Protorosauria

Not to be confused with Protorosaurus.

Protorosaurs Temporal range: Latest Middle Permian - Late Triassic, 260–201.3 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Fossil specimen of Protorosaurus speneri, Teyler's Museum
Skeletal reconstructions of various members of Tanysauria, including Trachelosaurus fischeri, Dinocephalosaurus orientalis, Tanystropheus hydroides, and Tanystropheus longobardicus
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Clade:	Archosauromorpha
Order:	†Protorosauria Huxley, 1871
Subtaxa
†Czatkowiella †Exilisuchus? †Jesairosaurus †Malutinisuchus †Mecistotrachelos? †Megacnemus †Microcnemus †Protorosaurus †Rhombopholis † Tanysauria † Sharovipterygidae? †Trachelosauridae †Tanystropheidae

Protorosauria is an extinct, likely paraphyletic group of basal archosauromorph reptiles from the latest Middle Permian (Capitanian stage) to the end of the Late Triassic (Rhaetian stage) of Asia, Europe and North America. It was named by the English anatomist and paleontologist Thomas Henry Huxley in 1871 as an order, originally to solely contain Protorosaurus. Other names which were once considered equivalent to Protorosauria include Prolacertiformes and Prolacertilia.[1]

Protorosaurs are distinguished by their long necks formed by elongated cervical vertebrae, which have ribs that extend backward to the vertebrae behind them. Protorosaurs also have a gap between the quadrate bones and the jugal bones in the back of the skull near the jaw joint, making their skulls resemble those of lizards.[1] While previously thought to be monophyletic, the group is now thought to consist of various groups of basal archosauromorph reptiles that lie outside Crocopoda,[2] though some recent studies have recovered the group as monophyletic.[3] A number of members of Protosauria have been found to belong to a monophyletic group (though not including Protorosaurus) which was named Tanysauria in 2024.[4]

Classification

Protorosauria was considered to be a synonym of Prolacertiformes for many years.[5]

Since 1998, many phylogenetic analyses have found Protorosauria, as used in its widest sense, to be a polyphyletic or paraphyletic taxon. Protorosaurus, Macrocnemus, tanystropheids, and various other protorosaurs are usually placed near the base of Archosauromorpha, while Prolacerta and Pamelaria, two Gondwanan Triassic protorosaurs, are now thought to be in a more derived position as close relatives of Archosauriformes.[6] Most phylogenetic analyses since 1998 have found a strongly supported clade that includes only the genus Prolacerta and the Archosauriformes.[7]

For this reason Prolacerta, Pamelaria, and several other related forms (collectively called prolacertids) have been removed from Protorosauria. Because the name Prolacertiformes is defined based on the genus Prolacerta, the name Protorosauria is used for the remaining group.

Only recently has Protorosauria been defined in a phylogenetic sense as the most inclusive clade containing taxa such as Protorosaurus, Macrocnemus, and Tanystropheus. Analyses, such as Dilkes (1998), Sues (2003), Modesto & Sues (2004), Rieppel, Fraser & Nosotti (2003), Rieppel, Li & Fraser (2008), Gottmann-Quesada and Sander (2009) and Renesto et al. (2010),[7][8][9][10][11] recovered a large Protorosauria, that includes Protorosaurus, Drepanosauridae (and relatives) and Tanystropheidae (and relatives). However, some analysis found Protorosaurus (and sometimes the closely related Czatkowiella) to be more advanced[12] or more basal[13] than the node Drepanosauridae+Tanystropheidae, but always more basal than Prolacerta.

Some studies still use the term Prolacertiformes to include prolacertids and traditional protorosaurs, while restricting the term Protorosauria to the smallest clade that includes Protorosaurus, Macrocnemus, and Tanystropheus; thus Protorosauria is a true clade, while Prolacertiformes is an evolutionary grade of early archosauromorphs.[14]

Pritchard et al. (2015),[15] Nesbitt et al. (2015),[16] Ezcurra (2016)[17] and Spiekman et al., 2021[2] found that even this definition of Protorosauria, like Prolacertiformes, was an unnatural group of various non-Crocopodan archosauromorphs. These studies found that tanystropheids were archosauromorphs more closely related to crocopods than to Protorosaurus. Nevertheless, Ezcurra noted that archosauromorph systematics required further study, and that phylogenetic support for Protorosauria being a natural group was only barely weaker than the support for the group being unnatural.

Included groups

The Protorosauria includes the Permian genus Protorosaurus, closely related to Czatkowiella.[18] A wide variety of Permian and Triassic reptiles have been classified within Protorosauria, including the arboreal gliding reptile Sharovipteryx and the aquatic tanystropheids, which have extremely long necks.

