''Titanomachya'' (named after the

Titanomachy In Greek mythology, the Titanomachy (; ) was a ten-year war fought in ancient Thessaly, consisting of most of the Titans (the older generation of gods, based on Mount Othrys) fighting against the Twelve Olympians, Olympians (the younger generati ...

of Greek mythology) is an

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

genus of

titanosauria Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of Sauropoda, sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with tax ...

n sauropod dinosaur from the

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...

La Colonia Formation The La Colonia Formation is a geological Formation (geology), formation in Argentina whose strata date back to the Late Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.Weishampel et al., 2004, pp.600- ...

of Argentina. The genus contains a single species, ''T. gimenezi''. It is a relatively small titanosaur, weighing around .

Discovery and naming

The ''Titanomachya''

holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...

specimen, MPEF Pv 11547, was discovered in sediments of the

near the

Cerro Bayo Cerro Bayo is a mountain of the Andes range located 9 km from the town Villa La Angostura, Neuquén Province, Argentina, within the Valdivian temperate rain forests, in an area with numerous lakes. The mountain hosts a ski area with 25 run ...

mountain and Bajada del Diablo crater in

Chubut Province Chubut ( from Tehuelche language, Tehuelche 'transparent'; ) is a provinces of Argentina, province in southern Argentina, situated between the 42nd parallel south (the border with Río Negro Province), the 46th parallel south (bordering Santa ...

Patagonia Patagonia () is a geographical region that includes parts of Argentina and Chile at the southern end of South America. The region includes the southern section of the Andes mountain chain with lakes, fjords, temperate rainforests, and glaciers ...

, Argentina. The specimen consists of an incomplete, partially

articulated An articulated vehicle is a vehicle which has a permanent or semi-permanent coupling in its construction. This coupling works as a large pivot joint, allowing it to bend and turn more sharply. There are many kinds, from heavy equipment to buse ...

skeleton, including a

caudal vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...

, several fragmentary ribs, two

haemal arch A haemal arch, also known as a chevron, is a bony arch on the ventral side of a tail vertebra of a vertebrate. The canal formed by the space between the arch and the vertebral body is the haemal canal. A spinous ventral process emerging from the ha ...

es, the left

humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...

, fragments of the

pelvic girdle The hip bone (os coxae, innominate bone, pelvic bone or coxal bone) is a large flat bone, constricted in the center and expanded above and below. In some vertebrates (including humans before puberty) it is composed of three parts: the Ilium (bone) ...

, part of both

femora The femur (; : femurs or femora ), or thigh bone is the only bone in the thigh — the region of the lower limb between the hip and the knee. In many four-legged animals the femur is the upper bone of the hindleg. The top of the femur fits in ...

, both

tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...

e and

fibula The fibula (: fibulae or fibulas) or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. ...

e, and parts of both astragali. Joachim Wtewael - The Battle Between the Gods and the Titans - WGA25902

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

—the battle where the

Olympian gods upright=1.8, Fragment of a relief (1st century BC1st century AD) depicting the twelve Olympians carrying their attributes in procession; from left to right: Hestia (scepter), Hermes (winged cap and staff), Aphrodite (veiled), Ares (helmet and s ...

fought and defeated the

Titans In Greek mythology, the Titans ( ; ) were the pre-Twelve Olympians, Olympian gods. According to the ''Theogony'' of Hesiod, they were the twelve children of the primordial parents Uranus (mythology), Uranus (Sky) and Gaia (Earth). The six male ...

—referring to the species' proximity to the extinction of the titanosaurs. The

specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...

, ''gimenezi'', honors Olga Giménez and her paleontological contributions to the study of Argentinian dinosaurs from

. ''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 The North Patagonian Massif or Somún Cura Massif ( Spanish: ''Macizo Norpatagónico'', ''Macizo Nordpatagónico'' or ''Macizo de Somún Cura'') is a massif in northern Patagonia located in the Argentine provinces of Río Negro and Chubut. The ma ...

is the similarly aged ''

Dreadnoughtus ''Dreadnoughtus'' is a genus of titanosaurian sauropod dinosaur containing a single species, ''Dreadnoughtus schrani''. ''D. schrani'' is known from two partial skeletons discovered in Upper Cretaceous (Campanian to Maastrichtian; approximately ...

'' from Argentina's

Cerro Fortaleza Formation The Cerro Fortaleza Formation, in older literature described as Pari Aike Formation, is a Late Cretaceous geologic formation of Campanian to Maastrichtian age (although it has formerly been reported to be Cenomanian to Santonian) of the Austra ...

Description

''Titanomachya'' is a small titanosaur, with an estimated length of . It probably weighed between , with a mean body mass of . The holotype specimen was an adult similar in size to some related

saltasaurids Saltasauridae (named after the Salta region of Argentina where they were first found) is a family of armored herbivorous Sauropoda, sauropods from the Upper Cretaceous. They are known from fossils found in South America, Africa, Asia, North Ameri ...

(''

Neuquensaurus ''Neuquensaurus'' (meaning "Neuquén lizard") is a genus of saltasaurid sauropod dinosaur that lived in the Late Cretaceous, about 80 million years ago in Argentina in South America. Its fossils were recovered from outcrops of the Anacleto Forma ...

