Lithostrotia

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Lithostrotians
Temporal range: Early to Late Cretaceous, 120–66  Ma
Saltasaurus dinosaur.png
Life restorations of a pair of Saltasaurus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Sauropodomorpha
Clade: Sauropoda
Clade: Macronaria
Clade: Titanosauria
Clade: Lithostrotia
Upchurch et al., 2004
Subgroups [1] [2] [3]

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.

Contents

History of research

In 1895, Richard Lydekker named the family Titanosauridae to summarize sauropods with procoelous (concave on the front) caudal vertebrae. [8] The name Titanosauridae has since been widely used, and was defined by Salgado and colleagues (1997), Gonzalaz-Riga (2003), and Salgado (2003) as a node-based taxon. [9] According to a proposal by Wilson and Upchurch (2003) looks today much of the research on the use of that name from[ clarification needed ]: Wilson and Upchurch published a revision of the genus Titanosaurus and declare the type species Titanosaurus indicus as invalid because it is based only on two vertebrae of the tail, showing no diagnostically usable features. Consistently these authors consider ranking groups that are based on Titanosaurus as the nominal taxon, Titanosauridae, Titanosaurinae and Titanosauroidea - also considered invalid. [8] In 2004 Upchurch and colleagues presented the new group Lithostrotia to describe the same group as Titanosauridae, but instead it was not based upon a specific taxon. [10] The name Lithostrotia is not currently recognized by all researchers. [11]

Definition and synapomorphies

Caudal vertebrae of Malawisaurus (top right) and Saltasaurus (bottom) compared with Tornieria Sauropod caudal pneumaticity.png
Caudal vertebrae of Malawisaurus (top right) and Saltasaurus (bottom) compared with Tornieria

Upchurch and colleagues (2004) define the Lithostrotia as a node-based taxon that includes the last common ancestor of Malawisaurus and Saltasaurus and all descendants of that ancestor. According to this definition the Lithostrotia includes all forms that are more derived than Malawisaurus in phylogenies. [10]

In addition to defining the group, Upchurch and colleagues gave two common derived features (synapomorphies), which serve to distinguish the group from non-members. The first is that all caudal vertebrae apart from the farthest distal were procoelous, meaning their front face was concave. Also, the front (proximal) caudal vertebrae were particularly strong procoelous. This first feature is also shared with Mamenchisauridae. [10]

Unchurch et al. named Lithostrotia based on the presence of osteoderms in many members, but the eponymous osteoderms do not represent synapomorphy, as the evolutionary history of osteoderms is unknown within the titanosaurs. It may be this trait has developed multiple times independently within the titanosaurs and Lithostrotia, as osteoderms are known in many saltasaurids, Mendozasaurus , Aeolosaurus , Ampelosaurus , and various other genera both within and outside Lithostrotia with different morphologies. [12]

Systematics

Malawisaurus skeleton, Royal Ontario Museum August 1, 2012 - Malawisaurus on Display at the Royal Ontario Museum.jpg
Malawisaurus skeleton, Royal Ontario Museum

Lithostrotia is a derived group of titanosaurs, excluding primitive forms such as Andesaurus and Phuwiangosaurus . [10] The possibly equivalent clade Titanosauridae was positioned in a phylogenetic analysis by Calvo et al. (2007), where it included all titanosaurs apart from Andesaurus , though multiple primitive forms were not analyzed. [11] Other phylogenies, by Unchurch et al. (2015), instead have found a few, non-lithostrotian titanosaurs, or nearly all non-brachiosaurid titanosauriformes within the group. [13] Poropat et al. (2015) conducted a similar analysis to one of Unchurch et al. (2015). This analysis found that Andesaurus, Argentinosaurus and Epachthosaurus were within Titanosauria but outside Lithostrotia, and the latter group included Malawisaurus , Nemegtosaurus , Diamantinasaurus , Tapuiasaurus and Alamosaurus as basal lithostrotians outside Saltasauridae. [14] Another phylogenetic analysis by Poropat and colleagues in 2016, partially reproduced below, found Diamantinasaurus as a non-lithostrotian titanosaur and the sister taxon of the contemporary Savannasaurus . [15] [16] The cladogram below follows Mocho et al. (2019) with the new subgroup called Lirainosaurinae. [17]

While the clade Lithostrotia has been used in many different phylogenetic analyses as a description for a clade of titanosaurs, its use is not universal. The clade has been omitted from results as it has been considered a synonym of Titanosauria, or it has been used as either a very large clade encompassing almost all titanosaurs, or a smaller clade that excludes the taxa of Colossosauria. These incongruent results are because of the instability of the defining taxon Malawisaurus, which may in fact represent a chimaeric assemblage of remains and requires re-evaluation to determine what Lithostrotia includes. [1]

Related Research Articles

<span class="mw-page-title-main">Titanosauria</span> Extinct clade of dinosaurs

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.

<span class="mw-page-title-main">Nemegtosauridae</span> Extinct family of dinosaurs

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.

<i>Aeolosaurus</i> Extinct genus of dinosaurs

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.

<i>Andesaurus</i> Extinct genus of dinosaurs

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.

