Neuquensaurus
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''Neuquensaurus'' (meaning "
Neuquén Neuquén (; ) is the capital city of the Argentine province of Neuquén and of the Confluencia Department, located in the east of the province. It occupies a strip of land west of the confluence of the Limay and Neuquén rivers which form t ...
lizard") is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of saltasaurid
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
that lived in the Late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
, about 80 million years ago in
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
in South America. Its fossils were recovered from outcrops of the
Anacleto Formation The Anacleto Formation is a geologic formation with outcrops in the Argentina, Argentine Patagonian provinces of Mendoza Province, Mendoza, Río Negro Province, Río Negro, and Neuquén Province, Neuquén. It is the youngest formation within the ...
around
Cinco Saltos Cinco Saltos is a city in the province of Río Negro, Argentina, located on the eastern side of the valley of the Neuquén River, near the Pellegrini Lake, about northwest of Neuquén and from Allen. It has 19,819 inhabitants as per the . T ...
, near the Neuquén river from which its name is derived.


History

In 1893,
Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was a British naturalist, geologist and writer of numerous books on natural history. He was known for his contributions to zoology, paleontology, and biogeography. He worked extensively in cata ...
named ''
Titanosaurus ''Titanosaurus'' (; ) is a dubious genus of sauropod dinosaurs, first described by Richard Lydekker in 1877.R. Lydekker. (1877). Notices of new and other Vertebrata from Indian Tertiary and Secondary rocks. ''Records of the Geological Survey of ...
australis'', based on a series of caudal vertebrae and limb elements. The remains had been found by
Santiago Roth Santiago Roth (14 June 1850 – 4 August 1924) was a Swiss Argentine paleontologist and academic known for his fossil collections and Patagonian expeditions. Life Kaspar Jakob (Spanish: Santiago) was born and raised in Herisau, Canton Appenzell ...
and F. Romero in the
Neuquén Province Neuquén () is a Provinces of Argentina, province of Argentina, located in the west of the country, at the northern end of Patagonia. It borders Mendoza Province to the north, Rio Negro Province to the southeast, and Chile to the west. It also me ...
of
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
at the Neuquén River, and were by Lydekker assigned to a single individual. Six caudal vertebrae, with the inventory numbers MLP Ly 1-6-V-28-1, were the
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 ...
of the species. They had probably been found in a layer of the
Anacleto Formation The Anacleto Formation is a geologic formation with outcrops in the Argentina, Argentine Patagonian provinces of Mendoza Province, Mendoza, Río Negro Province, Río Negro, and Neuquén Province, Neuquén. It is the youngest formation within the ...
. Some elements that had been referred to ''Titanosaurus australis'' were reassigned to ''
Laplatasaurus ''Laplatasaurus'' (meaning "La Plata lizard", named for La Plata, Argentina) is a genus of titanosaurian sauropod dinosaur that lived during the Late Cretaceous in South America, with the holotype and only known specimen found in the Anacleto For ...
araukanicus'' by
Friedrich von Huene Baron Friedrich Richard von Hoyningen-Huene (22 March 1875 – 4 April 1969) was a German nobleman paleontologist who described a large number of dinosaurs, more than anyone else in 20th-century Europe. He studied a range of Permo-Carbonife ...
in 1929. The same year, von Huene named a ''Titanosaurus robustus'', and claimed it differed from ''T. australis'' in the limb material. Von Huene described all the slender limb material to ''T. australis'', but did not identify any differentiating features between the vertebrae. When describing ''T. robustus'', von Huene did not really compare other genera and species to it. From the
syntype In biological nomenclature, a syntype is any one of two or more biological types that is listed in a description of a taxon where no holotype was designated. Precise definitions of this and related terms for types have been established as part o ...
material assigned by von Huene to ''T. robustus''
José Fernando Bonaparte José is a predominantly Spanish and Portuguese form of the given name Joseph. While spelled alike, this name is pronounced very differently in each of the two languages: Spanish ; Portuguese (or ). In French, the name ''José'', pronounced , ...
''et al.'' in 1978 chose four
lectotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to anchor or centralizes ...
s, specimens MLP 26-250, MLP 26-252, MLP 26-254, and MLP 26-259, a left femur, both ulnae, and a left radius. In 1986, Jaime Eduardo Powell, concluding that ''Titanosaurus australis'' was less similar than ''Laplatasaurus araukanicus'' to ''Titanosaurus indicus'', named a separate genus: ''Neuquensaurus''. However, he did so in an unpublished dissertation which caused ''Neuquensaurus australis'' and ''Neuquensaurus robustus'' to remain invalid ''nomina ex dissertatione''. In 1990, the two species were assigned to ''
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 ...
'' by
John Stanton McIntosh John Stanton "Jack" McIntosh ( - ) was an American physicist and paleontologist who heavily influenced the study of sauropods. McIntosh worked professionally as a physicist at Wesleyan University until retirement in 1998, pursuing the study of sa ...
, as a ''Saltasaurus australis'' and a ''Saltasaurus robustus'', claiming that the features found by Bonaparte were not of sufficient taxonomic importance to justify a generic separation. In 1992 Powell validly named ''Neuquensaurus'', with as
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
''Titanosaurus australis'' of which the ''
combinatio nova In biological taxonomy, a combinatio nova (abbreviated comb. nov. or n. comb.) refers to the formal renaming of an organism's scientific name when it is transferred to a different genus, reclassified within a different species, or its taxonomic ...
'' then is ''Neuquensaurus australis''. He also found ''Titanosaurus robustus'' to be assignable to the new genus, but considered it non-diagnostical, and so a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
''.


