Lithostrotia

2025-06-27T16:57:20Z

92.46.7.6: If ninjatitan was Lithostrotia probably they originated in 140 mya least

{{Short description|Extinct clade of dinosaurs}}
{{automatic taxobox
|name = Lithostrotians
|fossil_range = [[Early Cretaceous|Early]] to [[Late Cretaceous]], {{fossil range|120|earliest=140|66}}
|image = Saltasaurus dinosaur.png
|image_caption = Life restorations of a pair of ''Saltasaurus''
|taxon = Lithostrotia
|authority = Upchurch ''et al.'', 2004
|subdivision_ranks = Subgroups
|subdivision_ref=<ref name="carballido2022">{{cite book|last1=Carballido|first1=J.L.|last2=Otero|first2=A.|last3=Mannion|first3=P.D.|last4=Salgado|first4=L.|last5=Moreno|first5=A.P.|year=2022|chapter=Titanosauria: A Critical Reappraisal of Its Systematics and the Relevance of the South American Record|pages=269–298|editor-last=Otero|editor-first=A.|editor2-last=Carballido|editor2-first=J.L.|editor3-last=Pol|editor3-first=D.|title=South American Sauropodomorph Dinosaurs. Record, Diversity and Evolution|publisher=Springer|isbn=978-3-030-95958-6|doi=10.1007/978-3-030-95959-3|issn=2197-9596}}</ref><ref name="gallina2022">{{cite book|last1=Gallina|first1=P.A.|last2=González Riga|first2=B.J.|last3=Ortiz David|first3=L.D.|year=2022|chapter=Time for Giants: Titanosaurs from the Berriasian–Santonian Age|pages=299–340|editor-last=Otero|editor-first=A.|editor2-last=Carballido|editor2-first=J.L.|editor3-last=Pol|editor3-first=D.|title=South American Sauropodomorph Dinosaurs. Record, Diversity and Evolution|publisher=Springer|isbn=978-3-030-95958-6|doi=10.1007/978-3-030-95959-3|issn=2197-9596}}</ref><ref name="santucci2022">{{cite book|last1=Santucci|first1=R.M.|last2=Filippi|first2=L.S.|year=2022|chapter=Last Titans: Titanosaurs From the Campanian–Maastrichtian Age|pages=341–391|editor-last=Otero|editor-first=A.|editor2-last=Carballido|editor2-first=J.L.|editor3-last=Pol|editor3-first=D.|title=South American Sauropodomorph Dinosaurs. Record, Diversity and Evolution|publisher=Springer|isbn=978-3-030-95958-6|doi=10.1007/978-3-030-95959-3|issn=2197-9596}}</ref>
|subdivision =
*{{extinct}}''[[Arackar]]''
*{{extinct}}''[[Atacamatitan]]''
*{{extinct}}''[[Elaltitan]]''
*{{extinct}}''[[Epachthosaurus]]''
*{{extinct}}''[[Isisaurus]]''
*{{extinct}}''[[Malawisaurus]]''
*{{extinct}}''[[Mansourasaurus]]''
*{{extinct}}''[[Mnyamawamtuka]]''<ref name="gorscak19"/>
*{{extinct}}''[[Normanniasaurus]]''<ref name="gorscak19">{{cite journal|last1=Gorscak|first1=E.|last2=O'Connor|first2=P.|year=2019|title=A new African Titanosaurian Sauropod Dinosaur from the middle Cretaceous Galula Formation (Mtuka Member), Rukwa Rift Basin, Southwestern Tanzania|journal=PLOS ONE|volume=14|issue=2|pages=e0211412|doi=10.1371/journal.pone.0211412|doi-access=free|pmid=30759122 |pmc=6374010|bibcode=2019PLoSO..1411412G }}</ref>
*{{extinct}}''[[Paralititan]]''
*{{extinct}}''[[Petrobrasaurus]]''
*{{extinct}}''[[Rapetosaurus]]''
*{{extinct}}''[[Rukwatitan]]''<ref name="gorscak19"/>
*{{extinct}}''[[Sonidosaurus]]''<ref name="averianov2017">{{cite journal|last1=Averianov|first1=A.O.|last2=Sues|first2=H.D.|year=2017|title=Review of Cretaceous sauropod dinosaurs from Central Asia|journal=Cretaceous Research|volume=69|pages=184–197|doi=10.1016/j.cretres.2016.09.006|doi-access=free|bibcode=2017CrRes..69..184A }}</ref>
*{{extinct}}''[[Tapuiasaurus]]''
*{{extinct}}''[[Uriash]]''
*{{extinct}}''[[Volgatitan]]''<ref name="averianov2018">{{cite journal|last1=Averianov|first1=A.O.|last2=Efimov|first2=V.|year=2018|title=The oldest titanosaurian sauropod of the Northern Hemisphere|journal=Biological Communications|volume=63|issue=6|pages=145–162|doi=10.21638/spbu03.2018.301|doi-access=free|hdl=11701/15099|hdl-access=free}}</ref>
*{{extinct}}[[Lirainosaurinae]]
*{{extinct}}'''[[Eutitanosauria]]'''
}}

