinosaur phylogeny: critique and cladistic analysis. Zoological Journal of the Linnean Society 136: 217-276.

Wilson, J.A. 2005. Redescription of the Mongolian sauropod Nemegtosaurus mongoliensis Nowinski (Dinosauria: Saurischia) and comments on Late Cretaceous sauropod diversity. Journal of Systematic Palaeontology 3: 283–318.

Wilson, J.A. 2006a. An Overview of Titanosaur Evolution and Phylogeny. En: Colectivo Arqueológico-Paleontológico, Actas de las III Jornadas sobre Dinosaurios y su Entorno: 169-190.

Wilson, J.A. 2006b. Anatomical nomenclature of fossil vertebrates: standardized terms or ‘lingua franca’. Journal of Vertebrate Paleontology 26(3): 511-518.

Wilson, J.A., and Carrano, M.T. 1999. Titanosaurs and the origin of “wide-gauge” trackways: a biomechanical and systematic perspective on sauropod locomotion. Paleobiology 25 (2): 252-267.

Wilson, J.A., and Upchurch, P. 2003. A revision of Titanosaurus Lydekker (Dinosauria - Sauropoda), the first dinosaur genus with a 'Gondwanan' distribution. Journal of Systematic Palaeontology 1 (3): 125-160.

Xu, X., Zhang, X., Tan, Q., Zhao, X, and Tan, L. 2006. A New Titanosaurian Sauropod from Late Cretaceous of Nei Mongol, China. Acta Geologica Sinica 80 (1): 20-26.

You H., Tang F. and Luo Z. 2003. A new basal titanosaur (Dinosauria: Sauropoda) from the Early Cretaceous of China. Acta Geologica Sinica (English edition) 77: 424–429.

Zaher H, Pol D, Carvalho AB, Nascimento PM, Riccomini C, et al. 2011. A Complete Skull of an Early Cretaceous Sauropod and the Evolution of Advanced Titanosaurians. PLoS ONE 6(2): e16663. doi:10.1371/journal.pone.0016663.