Titanomachya Temporal range: Late Cretaceous, | |
---|---|
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | † Sauropodomorpha |
Clade: | † Sauropoda |
Clade: | † Macronaria |
Clade: | † Titanosauria |
Superfamily: | † Saltasauroidea |
Genus: | † Titanomachya Pérez-Moreno et al., 2024 |
Type species | |
Titanomachya gimenezi Pérez-Moreno et al., 2024 |
Titanomachya (named after the Titanomachy of Greek mythology) is an extinct genus of titanosaurian sauropod dinosaur from the Late Cretaceous La Colonia Formation of Argentina. The genus contains a single species, T. gimenezi. It is a relatively small titanosaur, weighing around 7.8 tonnes (8.6 short tons).
The Titanomachya holotype specimen, MPEF Pv 11547, was discovered in sediments of the La Colonia Formation near the Cerro Bayo mountain and Bajada del Diablo crater in Chubut Province of Patagonia, Argentina. The specimen consists of an incomplete, partially articulated skeleton, including a caudal vertebra, several fragmentary ribs, two haemal arches, the left humerus, fragments of the pelvic girdle, part of both femora, both tibiae and fibulae, and parts of both astragali. [1]
In 2024, Pérez-Moreno et al. described Titanomachya gimenezi as a new genus and species of titanosaurian sauropod based on these fossil remains. Two partial caudal vertebrae belonging to saltasauroid titanosaurs are also known from the formation, but were not referred to Titanomachya. The generic name, Titanomachya, alludes to the mythological Greek Titanomachy—the battle where the Olympian gods fought and defeated the Titans—referring to the species' proximity to the extinction of the titanosaurs. The specific name, gimenezi, honors Olga Giménez and her paleontological contributions to the study of Argentinian dinosaurs from Chubut Province. [1]
Titanomachya represents the first saltasauroid titanosaur to be discovered in Central Patagonian sediments dated to the end of the Cretaceous. The only other saltasauroid from the North Patagonian Massif is the similarly aged Dreadnoughtus from Argentina's Cerro Fortaleza Formation. [1]
Titanomachya is a small titanosaur, with an estimated length of 6 metres (20 ft). [2] It probably weighed between 5.8 and 9.8 tonnes (6.4 and 10.8 short tons), with a mean body mass of 7.8 tonnes (8.6 short tons). The holotype specimen was an adult similar in size to some related saltasaurids ( Neuquensaurus and Saltasaurus ), but it may have weighed more, as indicated by an astragalus modified for improved weight-bearing. The morphology of the astragalus is intermediate between colossosaurs and saltasauroids. [1]
Pérez-Moreno et al. (2024) entered Titanomachya into a phylogenetic analysis and found it to be in the clade Lithostrotia, itself recovered as a subclade of the Saltasauroidea. This result is similar to that recovered in the 2023 description of Bustingorrytitan . [3] The results of Pérez-Moreno et al. are shown in the cladogram below: [1]
Titanomachya was found in outcrops of the La Colonia Formation, which dates to the Maastrichtian (Cretaceous)–Paleocene (Paleogene) boundary. [4] This formation is best known for fossils of the abelisaurid theropod Carnotaurus , as well as fossils of other reptiles including the turtle Patagoniaemys , the snake Alamitophis and the plesiosaurs Kawanectes , Chubutinectes and Sulcusuchus . [5] [6] [7] [8] While Titanomachya represents the first herbivorous dinosaur named from this locality, remains of indeterminate hadrosaurs and ankylosaurs have also been found. The small adult size of Titanomachya compared to giant titanosaurs may have been a result of competition with an increased diversity of these other herbivores. [1] [2]
Aeolosaurus is a genus of titanosaurian sauropod dinosaur from the Late Cretaceous Period of what is now South America. Like most sauropods, it would have been a quadrupedal herbivore with a long neck and tail. Aeolosaurus is well known for a titanosaur, as it is represented by the remains of several individuals belonging to at least two species. However, like most titanosaurs, no remains of the skull are known. The holotype of Aeolosaurus rionegrinus consists of a series of seven tail vertebrae, as well as parts of both forelimbs and the right hindlimb. It was discovered in the Angostura Colorada Formation in Argentina, which dates from the Campanian stage of the Late Cretaceous, about 83 to 74 million years ago. The species A. maximus was transferred over to the new genus Arrudatitan in 2021.
Bonitasaura is a genus of titanosaurian dinosaur hailing from uppermost layers of the Late Cretaceous (Santonian) Bajo de la Carpa Formation, Neuquén Group of the eastern Neuquén Basin, located in Río Negro Province, Northwestern Patagonia, Argentina. The remains, consisting of a partial sub-adult skeleton jumbled in a small area of fluvial sandstone, including a lower jaw with teeth, a partial vertebrae series, and limb bones, were described by Sebastian Apesteguía in 2004.
Rocasaurus is a genus of titanosaurian sauropod that lived in South America. Rocasaurus was discovered in Argentina in 2000, within the Allen Formation which is dated to be middle Campanian to early Maastrichtian in age. This genus grew up to 8 metres (26 ft) long, making it one of the smaller sauropods. It seems to be closely related to saltasaurid dinosaurs, like Saltasaurus and Neuquensaurus.
The Huincul Formation is a geologic formation of Late Cretaceous age of the Neuquén Basin that outcrops in the Mendoza, Río Negro and Neuquén Provinces of northern Patagonia, Argentina. It is the second formation in the Río Limay Subgroup, the oldest subgroup within the Neuquén Group. Formerly that subgroup was treated as a formation, and the Huincul Formation was known as the Huincul Member.