'''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]] {{Transliteration|grc|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.

==History of research==
In 1895, Richard Lydekker named the family Titanosauridae to summarize sauropods with procoelous (concave on the front) caudal vertebrae.<ref name="wilson03"/> 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]].<ref name="titanosauridae-ts">{{cite web |url=http://www.taxonsearch.org/dev/taxon_edit.php?Action=View&tax_id=316 |title=Titanosauridae |author=Paul Sereno |work=Taxon Search |access-date=2014-08-28 |url-status=dead |archive-url=https://web.archive.org/web/20140903154246/http://www.taxonsearch.org/dev/taxon_edit.php?tax_id=316&Action=View |archive-date=2014-09-03 }}</ref> According to a proposal by Wilson and Upchurch (2003) looks today much of the research on the use of that name from{{clarify|date=November 2022}}: 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.<ref name="wilson03">{{cite journal|last1=Wilson|first1=J.A.|last2=Upchurch|first2=P.|date=2003|title=A revision of ''Titanosaurus'' Lydekker (Dinosauria – Sauropoda), the first dinosaur genus with a "Gondwanan" distribution|journal=Journal of Systematic Palaeontology|volume=1|issue=3|pages=125–160|issn=1477-2019|doi=10.1017/S1477201903001044|bibcode=2003JSPal...1..125W |s2cid=53997295}}</ref> 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.<ref name="upchurch04">{{cite book|last1=Upchurch|first1=P.|last2=Barrett|first2=P.M.|last3=Dodson|first3=P.|date=2004|chapter=Sauropoda|editor-last=Weishampel|editor-first=David B.|editor2-last=Dodson|editor2-first=Peter|editor3-last=Osmolska|editor3-first=H.|title=The Dinosauria|publisher=University of California Press|pages=259–322|edition=2nd}}</ref> The name Lithostrotia is not currently recognized by all researchers.<ref name="calvo07">{{cite journal|last1=Calvo|first1=J.O.|last2=Porfiri|first2=J.D.|last3=González-Riga|first3=B.J.|last4=Kellner|first4=A.W.|date=2007|title=A new Cretaceous terrestrial ecosystem from Gondwana with the description of a new sauropod dinosaur|journal=Anais da Academia Brasileira de Ciências|volume=79|issue=3|pages=529–41|doi=10.1590/S0001-37652007000300013|pmid=17768539|doi-access=free}}</ref>

==Definition and synapomorphies==
[[File:Sauropod caudal pneumaticity.png|thumb|left|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 trait|derived]] than ''Malawisaurus'' in phylogenies.<ref name="upchurch04"/>

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 [[anatomical terms of location#Proximal and distal|distal]] were procoelous, meaning their front face was [[wikt:concave|concave]]. Also, the front (proximal) caudal vertebrae were particularly strong procoelous. This first feature is also shared with [[Mamenchisauridae]].<ref name="upchurch04"/>

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.<ref name="d_Emic_2009">{{cite journal|last1=D'Emic|first1=M.D.|last2=Wilson|first2=J.A.|last3=Chatterjee|first3=S.|date=2009|title=The titanosaur (Dinosauria: Sauropoda) osteoderm record: review and first definitive specimen from India|journal=Journal of Vertebrate Paleontology|volume=29|issue=1|issn=0272-4634|pages=165–177|doi=10.1671/039.029.0131|bibcode=2009JVPal..29..165D |s2cid=85991253}}</ref>

