Crurotarsi

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Crurotarsans
Temporal range:
Early Triassic–Present, 250–0 Ma

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Protome batalaria.jpg
Life restoration of Protome batalaria

Ornithosuchus BW.jpg
Life restoration of Ornithosuchus woodwardi

Scientific classification e
Kingdom:
Animalia
Phylum:
Chordata
Class:
Reptilia

Clade:

Archosauriformes

Clade:

Crurotarsi
Sereno & Arcucci, 1990
Subgroups


  • Phytosauria


  • Archosauria
    • Avemetatarsalia

    • Pseudosuchia


Crurotarsi is a group of archosauriform reptiles that includes the archosaurs (represented today by birds and crocodilians) and the extinct, crocodile-like phytosaurs.[1] The name is derived from the Latin word crus and the Greek word tarsos; it refers to the specialized articulation between crus and tarsus—specifically between fibula and calcaneum—present in the skeletons of suchians and phytosaurs, with a hemicylindrical condyle on the calcaneum articulating against fibula.[2][3]




Contents





  • 1 Taxonomic history


  • 2 Phylogeny


  • 3 References


  • 4 External links




Taxonomic history


The name Crurotarsi was erected as a node-based clade by Paul Sereno and A. B. Arcucci in 1990 to supplant the old term Pseudosuchia, but with a different definition.[2] Crurotarsi includes, by most published definitions, all descendants of the common ancestor of modern crocodiles, ornithosuchids, aetosaurs, and phytosaurs; Nesbitt (2011) provided a shorter definition, defining Crurotarsi as "the least inclusive clade containing Rutiodon carolinensis Emmons, 1856, and Crocodylus niloticus Laurenti, 1768".[1] According to two studies published in 2011 by Nesbitt and coworkers, using either of these definitions leads to the inclusion of all other true archosaurs in Crurotarsi, due to the possibly basal phylogenetic position of the phytosaurs. This means that grouping the phytosaurs and crocodilians into a clade while excluding the avemetatarsalians (pterosaurs, dinosaurs, and birds) would result in a paraphyletic grouping. A more definitive group is Pseudosuchia, which is defined as all archosaurs closer to crocodiles than to birds (matching the traditional content of Crurotarsi).[1][4]



Phylogeny


Paul Sereno and A. B. Arcucci named Crurotarsi in 1990, defining it as "Parasuchia [phytosaurs], Ornithosuchidae, Prestosuchus, Suchia, and all descendants of their common ancestor".[2] The groups in this definition were considered crocodile-line archosaurs, as opposed to the bird-line archosaurs. Ornithosuchids were once considered bird-line archosaurs (as implied by their name, which means "bird crocodiles" in Greek), but were later recognized as crocodile-line archosaurs. This reclassification may have inspired Sereno's Crurotarsi, a node-based clade defined by the inclusion of ornithosuchids and other early archosaurs.


Two names were proposed for crocodile-line archosaurs before Crurotarsi was erected. The first, Pseudosuchia, was established as a stem-based clade in 1985.[5] It includes crocodiles and all archosaurs more closely related to crocodiles than to birds. The second, Crocodylotarsi, was named in 1988, possibly as a replacement for Pseudosuchia.[6] The name Pseudosuchia, meaning "false crocodiles", has been used for over a century, and traditionally included aetosaurs. As a clade, Pseudosuchia includes the group Eusuchia, or "true crocodiles". Crocodylotarsi may have been named to remove confusion, but as a stem-based clade it is synonymous with Pseudosuchia. Because Pseudosuchia was named first, it has precedence. Crurotarsi traditionally contains the same archosaurs as Pseudosuchia, but as a node-based clade it is not synonymous.[7]


In 2011, Sterling J. Nesbitt found phytosaurs to be the sister taxon of Archosauria, and therefore not crocodile-line archosaurs. Because phytosaurs are included in the definition of Crurotarsi, this change in their phylogenetic placement expanded the scope of Crurotarsi, which therefore now includes phytosaurs, crocodiles, pterosaurs and dinosaurs. However, Pseudosuchia still contains only crocodile-line archosaurs.


Below is a cladogram modified from Nesbitt (2011) showing the new changes:[1]


.mw-parser-output table.cladeborder-spacing:0;margin:0;font-size:100%;line-height:100%;border-collapse:separate;width:auto.mw-parser-output table.clade table.cladewidth:100%.mw-parser-output table.clade tdborder:0;padding:0;vertical-align:middle;text-align:center.mw-parser-output table.clade td.clade-labelwidth:0.8em;border:0;padding:0 0.2em;vertical-align:bottom;text-align:center.mw-parser-output table.clade td.clade-slabelborder:0;padding:0 0.2em;vertical-align:top;text-align:center.mw-parser-output table.clade td.clade-barvertical-align:middle;text-align:left;padding:0 0.5em.mw-parser-output table.clade td.clade-leafborder:0;padding:0;text-align:left;vertical-align:middle.mw-parser-output table.clade td.clade-leafRborder:0;padding:0;text-align:right



Archosauriformes 








†ProterosuchidaeProterosuchusDB flipped.jpg












†ErythrosuchidaeErythrosuchus africanus.jpg












VancleaveaVancleavea white background.jpg












†ProterochampsiaChanaresuchus.jpg












EuparkeriaEuparkeria BW flipped.jpg



 Crurotarsi 








†PhytosauriaSmilosuchus adamanensis flipped.jpg



 Archosauria 








Avemetatarsalia (bird-line archosaurs)Meyers grosses Konversations-Lexikon - ein Nachschlagewerk des allgemeinen Wissens (1908) (Antwerpener Breiftaube).jpg



 Pseudosuchia (crocodile-line archosaurs) 








†OrnithosuchidaeOrnithosuchus BW white background.jpg





SuchiaDescription des reptiles nouveaux, ou, Imparfaitement connus de la collection du Muséum d'histoire naturelle et remarques sur la classification et les caractères des reptiles (1852) (Crocodylus moreletii).jpg











Below is a cladogram after Nesbitt & Norell (2006) and Nesbitt (2007) with Crurotarsi in its traditional sense encompassing just crocodile-line archosaurs:[8][9]






Archosauriformes 








EuparkeriaEuparkeria BW flipped.jpg












†ProterochampsidaeChanaresuchus.jpg



 Archosauria 








to AvemetatarsaliaMeyers grosses Konversations-Lexikon - ein Nachschlagewerk des allgemeinen Wissens (1908) (Antwerpener Breiftaube).jpg




Crurotarsi 








†PhytosauriaSmilosuchus adamanensis flipped.jpg



 Suchia 








†AetosauriaDesmatosuchus spurensis flipped.jpg












CrocodylomorphaDescription des reptiles nouveaux, ou, Imparfaitement connus de la collection du Muséum d'histoire naturelle et remarques sur la classification et les caractères des reptiles (1852) (Crocodylus moreletii).jpg












†OrnithosuchidaeOrnithosuchus BW white background.jpg



 Rauisuchia 















†RauisuchidaePostosuchus kirkpatricki flipped.jpg





†PrestosuchidaePrestosuchus-chiniquensis (2).jpg




 "Group X" 








ArizonasaurusArizonasaurus BW white background.jpg












LotosaurusLotosaurus BW white background.jpg



 "Group Y" or Shuvosaurinae 








Sillosuchus












ShuvosaurusShuvosaurus BW flipped.jpg





EffigiaEffigia BW white background.jpg















Cladogram after Brusatte, Benton, Desojo and Langer (2010) [10]






Archosauriformes 








ErythrosuchusErythrosuchus africanus.jpg












EuparkeriaEuparkeria BW flipped.jpg












†ProterochampsidaeChanaresuchus.jpg



 Archosauria 








to AvemetatarsaliaMeyers grosses Konversations-Lexikon - ein Nachschlagewerk des allgemeinen Wissens (1908) (Antwerpener Breiftaube).jpg




Crurotarsi 








†PhytosauriaSmilosuchus adamanensis flipped.jpg



 Suchia 















†AetosauriaDesmatosuchus spurensis flipped.jpg



 Paracrocodylomorpha 








GracilisuchusGracilisuchus BW white background.jpg



 Bathyotica 








ErpetosuchusErpetosuchus BW white background.jpg





CrocodylomorphaDescription des reptiles nouveaux, ou, Imparfaitement connus de la collection du Muséum d'histoire naturelle et remarques sur la classification et les caractères des reptiles (1852) (Crocodylus moreletii).jpg






















Revueltosaurus





†OrnithosuchidaeOrnithosuchus BW white background.jpg




 Rauisuchia 






 †Rauisuchoidea 















Arganasuchus












Fasolasuchus












Stagonosuchus





TicinosuchusTicinosuchus BW white background.jpg













 Prestosuchidae 








SaurosuchusSaurosuchus BW white background.jpg












Batrachotomus





PrestosuchusPrestosuchus-chiniquensis (2).jpg





 Rauisuchidae 








Tikisuchus












Rauisuchus












PostosuchusPostosuchus kirkpatricki flipped.jpg





Teratosaurus








 †Poposauroidea 








Yarasuchus












Qianosuchus




















ArizonasaurusArizonasaurus BW white background.jpg





Bromsgroveia





LotosaurusLotosaurus BW white background.jpg





PoposaurusPoposaurus gracilis (1) flipped.jpg





Sillosuchus



 Shuvosauridae 








ShuvosaurusShuvosaurus BW flipped.jpg





EffigiaEffigia BW white background.jpg
















References




  1. ^ abcd Nesbitt, S.J. (2011). "The early evolution of archosaurs: relationships and the origin of major clades" (PDF). Bulletin of the American Museum of Natural History. 352: 1–292. doi:10.1206/352.1. 


  2. ^ abc Sereno, P.C.; Arcucci, A.B. (1990). "The monophyly of crurotarsal archosaurs and the origin of bird and crocodile ankle joints". Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 180: 21–52. 


  3. ^ Sereno, Paul (1991). "Basal archosaurs: phylogenetic relationships and functional implications". Journal of Vertebrate Paleontology. 11 (Suppl. 4): 1–51. doi:10.1080/02724634.1991.10011426. 


  4. ^ Gauthier, J. A.; Nesbitt, S. J.; Schachner, E. R.; Bever, G. S.; Joyce, W. G. (2011). "The bipedal stem-crocodilian Poposaurus gracilis: inferring function in fossils and innovation in archosaur locomotion". Bulletin of the Peabody Museum of Natural History. 52: 107–126. doi:10.3374/014.052.0102. 


  5. ^ Gauthier, J.A.; Padian, K. (1985). "Phylogenetic, functional, and aerodynamic analyses of the origin of birds and their flight". In Hecht, M.K.; Ostrom, J.H.; Viohl, G.; Wellnhofer, P. The Beginnings of Birds. Eichstatt: Freunde des Jura-Museums. pp. 185–197. 


  6. ^ Benton, M.J.; Clark, J.M. (1988). "Archosaur phylogeny and the relationships of the Crocodylia". In Benton, M.J. Phylogeny and Classification of the Tetrapods. 1. Oxford: Clarendon Press. pp. 295–338. 


  7. ^ Brochu, C.A. (1997). "Synonymy, redundancy, and the name of the crocodile stem-group". Journal of Vertebrate Paleontology. 17 (2): 448–449. doi:10.1080/02724634.1997.10010992. 


  8. ^ Nesbitt, SJ; Norell, MA. (2006). "Extreme convergence in the body plans of an early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda)". Proceedings of the Royal Society of London B: Biological Sciences. 273 (1590): 1045–1048. doi:10.1098/rspb.2005.3426. PMC 1560254 Freely accessible. PMID 16600879. 


  9. ^ Nesbitt, S. (2007). "The anatomy of Effigia okeeffeae (Archosauria, Suchia), theropod-like convergence, and the distribution of related taxa" (PDF). Bulletin of the American Museum of Natural History. 302: 84. doi:10.1206/0003-0090(2007)302[1:taoeoa]2.0.co;2. 


  10. ^ Brusatte, Stephen L.; Benton, Michael J.; Desojo, Julia B.; Langer, Max C. (2010). "The higher-level phylogeny of Archosauria (Tetrapoda: Diapsida)". Journal of Systematic Palaeontology. 8 (1): 3–47. doi:10.1080/14772010903537732. 



  • Benton, M. J. (2004). Vertebrate Paleontology (3rd ed.). Blackwell Science. 


  • Dunham, W. (2008-09-12). "Lucky break allowed dinosaurs to rule Earth". Reuters. Retrieved 2012-01-14. 


  • Brusatte, S. L.; Benton, M. J.; Ruta, M.; Lloyd, G. T. (2008-09-12). "Superiority, Competition, and Opportunism in the Evolutionary Radiation of Dinosaurs" (PDF). Science. 321 (5895): 1485–1488. doi:10.1126/science.1161833. PMID 18787166. Retrieved 2012-01-14. 


External links




  • Palaeos

  • taxonomic hierarchy according to Benton 2004

  • Mikko's Phylogeny






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