Pellegrinisaurus is a genus of titanosaurian sauropod dinosaur that lived in South America during the Late Cretaceous period. The holotype was found in the Allen Formation, Argentina.
Rinconsaurus is a genus of titanosaur sauropod dinosaur from the Late Cretaceous in what is now Argentina. The type species, Rinconsaurus caudamirus, was described by Calvo and Riga in 2003, and is based on three partial skeletons.
The Anacleto Formation is a geologic formation with outcrops in the Argentine Patagonian provinces of Mendoza, Río Negro, and Neuquén. It is the youngest formation within the Neuquén Group and belongs to the Río Colorado Subgroup. Formerly that subgroup was treated as a formation, and the Anacleto Formation was known as the Anacleto Member.
Tuarangisaurus is an extinct genus of elasmosaurid known from New Zealand. The type and only known species is Tuarangisaurus keyesi, named by Wiffen and Moisley in 1986.
Uberabatitan is a genus of titanosaurian sauropod dinosaur from the Late Cretaceous of Brazil. It is known from bones including neck, back, and tail vertebrae, pelvic bones, and limb bones. These fossils were found in the uppermost portion of the Maastrichtian-age Serra da Galga Formation of the Bauru Group, in Uberaba, Minas Gerais. The type species, described by Salgado and Carvalho in 2008, is U. ribeiroi. To date, it is the most recent titanosaur from Bauru Group rocks; other titanosaurs from the Bauru Group, including Baurutitan and Trigonosaurus, come from lower levels.
The La Colonia Formation is a geological formation in Argentina whose strata date back to the Late Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.
The Allen Formation is a geological formation in Argentina whose strata date back to the Late Cretaceous (middle Campanian to early Maastrichtian. Dinosaur remains are among the fossils that have been recovered from the formation. Indeterminate chelid remains and other vertebrates have also been discovered in this formation.
The Snow Hill Island Formation is an Early Maastrichtian geologic formation found on James Ross Island, James Ross Island group, Antarctica. Remains of a paravian theropod Imperobator antarcticus have been recovered from it, as well as the elasmarian ornithopods Trinisaura santamartaensis, Biscoveosaurus and Morrosaurus antarcticus, the ankylosaurian Antarctopelta oliveroi, and the shark Notidanodon sp. Alongside these described genera are also the remains of indeterminate elasmosaurids, lithostrotian titanosaurs and an indeterminate pterosaur.
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.
Eutitanosauria is a clade of titanosaurs, encompassing the more derived members of the group and characterized by the absence of the hyposphene-hypantrum articulation and possibly the presence of osteoderms. The group was first named by Sanz and colleagues in 1999, who used it to unite the group of Argyrosaurus, Lirainosaurus, Saltasaurus and the Peiropolis titanosaur. However, this definition was not used as it made the group equivalent to Saltasauridae, so Saldago redefined it in 2003 to be all titanosaurs closer to Saltasaurus than Epachthosaurus. This definition created Eutitanosauria as the sister group to Epachthosaurinae, but was problematic due to the variable nature of Epachthosaurus. Eutitanosauria was often broadly similar to Lithostrotia, and has often been unused or unlabelled on phylogenies. Sometimes Epachthosaurus would be more primitive than Malawisaurus, making Eutitanosauria more encompassing than Lithostrotia, or Epachthosaurus could nest close to Colossosauria and limit Eutitanosauria to a smaller group of saltasauroids. Because of the flexible nature of Epachthosaurus in basal titanosaur phylogeny, Carballido and colleagues redefined the group in 2022 to include the smallest clade of both Patagotitan, a colossosaur, and Saltasaurus, creating a node-stem clade with Colossosauria and Saltasauroidea, presenting the informal cladogram of stable titanosaur clades below.
Kawanectes is a genus of elasmosaurid plesiosaur, a type of long-necked marine reptile, that lived in the marginal marine environment of Late Cretaceous Patagonia. It contains one species, K. lafquenianum, described in 2016 by O'Gorman.
Saltasauroidea is a superfamily of titanosaurs named by França and colleagues in 2016 based on their phylogenetic results, for a clade uniting Aeolosaurini and Saltasauridae, as well as the intermediate genera Baurutitan, Diamantinasaurus and Isisaurus. The group was not defined or discussed in the text, but was supported by Carballido and colleagues in 2022 as a useful designation for subdividing titanosaurs. As there was no discussion about the intentions for the clade, Carballido gave it the definition of all taxa closer to Saltasaurus than Patagotitan, encompassing half of Eutitanosauria as the sister taxon to the inversely defined Colossosauria. Carballido et al. placed Nemegtosauridae and Saltasauridae within the group, though they had Aeolosaurini within Colossosauria. The informal cladogram of titanosaur relationships they proposed is shown below.
Menucocelsior is a genus of medium-sized titanosaurian sauropod dinosaur from the Late Cretaceous Allen Formation of Argentina. The type and only species is Menucocelsior arriagadai.
Teratopodus is an ichnogenus of titanosaurian sauropod footprint. It includes a single species, T. malarguensis, known from prints found in the Late Cretaceous Anacleto Formation of Argentina. The Teratopodus tracks represent some of the best sauropod pes tracks currently known from South America.
Chubutinectes is an extinct genus of elasmosaurid plesiosaur from the Late Cretaceous La Colonia Formation of Argentina. The genus contains a single species, C. carmeloi, known from a partial skeleton and associated gastroliths.