==Systematics==
[[File:August 1, 2012 - Malawisaurus on Display at the Royal Ontario Museum.jpg|thumb|''Malawisaurus'' skeleton, [[Royal Ontario Museum]]]]
Lithostrotia is a derived group of titanosaurs, excluding primitive forms such as ''[[Andesaurus]]'' and ''[[Phuwiangosaurus]]''.<ref name="upchurch04"/> 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.<ref name="calvo07"/> Other phylogenies, by Unchurch ''et al.'' (2015), instead have found a few, non-lithostrotian titanosaurs, or nearly all non-brachiosaurid titanosauriformes within the group.<ref name="upchurch15">{{cite journal|doi=10.1371/journal.pone.0125819|pmid=26039587|title=The Anatomy and Phylogenetic Relationships of "Pelorosaurus" becklesii (Neosauropoda, Macronaria) from the Early Cretaceous of England|journal=PLOS ONE|volume=10|issue=6|pages=e0125819|year=2015|last1=Upchurch|first1=Paul|last2=Mannion|first2=Philip D.|last3=Taylor|first3=Michael P.|pmc=4454574|bibcode=2015PLoSO..1025819U|doi-access=free}}</ref> 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]].<ref name="poropatetal14">{{cite journal|last1=Poropat|first1=S.F.|last2=Upchurch|first2=P.|last3=Mannion|first3=P.D.|last4=Hocknull|first4=S.A.|last5=Kear|first5=B.P.|last6=Sloan|first6=T.|last7=Sinapius|first7=G.H.K.|last8=Elliot|first8=D.A.|year=2014|title=Revision of the sauropod dinosaur ''Diamantinasaurus matildae'' Hocknull et al. 2009 from the mid-Cretaceous of Australia: Implications for Gondwanan titanosauriform dispersal|journal=Gondwana Research|volume=27|issue=3|pages=995–1033|doi=10.1016/j.gr.2014.03.014|doi-access=free|hdl=10044/1/27497|hdl-access=free}}</ref> 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]]''.<ref name="savannasaurus2016">{{cite journal | first1 = S.F. | last1 = Poropat | first2 = P.D. | last2 = Mannion | first3 = P. | last3 = Upchurch | first4 = S.A. | last4 = Hocknull | first5 = B.P. | last5 = Kear | first6 = M. | last6 = Kundrát | first7 = T.R. | last7 = Tischler | first8 = T. | last8 = Sloan | first9 = G.H.K. | last9 = Sinapius | first10 = J.A. | last10 = Elliott | first11 = D.A. | last11 = Elliott | title = New Australian sauropods shed light on Cretaceous dinosaur palaeobiogeography | journal = Scientific Reports | volume = 6 | pages = 34467 | date = 2016 | doi = 10.1038/srep34467 | pmid=27763598 | pmc=5072287| bibcode = 2016NatSR...634467P }}</ref><ref name="NYT-20161020">{{cite news |last=St. Fleur |first=Nicholas |title=Meet the New Titanosaur. You Can Call It Wade. |url=https://www.nytimes.com/2016/10/21/science/dinosaur-australia-titanosaur.html |date=20 October 2016 |work=[[New York Times]] |access-date=21 October 2016 }}</ref>
The cladogram below follows Mocho ''et al.'' (2019) with the new subgroup called [[Lirainosaurinae]].<ref>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</ref>
{{col-begin|width=75%}}
{{col-break}}
'''Poropat ''et al.'' (2016)'''
