Saltasaurids Temporal range: Late Cretaceous, | |
---|---|
![]() | |
Restoration of Saltasaurus | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | † Sauropodomorpha |
Clade: | † Sauropoda |
Clade: | † Macronaria |
Clade: | † Titanosauria |
Superfamily: | † Saltasauroidea |
Family: | † Saltasauridae Powell, 1992 |
Genera | |
Synonyms | |
|
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, [4] Asia, North America, and Europe. They are characterized by their vertebrae and feet, which are similar to those of Saltasaurus , the first of the group to be discovered and the source of the name. The last and largest of the group and only one found in North America, Alamosaurus , was thirty-four metres (112 feet) in length and one of the last sauropods to go extinct.
Most of the saltasaurids were smaller, around fifteen metres (49 feet) in length, and one, Rocasaurus , was only eight metres (26 feet) long. Like all sauropods, the saltasaurids were quadrupeds, their necks and tails were held almost parallel to the ground, and their small heads had only tiny, peg-like teeth. They were herbivorous, stripping leaves off of plants and digesting them in their enormous guts. [5] Although large animals, they were smaller than other sauropods of their time, and many possessed distinctive additional defenses in the form of scutes along their backs.
As sauropods, the Saltasauridae are herbivorous saurischians with the characteristic body plan of a small head, long neck, four erect legs, and a counterbalancing tail. Most sauropods are from the clade Neosauropoda, which is further split into the narrow-toothed Diplodocoidea and the broad-toothed Macronaria. The Macronarians emerged in the Jurassic and a subclade, the Titanosauria, survived into the Cretaceous and spread across the continents. Because of their diversity, wide distribution, and the fragmentary or incomplete nature of most specimens, little is known about the titanosaurs beyond their size and tendency to have scutes. [6]
The saltasaurids, one of the several titanosaur families, are recognized by the convexities in certain caudal vertebrae and the markings on their coracoid bones. [7] All saltasaurids have thirty-five or fewer caudal vertebrae, [8] each of which is convex on both sides of its centrum, and the one closest to the tail is shorter than the others. [9] Their coracoid bones have rectangular margins on the anteroventral side, as well as a lip where they meet the infraglenoid. The Opisthocoelicaudiinae, a subfamily of the saltasaurids, are unique in that they lack phalanges in their forelimbs. [8] [10] Although Saltasaurus is known to possess dorsal osteoderms, scutes have not been discovered in all saltasaurids, and it is unclear when and where the evolution of osteoderms occurred in saltasaurids and titanosaurs in general. [11]
The first saltasaurid to be discovered was Alamosaurus, found by paleontologist Charles Gilmore in Utah in 1922. The next species would not be described until Opisthocoelicaudia was named by Magdalena Borsuk-Bialynicka from a postcranial material in Mongolia in 1977. In 1980, Jose Bonaparte and Jaime Powell discovered Saltasaurus in Argentina. This was the first sauropod to be discovered with armor and proved that sauropods had thrived in Cretaceous South America. Paul Sereno eventually recognized a cladistic relationship between Opisthocoelicauda and Saltasaurus to create the family Saltasauridae. [12]
The group is defined by the characteristics shared by all with the two best-known members, Saltasaurus and Opisthocoelicaudia. Paleontologists J Wilson and P Upchurch defined the Saltasauridae in 2003 as the least inclusive clade containing Opisthocoelicaudia skarzynskii, and Saltasaurus loricatus, their most recent common ancestor, and all that species’ descendants.
This taxonomy is based on those of González Riga et al. (2009) and Curry Rogers & Wilson (2005). [13] [14]
The family is then further divided into two subfamilies. Wilson and Upchurch defined Saltasaurinae in 2003 as the least-inclusive clade containing Saltasaurus but not Opisthocoelicaudia. The same paleontologists defined Opisthocoelicaudiinae as the inverse: the least-inclusive clade containing Opisthocoelicaudia but not Saltasaurus. Some species, due to the incompleteness of their skeletons, cannot yet be placed in either subfamily.
Saltasauridae in a cladogram after Navarro et al., 2022: [15]
Saltasauridae | |
Many fragmentary saltasaurids have been discovered since 1980, placing members of the family in territories as widely dispersed as today's Australia, Madagascar, and France, in addition to their earlier-known residencies in North and South America. Like the other titanosaurs, the saltasaurids where a widespread, successful group that colonized all continents in the Cretaceous. [16]
Like all titanosaurs, the saltasaurids possessed small, peg-like teeth that were not usable for chewing. Coproliths from an unidentified titanosaur found in India suggest a diet of conifers, cycads, and early species of grasses. [17] Unable to chew and probably lacking gastroliths, sauropods survived by retaining plant matter in their stomachs for long periods of time, fermenting it to extract as many resources as possible. Their long necks allowed them to graze over a large area while standing, reducing energy use.
