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

South American titanosaurs

Discovery of titanosaur sauropods by B. Gonzalez Riga. (photograph, all right reserved)

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).

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