Description

This dinosaur is believed to have possessed armor-like osteoderms. A relatively small sauropod, with a femur only long. It is one of the most completely known of Patagonian sauropods, measuring long and weighing . In addition to the original fossils described by Lydekker in 1893, it is represented by fossils collected in the early twentieth century, and more recent material, including a well preserved and partially articulated specimen described in 2005 (with two associated osteoderms).
Cervical vertebrae In tetrapods, cervical vertebrae (: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In saurop ...
are known from multiple specimens of ''Neuquensaurus'', although few are definitively associated with any other material. The centra are elongate, compressed vertically, and strongly , with an ovoid and rounded neural spine. Diagnostic of
Saltasaurinae Saltasaurinae is a subfamily of titanosaurian sauropods known from the late Cretaceous period of South America, India and Madagascar. Description Saltasaurines are relatively small sauropods with the general body shape of a small head, long nec ...
, the are shortened and robust and the extend far posteriorly to compensate. The lateral projections of the neural arch (transverse processes) are far more robust than in ''
Isisaurus ''Isisaurus'' (named after the Indian Statistical Institute) is a genus of titanosaurian dinosaur from the Late Cretaceous Lameta Formation of India and Pab Formation of Pakistan. The genus contains a single species, ''Isisaurus colberti''. Di ...
''. As in the cervicals, the dorsal vertebrae bear strongly opisthocoelous centra with lateral pneumatic openings. In the dorsals, however, the pleurocoels are posteriorly pinched and internally subdivided, a regular feature of titanosaurs. The pleurocoels of ''Neuquensaurus'' are proportionally taller than in '' Trigonosaurus'' or ''
Lirainosaurus ''Lirainosaurus'' (meaning "slender lizard"; from the Basque ''lirain'', meaning "slender", and the Greek ''sauros'', meaning "lizard") is a genus of titanosaur sauropod which lived in what is now Spain. The type species, ''Lirainosaurus astibiae ...
''. A ventral keel is lost throughout the dorsal series of probably 10 vertebrae, although the total count is uncertain. The vertebrae also lean less posteriorly towards the pelvis, initially inclined far beyond their individual centrum. However, they lack the of non-titanosaurs that limit vertebral motion. Two accessory are present in the dorsals of ''Neuquensaurus'' that are absent even in the close relatives ''Saltasaurus'' and ''
Rocasaurus ''Rocasaurus'' (meaning " General Roca lizard") 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 Maastr ...
''. Unique among all sauropods, the
sacrum The sacrum (: sacra or sacrums), in human anatomy, is a triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part of the pelvic cavity, ...
of ''Neuquensaurus'' is composed of seven vertebrae. The last vertebra is convex anteriorly and posteriorly, and diagnostically the middle vertebrae are strongly compressed in width. Like other derived titanosaurs, all
caudal vertebrae Caudal vertebrae are the vertebrae of the tail in many vertebrates. In birds, the last few caudal vertebrae fuse into the pygostyle, and in apes, including humans, the caudal vertebrae are fused into the coccyx. In many reptiles, some of the caud ...
