Lithostrotians
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Lithostrotia is a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
of derived
titanosaur Titanosaurs (or titanosaurians; members of the group Titanosauria) 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 thr ...
sauropods 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 ...
that lived during the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
and
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 ...
. The group was defined by Upchurch ''et al.'' in 2004 as the most recent common ancestor of ''
Malawisaurus ''Malawisaurus'' (meaning "Malawi lizard") is an extinct genus of titanosaurian sauropod dinosaur. It is known from the Dinosaur Beds of northern Malawi, which probably date to the Aptian stage of the Early Cretaceous. The type species is ''M. di ...
'' 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 ...
'' and all the descendants of that ancestor. Lithostrotia is derived from the
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
, meaning "inlaid with stones", referring to the fact that many known lithostrotians are preserved with
osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of Extant taxon, extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, Temnospondyli, ...
. However, osteoderms are not a distinguishing feature of the group, as the two noted by Unchurch ''et al.'' include
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 ...
with strongly concave front faces (procoely), although the farthest vertebrae are not procoelous.


History of research

In 1895, Richard Lydekker named the family Titanosauridae to summarize sauropods with procoelous (concave on the front) caudal vertebrae. 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 Phylogenetic nomenclature is a method of nomenclature for taxon, taxa in biology that uses phylogenetics, phylogenetic definitions for taxon names as explained below. This contrasts with Biological classification, the traditional method, by which ...
. According to a proposal by Wilson and Upchurch (2003) looks today much of the research on the use of that name from: Wilson and Upchurch published a revision of the genus ''
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 ...
'' 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. 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. The name Lithostrotia is not currently recognized by all researchers.


Definition and synapomorphies

Upchurch and colleagues (2004) define the Lithostrotia as a node-based taxon that includes the last common ancestor of ''
Malawisaurus ''Malawisaurus'' (meaning "Malawi lizard") is an extinct genus of titanosaurian sauropod dinosaur. It is known from the Dinosaur Beds of northern Malawi, which probably date to the Aptian stage of the Early Cretaceous. The type species is ''M. di ...
'' 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 ...
'' and all descendants of that ancestor. According to this definition the Lithostrotia includes all forms that are more derived than ''Malawisaurus'' in phylogenies. In addition to defining the group, Upchurch and colleagues gave two common derived features (
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ...
), which serve to distinguish the group from non-members. The first is that 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 ...
apart from the farthest
distal Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position provi ...
were procoelous, meaning their front face was
concave Concave or concavity may refer to: Science and technology * Concave lens * Concave mirror Mathematics * Concave function, the negative of a convex function * Concave polygon A simple polygon that is not convex is called concave, non-convex or ...
. Also, the front (proximal) caudal vertebrae were particularly strong procoelous. This first feature is also shared with
Mamenchisauridae Mamenchisauridae is a family of sauropod dinosaurs belonging to the Eusauropoda known from the Jurassic and Early Cretaceous of Asia and Africa. Mamenchisaurids are characterized by their proportionately extremely long necks. Some members of the ...
. Unchurch ''et al.'' named Lithostrotia based on the presence of
osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of Extant taxon, extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, Temnospondyli, ...
in many members, but the eponymous osteoderms do not represent
synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel Phenotypic trait, character or character state that has evolution, evolved from its ancestral form (or Plesiomorphy and symplesiomorphy, plesiomorphy). A synapomorphy is an apomorphy sh ...
, 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 ''Aeolosaurus'' (; "Aeolus' lizard") is a genus of titanosaurian sauropod dinosaur from the Late Cretaceous Period (geology), Period of what is now South America. Like most sauropods, it would have been a quadrupedal herbivore with a long neck an ...
'', ''
Ampelosaurus ''Ampelosaurus'' ( ; meaning "vine lizard") 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 ...
'', and various other genera both within and outside Lithostrotia with different morphologies.


Systematics

Lithostrotia is a derived group of titanosaurs, excluding primitive forms such as ''
Andesaurus ''Andesaurus'' ( ; "Andes lizard") is a genus of basal titanosaurian sauropod dinosaur which existed during the middle of the Cretaceous Period in South America. Like most sauropods, it would have had a small head on the end of a long neck and a ...
'' and '' Phuwiangosaurus''. The possibly equivalent clade Titanosauridae was positioned in a phylogenetic analysis by Calvo ''et al.'' (2007), where it included all titanosaurs apart from ''
Andesaurus ''Andesaurus'' ( ; "Andes lizard") is a genus of basal titanosaurian sauropod dinosaur which existed during the middle of the Cretaceous Period in South America. Like most sauropods, it would have had a small head on the end of a long neck and a ...
'', though multiple primitive forms were not analyzed. Other phylogenies, by Unchurch ''et al.'' (2015), instead have found a few, non-lithostrotian titanosaurs, or nearly all non-brachiosaurid titanosauriformes within the group. Poropat ''et al.'' (2015) conducted a similar analysis to one of Unchurch ''et al.'' (2015). This analysis found that ''Andesaurus'', ''
Argentinosaurus ''Argentinosaurus'' (meaning "lizard from Argentina") is a genus of giant sauropod dinosaur that lived during the Late Cretaceous Period (geology), period in what is now Argentina. Although it is only known from fragmentary remains, ''Argentinos ...
'' and ''
Epachthosaurus ''Epachthosaurus'' (meaning "heavy lizard") was a genus of titanosaurian sauropod dinosaur from the Late Cretaceous. It was a basal Lithostrotia, lithostrotian titanosaur. Its fossils have been found in Central and Northern Patagonia in South Ame ...
'' were within Titanosauria but outside Lithostrotia, and the latter group included ''
Malawisaurus ''Malawisaurus'' (meaning "Malawi lizard") is an extinct genus of titanosaurian sauropod dinosaur. It is known from the Dinosaur Beds of northern Malawi, which probably date to the Aptian stage of the Early Cretaceous. The type species is ''M. di ...
'', ''
Nemegtosaurus ''Nemegtosaurus'' (meaning 'Reptile from the Nemegt') was a sauropod dinosaur from the Late Cretaceous Period of what is now Mongolia. Measuring an estimated long and weighing , it was named after the Nemegt Basin in the Gobi Desert, where the ...
'', '' Diamantinasaurus'', '' Tapuiasaurus'' 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 ...
'' as basal lithostrotians outside
Saltasauridae 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 ...
. 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 ''Savannasaurus'' is a genus of titanosaurian sauropoda, 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 cladogram below follows Mocho ''et al.'' (2019) with the new subgroup called
Lirainosaurinae Lirainosaurinae is a subfamily of lithostrotian titanosaur sauropods from the Late Cretaceous of France, Spain, and Romania. Systematics Lirainosaurinae was defined by Díez Díaz ''et al.'' (2018) as comprising "'' Lirainosaurus'', ''Ampelosaur ...
.Mocho P, Páramo A, Escaso F, Marcos-Fernández F, Vidal D, Ortega F. 2019. Titanosaurs from Lo Hueco (Campanian-Maastrichtian) reveal new information about the evolutionary history of European titanosaurs, pp. 111. In: The Palaeontological Association (ed.), 63rd Annual Meeting, 15th–21st December 2019, University of Valencia, Spain, Programme Abstracts, AGM papers Poropat ''et al.'' (2016) Mocho ''et al.'' (2019) 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.


References

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