<i>Sanpasaurus</i> Extinct species of reptile

Sanpasaurus is a poorly known sauropod dinosaur from the Early to Late Jurassic of Sichuan, China. The type species, S. yaoi, was described by Chung Chien Young, in 1944. The type remains, IVPP V.156, consists of 20 vertebrae, scapulae, forelimbs, and some hindlimb bones. Initially reported by Young as an ornithopod ornithischian, this specimen was unambiguously referred to Sauropoda in 2016 by McPhee et al., later refined to a basal gravisaurian position by Pol et al. in 2020 and 2022, closely related to Vulcanodon and Tazoudasaurus. Sanpasaurus is known from remains recovered from the Maanshan Member of the Ziliujing Formation.

<i>Neuquensaurus</i> Extinct genus of dinosaurs

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.

<i>Pellegrinisaurus</i> Extinct genus of dinosaurs

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.

Malarguesaurus is a genus of titanosauriform sauropod dinosaur from the Late Cretaceous of Mendoza Province, Argentina. Its fossils, consisting of tail vertebrae, chevrons, ribs, and limb bones, were found in the upper Turonian-lower Coniacian Portezuelo Formation of the Neuquén Group. The type species, described by González Riga et al. in 2008, is M. florenciae.

<i>Diamantinasaurus</i> Extinct genus of reptiles

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.

<i>Xianshanosaurus</i> Extinct genus of dinosaurs

Xianshanosaurus is a genus of sauropod dinosaur from the Early Cretaceous (Aptian-Albian) of the Ruyang Basin in Henan Province, China. Its type and only species is Xianshanosaurus shijiagouensis. It was described in 2009 by a team of paleontologists led by Lü Junchang. Xianshanosaurus may be a titanosaur, and Daxiatitan may be its closest relative, but its evolutionary relationships remain controversial.

<span class="mw-page-title-main">Lognkosauria</span> Clade of titanosaurian sauropods

Lognkosauria is a clade of giant long-necked sauropod dinosaurs within the clade Titanosauria. It includes some of the largest and heaviest dinosaurs known. They lived in South America and likely Asia during the Late Cretaceous period.

<span class="mw-page-title-main">Aeolosaurini</span> Extinct clade of dinosaurs

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.

<span class="mw-page-title-main">Eutitanosauria</span>

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.

<i>Patagotitan</i> Extinct genus of dinosaurs

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.

<i>Savannasaurus</i> Extinct genus of dinosaurs

Savannasaurus is a genus of titanosaurian sauropod dinosaur from the Late Cretaceous Winton Formation of Queensland, Australia. It contains one species, Savannasaurus elliottorum, named in 2016 by Stephen Poropat and colleagues. The holotype and only known specimen, originally nicknamed "Wade", is the most complete specimen of an Australian sauropod, and is held at the Australian Age of Dinosaurs museum. Dinosaurs known from contemporary rocks include its close relative Diamantinasaurus and the theropod Australovenator; associated teeth suggest that Australovenator may have fed on the holotype specimen.

Triunfosaurus is a genus of somphospondylan sauropod dinosaur from the Early Cretaceous of Brazil. It contains a single species, T. leonardii, described by Carvalho et al. in 2017. As a genus, Triunfosaurus can be distinguished from all other titanosaurs by the unique proportions of its ischium. It was initially described as a basal titanosaur, making it the earliest basal titanosaur known; however, subsequent research questioned the identification of the taxon as a titanosaur, instead reassigning it to the Somphospondyli.

<span class="mw-page-title-main">Rinconsauria</span> Extinct clade of dinosaurs

Rinconsauria is an extinct clade of giant titanosaurian sauropods known from the late Cretaceous period of Argentina.

<span class="mw-page-title-main">Colossosauria</span> Extinct clade of dinosaurs

Colossosauria is a clade of titanosaur sauropods from the latest Early Cretaceous through the Late Cretaceous of South America. The group was originally named by Bernardo González-Riga et al. in 2019 and defined as the "most inclusive clade containing Mendozasaurus neguyelap but not Saltasaurus loricatus or Epachthosaurus sciuttoi". The clade contains different taxa depending on the phylogenetic analysis used, in the defining paper the only subgroups were Rinconsauria and Lognkosauria, but alternate phylogenies published previously had also included various similar titanosaurs such as Aeolosaurus, Bonitasaura, Drusilasaura, Overosaurus and Quetecsaurus. The phylogenetic analysis of González-Riga et al. (2019) placed Colossosauria as sister taxa to Epachthosaurus, Pitekunsaurus and a larger clade including Saltasauridae.

<span class="mw-page-title-main">Diamantinasauria</span> Clade of titanosaurian sauropod dinosaurs

Diamantinasauria is an extinct clade of non-lithostrotian titanosaurian sauropod dinosaurs, known from the early Late Cretaceous (Cenomanian-Turonian) of South America and Australia. It was named by Poropat and colleagues in 2021, and contains four genera: Australotitan, Savannasaurus and Diamantinasaurus from the Winton Formation of Queensland, as well as Sarmientosaurus from the Bajo Barreal Formation of Patagonia. The existence of the clade indicates connectivity between Australia and South America via Antarctica during the Cretaceous period.

<span class="mw-page-title-main">Saltasauroidea</span>

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.

References

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