Another enigmatic group of Triassic reptiles, the Drepanosauromorpha, have often been classified as belonging to the Protorosauria.[19]

Pterosaurs have also been proposed as protorosaurs or close relatives of them,[20] although they are now regarded as a more derived group of archosaurs.

While Senter (2004) reassigned the bizarre, arboreal drepanosaurids and Longisquama to a group of more primitive diapsids called Avicephala,[21] subsequent studies failed to find the same result, instead supporting the hypothesis that they were protorosaurs.

Cladogram

The following cladogram shows the position of Protorosauria among the Sauria sensu Sean P. Modesto and Hans-Dieter Sues (2004).[7]

Petrolacosaurus Youngina  Choristodera  Lazarussuchus Champsosaurus Cteniogenys  Sauria   Lepidosauromorpha  Gephyrosaurus Squamata  Archosauromorpha   Protorosauria  Protorosaurus  Drepanosauridae  Drepanosaurus Megalancosaurus  Tanystropheidae  Macrocnemus Langobardisaurus Tanystropheus longobardicus Trilophosaurus  Rhynchosauria  Howesia Mesosuchus Prolacerta  Archosauriformes  Proterosuchus Euparkeria

Most recent studies have recovered Protorosauria as a whole as a paraphyletic, cladogram after Spiekman et al. 2021[2]

Diapsida

Araeoscelidia Neodiapsida Orovenator Claudiosaurus Acerosodontosaurus Youngina Sauria Lepidosauromorpha Archosauromorpha Jesairosaurus Protorosaurus Prolacerta Czatkowiella Sharovipterygidae (Ozimek) Fuyuansaurus Dinocephalosauridae Protanystropheus Pectodens Dinocephalosaurus Tanystropheidae Macrocnemus bassanii Macrocnemus fuyuanensis Augustaburiania Amotosaurus Langobardisaurus AMNH FARB 7206 Sclerostropheus Tanystropheus hydroides GMPKU P 1527 T. cf. hydroides Tanystropheus longobardicus Crocopoda Allokotosauria Rhynchosauria Teyujagua Archosauriformes "Protorosauria"

Although Protorosauria as a whole is often found to be a paraphyletic, a large group of former "protorosaurs" (excluding Protorosaurus) is frequently found to be monophyletic. This clade was given the name "Tanysauria" by Spiekman et al. in 2024.[4]

	Archosauromorpha Protorosaurus Aenigmastropheus Tanysauria Jesairosaurus Trachelosauridae Austronaga Pectodens Fuyuansaurus Trachelosaurus Gracilicollum Dinocephalosaurus Tanystropheidae Augustaburiania Luxisaurus Macrocnemus bassanii Macrocnemus fuyuanensis Macrocnemus obristi UFRGS-PV-492-T Amotosaurus Langobardisaurus Raibliania Tanystropheus spp. Sclerostropheus Ozimek Tanytrachelos Crocopoda

References

1. Sues, H.-D.; Fraser, N.C. (2010). "Early and early Middle Triassic in Gondwana". Triassic Life on Land: The Great Transition. New York: Columbia University Press. ISBN 9780231135221.
2. Spiekman, Stephan N. F.; Fraser, Nicholas C.; Scheyer, Torsten M. (2021-05-03). "A new phylogenetic hypothesis of Tanystropheidae (Diapsida, Archosauromorpha) and other "protorosaurs", and its implications for the early evolution of stem archosaurs". PeerJ. 9: e11143. doi:10.7717/peerj.11143. ISSN 2167-8359. PMC 8101476. PMID 33986981.
3. Simões, T. R.; Kammerer, C. F.; Caldwell, M. W.; Pierce, S. E. (2022). "Successive climate crises in the deep past drove the early evolution and radiation of reptiles". Science Advances. 8 (33): eabq1898. doi:10.1126/sciadv.abq1898. PMC 9390993. PMID 35984885.
4. Spiekman, Stephan N. F.; Ezcurra, Martín D.; Rytel, Adam; Wang, Wei; Mujal, Eudald; Buchwitz, Michael; Schoch, Rainer R. (2024-03-15). "A redescription of Trachelosaurus fischeri from the Buntsandstein (Middle Triassic) of Bernburg, Germany: the first European Dinocephalosaurus-like marine reptile and its systematic implications for long-necked early archosauromorphs". Swiss Journal of Palaeontology. 143 (1): 10. doi:10.1186/s13358-024-00309-6. ISSN 1664-2384.
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