The osteoderms of Saltasaurus consisted of numerous, large bony plates embedded in the dorsal skin, each surrounded by a pattern of smaller plates. The large osteoderms contained some hollow spaces for blood vessels and spongy trabecular bone, while the small ones were solid. [18] Patches of skin from unidentified Cretaceous titanosaurs have revealed similar scale patterns in embryos (a large scale surrounded by ten smaller ones) but no bone or mineralized structure, suggesting that, like crocodiles, those saltasaurids that possessed armor only developed it some time after hatching. Analysis of the osteoderms of the titanosaur Rapetosaurus revealed that the bones were hollow in adults, while those of juveniles were solid pieces similar to those in crocodiles. Paleontologist Kristina Curry Rogers, who made this discovery, theorized that the adult animals used their hollow osteoderms to store minerals during lean times. It is unknown whether any of the Saltasauridae used their osteoderms in a similar manner. [14]
The same Argentine dig site, Auca Mahuevo, that provided information on embryonic skin, has also yielded information on the nesting habits of titanosaurs, but not saltasaurids specifically. The nests were constructed on the surface by piling debris in a ring around the eggs, with the eggs themselves left uncovered. Each egg was porous and spherical, about 14 cm in diameter, and they were laid in clutches. The embryos show smaller rostrum and nares close to the anterior portion of the face compared to adult titanosaurs, suggesting that the nostrils may have moved towards the back of the head as the animal grew. [19]
Saltasaurus is a genus of saltasaurid dinosaur of the Late Cretaceous period of Argentina. Small among sauropods, though still heavy by the standards of modern creatures, Saltasaurus was characterized by a short neck and stubby limbs. It was the first genus of sauropod known to possess armour of bony plates embedded in its skin. Such small bony plates, called osteoderms, have since been found on other titanosaurians.
Titanosaurs were a diverse group of sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with taxa still thriving at the time of the extinction event at the end of the Cretaceous. This group includes some of the largest land animals known to have ever existed, such as Patagotitan—estimated at 37 m (121 ft) long with a weight of 69 tonnes —and the comparably-sized Argentinosaurus and Puertasaurus from the same region.
Nemegtosauridae is a family of titanosaurian sauropod dinosaurs based on their diplodocid-like skulls. Only three species are known: Nemegtosaurus, Quaesitosaurus and possibly Tapuiasaurus, each from the Cretaceous.
Alamosaurus is a genus of opisthocoelicaudiine titanosaurian sauropod dinosaurs containing a single known species, Alamosaurus sanjuanensis, from the Maastrichtian age of the Late Cretaceous period in what is now southwestern North America. Isolated vertebrae and limb bones indicate that it reached sizes comparable to Argentinosaurus and Puertasaurus, which would make it the absolute largest dinosaur known from North America. Its fossils have been recovered from a variety of rock formations spanning the Maastrichtian age. Specimens of a juvenile Alamosaurus sanjuanensis have been recovered from only a few meters below the Cretaceous-Paleogene boundary in Texas, making it among the last surviving non-avian dinosaur species. Alamosaurus is the only known sauropod to have inhabited North America after their nearly 30-million year absence from the North American fossil record and probably represents an immigrant from South America.
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.
Agustinia is a genus of sauropod dinosaur from the Early Cretaceous of South America. The genus contains a single species, Agustinia ligabuei, known from a single specimen that was recovered from the Lohan Cura Formation of Neuquén Province in Argentina. It lived about 116–108 million years ago, in the Aptian–Albian stages of the Early Cretaceous Period.
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 was unearthed in 1965 by Polish and Mongolian scientists, making Opisthocoelicaudia one of the best known sauropods from the Late Cretaceous. Tooth marks on this skeleton indicate that large carnivorous dinosaurs had fed on the carcass and possibly had carried away the now-missing parts. To date, only two additional, much less complete specimens are known, including part of a shoulder and a fragmentary tail. A relatively small sauropod, Opisthocoelicaudia measured about 11.4–13 m (37–43 ft) in length. Like other sauropods, it would have been characterised by a small head sitting on a very long neck and a barrel shaped trunk carried by four column-like legs. The name Opisthocoelicaudia means "posterior cavity tail", alluding to the unusual, opisthocoel condition of the anterior tail vertebrae that were concave on their posterior sides. This and other skeletal features lead researchers to propose that Opisthocoelicaudia was able to rear on its hindlegs.
Ampelosaurus is a titanosaurian sauropod dinosaur from the Late Cretaceous Period of what is now France. Its type species is A. atacis, named by Le Loeuff in 1995. Its remains were found in a level dating from 71.5 million years ago representing the early Maastrichtian.