'' and ''

Saltasaurus ''Saltasaurus'' (which means "lizard from Salta") is a genus of saltasaurid dinosaur of the Late Cretaceous Period (geology), period of Argentina. Small among sauropods, though still heavy by the standards of modern creatures, ''Saltasaurus'' was ...

''), but it may have weighed more, as indicated by an

astragalus Astragalus may refer to: * ''Astragalus'' (plant), a large genus of herbs and small shrubs *Astragalus (bone) The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...

modified for improved weight-bearing. The morphology of the astragalus is intermediate between colossosaurs and saltasauroids.

Classification

Pérez-Moreno et al. (2024) entered ''Titanomachya'' into a

phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...

and found it to be in the clade

Lithostrotia Lithostrotia is a clade of derived titanosaur sauropods that lived during the Early Cretaceous and Late Cretaceous. The group was defined by Upchurch ''et al.'' in 2004 as the most recent common ancestor of ''Malawisaurus'' and ''Saltasaurus'' an ...

, itself recovered as a subclade of the Saltasauroidea. This result is similar to that recovered in the 2023 description of '' Bustingorrytitan''. The results of Pérez-Moreno et al. are shown in the

cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...

below:

Paleoenvironment

''Titanomachya'' was found in outcrops of the

, which dates to the

Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...

(Cretaceous)–

Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...

(Paleogene) boundary. This formation is best known for fossils of the

abelisaurid Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are foun ...

theropods ''

Carnotaurus ''Carnotaurus'' (; ) is a genus of Theropoda, theropod dinosaur that lived in South America during the Late Cretaceous period, probably sometime between 72 and 69 million years ago. The only species is ''Carnotaurus sastrei''. Known from a si ...

'' and ''

Koleken ''Koleken'' (meaning "coming from clay and water") is a genus of Carnotaurini, carnotaurin abelisaurid from the Maastrichtian La Colonia Formation in the Chubut Province of Argentina. The type and only species is ''K. inakayali'', known from on ...

'', as well as fossils of other reptiles including the turtle '' Patagoniaemys'', the snake ''

Alamitophis ''Alamitophis'' is a genus of fossil snakes in the extinct family of Madtsoiidae. Its length is estimated at and it probably fed on frogs, lizards, and small mammals. It is found in Australia (Tingamarra Fauna, after which ''A. tingamarra'' is n ...

'' and the

plesiosaurs The Plesiosauria or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared in the latest Triassic Period (geology), Period, possibly in the Rhaetian st ...

Kawanectes ''Kawanectes'' (meaning "Kawas swimmer") is a genus of elasmosauridae, elasmosaurid plesiosaur, a type of long-necked marine reptile, that lived in the marginal marine (probably estuary, estuarine) environment of Late Cretaceous Patagonia. It co ...

'', ''

Chubutinectes ''Chubutinectes'' (meaning "Chubut Province, Chubut swimmer") is an extinct genus of elasmosaurid plesiosaur from the Late Cretaceous La Colonia Formation of Argentina. The genus contains a Monotypic taxon, single species, ''C. carmeloi'', known ...

'' and ''

Sulcusuchus ''Sulcusuchus'' is a genus of polycotylid plesiosaur from the Late Campanian to Early Maastrichtian of Argentina. Description The type species ''Sulcusuchus erraini'' was named in 1990 by Zulma Brandoni de Gasparini and Luis Spalletti. It w ...

''. Mammals discovered from the formation include ''

Reigitherium ''Reigitherium'' was a mammal that lived during the Late Cretaceous, in the (Late Campanian-Maastrichtian). Its fossils have been found in the Los Alamitos and the La Colonia Formations of Argentina. Description The original specimen of ''Rei ...

'', ''

Coloniatherium ''Coloniatherium'' is a meridiolestid mammal from the Late Cretaceous of Argentina. The single species, ''Coloniatherium cilinskii'', was a large member of the family Mesungulatidae. Taxonomy ''Coloniatherium'' was named in 2009 by Guillermo R ...

'', ''

Argentodites ''Argentodites'' is a possible multituberculate mammal from the Cretaceous of Argentina. The single species, ''Argentodites coloniensis'', is known from a single blade-like fourth lower premolar (p4) from the La Colonia Formation, which is mostl ...

'' and ''

Ferugliotherium ''Ferugliotherium'' is a genus of fossil mammals in the family Ferugliotheriidae from the Campanian and/or Maastrichtian period (Late Cretaceous; around 70 million years ago) of Argentina. It contains a single species, ''Ferugliotherium windhaus ...

''. Remains of an

enantiornithine The Enantiornithes, also known as enantiornithines or enantiornitheans in literature, are a group of extinct avialans ("birds" in the broad sense), the most abundant and diverse group known from the Mesozoic era. Almost all retained teeth and c ...

and, possibly, of a neornithine bird have been discovered. While ''Titanomachya'' represents the first herbivorous dinosaur named from this locality, remains of indeterminate

hadrosaurs Hadrosaurids (), also hadrosaurs or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod fami ...

and

ankylosaurs Ankylosauria is a group of herbivorous dinosaurs of the clade Ornithischia. It includes the great majority of dinosaurs with Armour (zoology), armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short ...

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.

References

{{Taxonbar, from=Q125421035 Lithostrotia Dinosaur genera Maastrichtian dinosaurs Fossil taxa described in 2024 Dinosaurs of Argentina