{{clade|style=font-size:85%; line-height:85%
|label1=[[Titanosaur]]ia
|1={{clade
|1=''[[Andesaurus]]''
|2={{clade
|1={{clade
|1=''[[Dongyangosaurus]]''
|2=''[[Baotianmansaurus]]'' }}
|2={{clade
|1=''[[Ligabuesaurus]]''
|2={{clade
|1={{clade
|1=''[[Savannasaurus]]''
|2=''[[Diamantinasaurus]]''}}
|2={{clade
|label2='''Lithostrotia'''
|1={{clade
|1=''[[Xianshanosaurus]]''
|2=''[[Daxiatitan]]''}}
|2={{clade
|1=''[[Malawisaurus]]''
|2={{clade
|1={{clade
|1=''[[Muyelensaurus]]''
|2={{clade
|1=''[[Futalognkosaurus]]''
|2=''[[Epachthosaurus]]'' }}}}
|2={{clade
|label1 = [[Nemegtosauridae]]
|1={{clade
|1=''[[Tapuiasaurus]]''
|2=''[[Nemegtosaurus]]'' }}
|2={{clade
|1=''[[Isisaurus]]''
|label2=[[Saltasauridae]]
|2={{clade
|1=''[[Saltasaurus]]''
|2={{clade
|1=''[[Opisthocoelicaudia]]''
|2={{clade
|1=''[[Jiangshanosaurus]]''
|2=''[[Alamosaurus]]'' }} }} }} }} }} }} }} }} }} }} }} }} }}
{{col-break}}
'''Mocho ''et al.'' (2019)'''
{{clade|style=font-size:85%; line-height:85%;
|label1='''Lithostrotia'''
|1={{clade
|1=''[[Malawisaurus]]''
|2={{clade
|1={{clade
|1=''[[Paludititan]]''
|2=''[[Lohuecotitan]]'' }}
|2={{clade
|1=''[[Epachthosaurus]]''
|2={{clade
|1={{clade
|label1=[[Opisthocoelicaudiinae]]
|1={{clade
|1=''[[Alamosaurus]]''
|2=''[[Opisthocoelicaudia]]'' }} }}
|2={{clade
|1={{clade
|label1=[[Saltasaurinae]]
|1={{clade
|1=''[[Neuquensaurus]]''
|2={{clade
|1=''[[Rocasaurus]]''
|2=''[[Saltasaurus]]'' }} }} }}
|2={{clade
|1={{clade
|label1=[[Lirainosaurinae]]
|1={{clade
|1=''[[Lirainosaurus]]''
|2={{clade
|1=''[[Atsinganosaurus]]''
|2=''[[Ampelosaurus]]'' }} }} }}
|2={{clade
|1=''[[Bonatitan]]''
|2={{clade
|1={{clade
|1=''[[Rapetosaurus]]''
|2=''[[Nemegtosaurus]]'' }}
|2={{clade
|label1=[[Aeolosaurinae]]
|1={{clade
|1=''[[Gondwanatitan]]''
|2=''[[Aeolosaurus]]'' }}
|label2=[[Rinconsauria]]
|2={{clade
|1=''[[Rinconsaurus]]''
|2={{clade
|1=''[[Muyelensaurus]]''
|2={{clade
|1=''[[Bonitasaura]]''
|label2=[[Lognkosauria]]
|2={{clade
|1=''[[Mendozasaurus]]''
|2=''[[Futalognkosaurus]]'' }} }} }} }} }} }} }} }} }} }} }} }} }} }}
{{col-end}}

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.<ref name="carballido2022">{{cite book|last1=Carballido|first1=J.L.|last2=Otero|first2=A.|last3=Mannion|first3=P.D.|last4=Salgado|first4=L.|last5=Moreno|first5=A.P.|year=2022|chapter=Titanosauria: A Critical Reappraisal of Its Systematics and the Relevance of the South American Record|pages=269–298|editor-last=Otero|editor-first=A.|editor2-last=Carballido|editor2-first=J.L.|editor3-last=Pol|editor3-first=D.|title=South American Sauropodomorph Dinosaurs. Record, Diversity and Evolution|publisher=Springer|isbn=978-3-030-95958-6|doi=10.1007/978-3-030-95959-3|issn=2197-9596}}</ref>

==References==
{{Reflist}}

{{Sauropodomorpha|T.}}
{{Taxonbar|from=Q140061}}
{{Portal bar|Dinosaurs}}

[[Category:Lithostrotia| ]]
[[Category:Dinosaur clades]]
[[Category:Cretaceous dinosaurs]]

wiki143 - User contributions [en]

Lithostrotia