of ''Neuquensaurus'' are strongly and vertically compressed. There is a prominent ventral depression like in ''Saltasaurus'' and ''Rocasaurus'', but contrasting the other genera there is no ridge dividing this depression. Additionally, the neural spines of vertebrae become progressively more reclined, and a prespinal lamina is present in all vertebrae. ''Neuquensaurus'' has an almost complete skeleton. Among the preserved elements are a scapula. The scapula is co-ossified with the coracoid, a feature also found in ''
Opisthocoelicaudia ''Opisthocoelicaudia'' is a genus of sauropod dinosaur of the Late Cretaceous Period discovered in the Gobi Desert of Mongolia. The type species is ''Opisthocoelicaudia skarzynskii''. A well-preserved skeleton lacking only the head and neck wa ...
''. In general aspect, the scapula resembles ''Saltasaurus'', ''Opisthocoelicaudia'', ''
Lirainosaurus ''Lirainosaurus'' (meaning "slender lizard"; from the Basque ''lirain'', meaning "slender", and the Greek ''sauros'', meaning "lizard") is a genus of titanosaur sauropod which lived in what is now Spain. The type species, ''Lirainosaurus astibiae ...
'', and ''
Alamosaurus ''Alamosaurus'' (; meaning "Ojo Alamo lizard") is a genus of titanosaurian sauropod dinosaurs containing a single known species, ''Alamosaurus sanjuanensis'', from the Maastrichtian age of the Late Cretaceous period in what is now southwestern ...
''. The coracoids of the scapulae are roughly quadrangular in shape, and are alike ''Saltasaurus'' and ''Lirainosaurus'', but not ''Opisthocoelicaudia'', ''
Rapetosaurus ''Rapetosaurus'' ( ) is a genus of titanosaurian sauropod dinosaur that lived in Madagascar from 70 to 66 million years ago, at the end of the Cretaceous Period. Only one species, ''Rapetosaurus krausei'', has been identified. Like other sauropod ...
'', or ''
Isisaurus ''Isisaurus'' (named after the Indian Statistical Institute) is a genus of titanosaurian dinosaur from the Late Cretaceous Lameta Formation of India and Pab Formation of Pakistan. The genus contains a single species, ''Isisaurus colberti''. Di ...
'', all of which have a rounded coracoid. Many characters distinguish ''Neuquensaurus'' from other titanosaurians. The features found by Otero in 2010 are: the possession of posterior caudal centra that are dorsoventrally flattened, and strongly developed fibular lateral tuberosity.


Classification

''Neuquensaurus'' is a derived
saltasaurine Saltasauridae (named after the Salta region of Argentina where they were first found) is a family of armored herbivorous sauropods from the Upper Cretaceous. They are known from fossils found in South America, Africa, Asia, North America, and Eur ...
. Its closest relatives were ''
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 ...
'', ''
Rocasaurus ''Rocasaurus'' (meaning " General Roca lizard") 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 Maastr ...
'', and ''
Bonatitan ''Bonatitan'' is a genus of titanosaurian dinosaur from the Late Cretaceous Allen Formation of Argentina. It was named in 2004. Description The type species is ''Bonatitan reigi'', first described by Martinelli and Forasiepi in 2004. The speci ...
'', and together they made up Saltasaurinae. The cladogram from Rubilar-Rogers ''et al.'' (2021) is shown below:


Paleoecology

''Neuquensaurus'' is known from fossils from the
Anacleto Formation The Anacleto Formation is a geologic formation with outcrops in the Argentina, Argentine Patagonian provinces of Mendoza Province, Mendoza, Río Negro Province, Río Negro, and Neuquén Province, Neuquén. It is the youngest formation within the ...
.


References


External links

{{Taxonbar, from=Q284931 Saltasauridae Dinosaur genera Campanian dinosaurs Anacleto Formation Taxa named by Jaime Powell Fossil taxa described in 1992 Dinosaurs of Argentina