A new archosaur from the Manda beds (Anisian, Middle Triassic) of southern Tanzania and its implications for character state optimizations at Archosauria and Pseudosuchia

LITERATURE CITED

• Abdala, F., and M. Allinson. 2005. The taxonomic status of Parathrinaxodon proops (Therapsida: Cynodontia), with comments on the morphology of the palate in basal cynodonts. Palaeontologia africana 41:45–52.
   Google Scholar
• Abdala, F., P. J. Hancox, and J. Neveling. 2005. Cynodonts from the uppermost Burgersdorp Formation, South Africa, and their bearing on the biostratigraphy and correlation of the Triassic Cynognathus Assemblage Zone. Journal of Vertebrate Paleontology 25:192–199.
   Web of Science ®Google Scholar
• Appleby, R. M., A. J. Charig, C. B. Cox, K. A. Kermack, and L. B. H. Tarlo. 1967. Reptilia; pp. 695–731 in W. B. Harland, C. H. Holland, M. R. House, N. F. Hughes, A. B. Reynolds, M. J. S. Rudwick, G. E. Satterthwaite, L. B. H. Tarlo, and E. C.Willey (eds.), The Fossil Record. The Geological Society of London, London.
   Google Scholar
• Bennett, S. C. 1996. The phylogenetic position of the Pterosauria within Archosauromorpha. Zoological Journal of the Linnean Society 118:261–308.
   Web of Science ®Google Scholar
• Benton, M. J. 1983. Dinosaur success in the Triassic; a noncompetitive ecological model. The Quarterly Review of Biology 58:29–55.
   Web of Science ®Google Scholar
• Benton, M. J. 1999. Scleromochlus taylori and the origin of dinosaurs and pterosaurs. Philosophical Transactions of the Royal Society of London, Series B 354:1423–1446.
   Web of Science ®Google Scholar
• Benton, M. J. 2010. Investigating evolutionary radiations; pp. 10–26 in M. Long and Z. Zhou (eds.), Darwin's Heritage Today—Proceedings of the Darwin 200 Beijing International Conference. Higher Education Press, Beijing.
   Google Scholar
• Benton, M. J., and J. M. Clark. 1988. Archosaur phylogeny and the relationships of the Crocodylia; pp. 295–338 in M. J. Benton (ed.), The Phylogeny and Classification of the Tetrapods. Volume 1: Amphibians and Reptiles. Clarendon Press, Oxford, U.K.
   Google Scholar
• Borsuk-Bialynicka, M., and S. E. Evans. 2009. Cranial and mandibular osteology of the Early Triassic archosauriform Osmolskina czatkowicensis from Poland. Palaeontologia Polonica 65:235–281.
   Google Scholar
• Brochu, C. A. 1992. Ontogeny of the postcranium in crocodylomorph archosaurs. M.S. thesis, University of Texas at Austin, Austin, Texas, 340 pp.
   Google Scholar
• Brusatte, S. L., M. J. Benton, J. B. Desojo, and M. C. Langer. 2010. The higher-level phylogeny of Archosauria (Tetrapoda: Diapsida). Journal of Systematic Palaeontology 8:3–47.
   Web of Science ®Google Scholar
• Brusatte, S. L., M. J. Benton, M. Ruta, and G. T. Lloyd. 2008. Superiority, competition, and opportunism in the evolutionary radiation of dinosaurs. Science 321:1485–1488.
   PubMed Web of Science ®Google Scholar
• Brusatte, S. L., M. J. Benton, G. T. Lloyd, M. Ruta, and S. C. Wang. 2011. Macroevolutionary patterns in the evolutionary radiation of archosaurs (Tetrapoda: Diapsida). Earth and Environmental Science Transactions of the Royal Society of Edinburgh 101:367–382.
   Web of Science ®Google Scholar
• Butler, R. J., P. M. Barrett, R. L. Abel, and D. J. Gower. 2009. A possible ctenosauriscid archosaur from the Middle Triassic Manda Beds of Tanzania. Journal of Vertebrate Paleontology 29:1022–1031.
   Web of Science ®Google Scholar
• Butler, R. J., S. L. Brusatte, M. Reich, S. J. Nesbitt, R. R. Schoch, and J. J. Hornung. 2011. The sail-backed reptile Ctenosauriscus from the latest Early Triassic of Germany and the timing and biogeography of the early archosaur radiation. PLoS ONE 6:e25693. doi:10.1371/journal.pone.0025693.
   PubMed Web of Science ®Google Scholar
• Carrano, M. T., and J. R. Hutchinson. 2002. Pelvic and hindlimb musculature of Tyrannosaurus rex (Dinosauria: Theropoda). Journal of Morphology 253:207–228.
   PubMed Web of Science ®Google Scholar
• Charig, A. J. 1956. New Triassic archosaurs from Tanganyika including Mandasuchus and Teleocrater. Ph.D. dissertation, University of Cambridge, Cambridge, U.K., 503 pp.
   Google Scholar
• Charig, A. J. 1972. The evolution of the archosaur pelvis and hind-limb: an explanation in functional terms; pp. 121–155 in K. A. Joysey and T. S. Kemp (eds.), Studies in Vertebrate Evolution. Oliver and Boyd, Edinburgh.
   Google Scholar
• Charig, A. J. 1984. Competition between therapsids and archosaurs during the Triassic Period: a review and synthesis of current theories. Zoological Society of London Symposium 52:597–628.
   Google Scholar
• Chatterjee, S. 1985. Postosuchus, a new thecodontian reptile from the Triassic of Texas and the origin of tyrannosaurs. Philosophical Transactions of the Royal Society of London, Series B 309:395–460.
   Web of Science ®Google Scholar
• Cope, E. D. 1869. Synopsis of the extinct Batrachia, Reptilia and Aves of North America. Transactions of the American Philosophical Society, New Series 14:1–252.
   Google Scholar
• Cox, C. B. 1991. The Pangaea dicynodont Rechnisaurus and the comparative biostratigraphy of Triassic dicynodont faunas. Palaeontology 34:767–784.
   Web of Science ®Google Scholar
• Cruickshank, A. R. I. 1972. The proterosuchian thecodonts; pp. 89–119 in K. A. Joysey and T. S. Kemp (eds.), Studies in Vertebrate Evolution. Oliver and Boyd, