South American titanosaurs

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

Cladogram showing the 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)

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

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Table 1. South American titanosaur species

	Especies	Autor-es	Unidad litoestratigráfica del hallazgo	País	Edad
1	Adamantisaurus mezzalirai	Santucci y Bertini 2006	Formación Adamantina	Brasil	Campaniano-Maastrichtiano?
2	Aeolosaurus rionegrinus	Powell, 1987	Formación Angostura Colorada	Argentina	Campaniano  tardío- Maastrichtiano
3	Aeolosaurus colhuehapensis	Casal, Martínez, Luna, Sciutto y Lamanna 2007	Formación Bajo Barreal	Argentina	Cenomaniano tardío-Turoniano temprano
4	Aeolosaurus maximus	Santucci y Campos 2011	Formación Adamantina	Brasil	Campaniano-Maastrichtiano
	Agustinia ligabuei	Bonaparte 1999	Formación Lohan Cura	Argentina	Apitano tardío-Albiano
5	Amargatitanis macni	Apesteguía 2007	Formación La Amarga	Argentina	Barremiano-Aptiano temprano
6	Andesaurus delgadoi	Calvo y Bonaparte 1991	Formación Cyeleros	Argentina	Cenomaniano temprano
7	Antarctosaurus wichmanianus	Huene 1929	Formación Anacleto?	Argentina	Campaniano temprano
8	Argentinosaurus huinculensis	Bonaparte y Coria 1993	Formación Huincul	Argentina	Cenomaniano temprano
9	Argyrosaurus superbus	Lydekker 1893	Formación Bajo Barreal	Argentina	Cenomaniano tardío-Turoniano temprano
10	Atacamatitan chilensis	Kellner, Rubilar-Rogers, Vargas y Suárez 2011	Formación Tolar	Chile	Maastrichtiano
11	Baurutitan britoi	Kellner, Campos y Trotta 2005	Formación Marília	Brasil	Maastrichtiano
12	Barrosasaurus casamiquelai	Salgado y Coria 2009	Formación Anacleto	Argentina	Campaniano temprano
13	Bonatitan reigi	Martinelli y Forasiepi 2004	Formación Allen	Argentina	Campaniano tardío-Maastrichtiano temprano
14	Bonitasaura salgadoi	Apesteguía 2004	Formación Bajo de la Carpa	Argentina	Santoniano
15	Chubutisaurus insignis	Del Corro 1975	Formación Cerro Barcino	Argentina	Aptiano-Albiano
16	Drusilasaura deseadensis	Navarrete, Casal, Martínez 2011	Formación Bajo Barreal	Argentina	Cenomaniano tardío- Turoniano temprano
17	Epachthosaurus sciuttoi	Powell, 1990	Formación Bajo Barreal	Argentina	Cenomaniano temprano-Turoniano tardío
18	Futalonkosaurus dukei	Calvo, Porfiri, González Riga y Kellner 2007	Formación Portezuelo	Argentina	Turoniano tardío-Coniaciano temprano
19	Gondwanatitan faustoi	Kellner y Azevedo 1999	Grupo Bauru	Brasil	Santoniano-Maastrichthiano
20	Laplatasaurus araukanicus	Huene 1929	Formación Anacleto?	Argentina	Campaniano temprano
21	Ligabuesaurus leanzai	Bonaparte, González Riga, y Apesteguía 2006	Formación Lohan Cura	Argentina	Aptiano tardío-Albiano
22	Malarguesaurus florenciae	González Riga, Previtera, y Pirrone 2009	Formación Portezuelo	Argentina	Turoniano tardío-Coniaciano temprano
23	Maxakalisaurus topai	Kellner, Campos, Azevedo, Trotta, Henriques, Craik, y Silva 2006	Formación Adamantina	Brasil	Campaniano-Maastrichtiano?
24	Mendozasaurus neguyelap	González Riga 2003	Formación Portezuelo-Plottier	Argentina	Turoniano tardío- Coniaciano
25	Muyelensaurus pecheni	Calvo, González Riga y Porfiri, 2007	Formación Portezuelo	Argentina	Turoniano tardío-Coniaciano temprano
26	Narambuenatitan palomoi	Filippi, García y Garrido 2011	Formación Anacleto	Argentina	Campaniano medio
27	Neuquensaurus australis	(Lydekker, 1893) Powell, 1986	Formación Anacleto	Argentina	Campaniano temprano
28	Panamericansaurus schroederi	Calvo y Porfiri 2010	Formación Allen	Argentina	Campaniano tardio-Maastrichtiano
29	Pellegrinisaurus powelli	Salgado 1996	Formación Anacleto	Argentina	Campaniano temprano
30	Petrobrasaurus puestohernandezi	Filippi, Canudo, Salgado, Garrido, García, Cerda y Otero 2011	Plottier Formation	Argentina	Coniaciano-Santoniano
31	Pitekunsaurus macayai	Filippi y Garrido 2008	Formación Anacleto	Argentina	Campaniano temprano
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32	Puertasaurus reuili	Novas, Salgado, Calvo, y Agnolin 2005	Formación Pari Aike	Argentina	Maastrichtiano temprano
33	Quetecsaurus rusconii	Gonzalez Riga y Ortiz David, 2014	Formacion Anacleto	Argentina	Campaniano inferior
34	Rinconsaurus caudamirus	Calvo y González Riga 2003	Formación Portezuelo-Plottier	Argentina	Turoniano tardío-Coniaciano
35	Rocasaurus muniozi	Salgado y Azpilicueta 2000	Formación Allen	Argentina	Campaniano tardío-Maastrichtiano temprano
36	Saltasaurus loricatus	Bonaparte y Powell 1980	Formación Lecho	Argentina	Campaniano-Maastrichtiano
37	Tapuiasaurus macedoi	Zaher, Pol, Carvalho, Nascimento, Riccomini, Larson, Juarez-Valieri, Pires-Domingues, da Silva, Campos 2011	Formación Quiricó	Brasil	Aptiano?
38	Traukutitan eocaudata	Juárez Valieri y Calvo 2011	Formación Bajo de la Carpa	Argentina	Santoniano
39	Trigonosaurus pricei	Campos, Kellner, Bertini, y Santucci 2005	Formación Marília	Brasil	Maastrichtiano
40	Uberabatitan ribeiroi	Salgado y Souza Carvalho 2008	Formación Marília	Brasil	Maastrichtiano

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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