Andesaurus is a genus of basal titanosaurian sauropod dinosaur which existed during the middle of the Cretaceous Period in South America. Like most sauropods, belonging to one of the largest animals ever to walk the Earth, it would have had a small head on the end of a long neck and an equally long tail.
Neuquensaurus 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 Formation around Cinco Saltos, near the Neuquén river from which its name is derived.
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.
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.
Neosauropoda is a clade within Dinosauria, coined in 1986 by Argentine paleontologist José Bonaparte and currently described as Saltasaurus loricatus, Diplodocus longus, and all animals directly descended from their most recent common ancestor. The group is composed of two subgroups: Diplodocoidea and Macronaria. Arising in the early Jurassic and persisting until the Cretaceous-Paleogene extinction event, Neosauropoda contains the majority of sauropod genera, including genera such as Apatosaurus, Brachiosaurus, and Diplodocus. It also includes giants such as Argentinosaurus, Patagotitan and Sauroposeidon, and its members remain the largest land animals ever to have lived.
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 and all the descendants of that ancestor. Lithostrotia is derived from the Ancient Greek lithostros, meaning "inlaid with stones", referring to the fact that many known lithostrotians are preserved with osteoderms. However, osteoderms are not a distinguishing feature of the group, as the two noted by Unchurch et al. include caudal vertebrae with strongly concave front faces (procoely), although the farthest vertebrae are not procoelous.
Diamantinasaurus is a genus of titanosaurian sauropod from Australia that lived during the early Late Cretaceous, about 94 million years ago. The type species of the genus is D. matildae, first described and named in 2009 by Scott Hocknull and colleagues based on fossil finds in the Winton Formation. Meaning "Diamantina lizard", the name is derived from the location of the nearby Diamantina River and the Greek word sauros, "lizard". The specific epithet is from the Australian song Waltzing Matilda, also the locality of the holotype and paratype. The known skeleton includes most of the forelimb, shoulder girdle, pelvis, hindlimb and ribs of the holotype, and one shoulder bone, a radius and some vertebrae of the paratype.
Opisthocoelicaudiinae is a subfamily of titanosaurian dinosaurs from the Late Cretaceous. It was named by John McIntosh in 1990. Opisthocoelicaudiines are known from Mongolia, Argentina, and the United States. Two genera were assigned to Opisthocoelicaudiinae by Gonzalez et al. (2009): Alamosaurus and Opisthocoelicaudia, a conclusion also found by Díez Díaz et al. (2018). The hands of opisthocoelicaudiines lacked wrist bones and phalanges.
Aeolosaurini is an extinct clade of titanosaurian dinosaurs known from the Cretaceous period of Argentina and Brazil. Rodrigo M. Santucci and Antonio C. de Arruda-Campos (2011) in their cladistic analysis found Aeolosaurus, Gondwanatitan, Maxakalisaurus, Panamericansaurus and Rinconsaurus to be aeolosaurids.
Saltasaurinae is a subfamily of titanosaurian sauropods known from the late Cretaceous period of South America, India and Madagascar.
Patagotitan is a genus of titanosaurian sauropod dinosaur from the Cerro Barcino Formation in Chubut Province, Patagonia, Argentina. The genus contains a single species known from at least six young adult individuals, Patagotitan mayorum, which was first announced in 2014 and then named in 2017 by José Carballido and colleagues. Preliminary studies and press releases suggested that Patagotitan was the largest known titanosaur and land animal overall, with an estimated length of 37 m (121 ft) and an estimated weight of 69 tonnes. Later research revised the length estimate down to 31 m (102 ft) and weight estimates down to approximately 50–57 tonnes, suggesting that Patagotitan was of a similar size to, if not smaller than, its closest relatives Argentinosaurus and Puertasaurus. Still, Patagotitan is one of the most-known titanosaurs, and so its interrelationships with other titanosaurs have been relatively consistent in phylogenetic analyses. This led to its use in a re-definition of the group Colossosauria by Carballido and colleagues in 2022.
Zhuchengtitan is a genus of titanosaurian sauropod dinosaur from the Late Cretaceous of Shandong, China. It contains a single species, Z. zangjiazhuangensis, named by Mo Jinyou and colleagues in 2017 from a single humerus. Zhuchengtitan can be identified by the extreme width of the top end of its humerus, as well as the expansion of the deltopectoral crest on its humerus; both of these characteristics indicate that it was likely closely related to Opisthocoelicaudia. However, it differs from the latter by the flatter bottom articulating surface of its humerus. Zhuchengtitan lived in a floodplain environment alongside Shantungosaurus, Zhuchengtyrannus, and Sinoceratops.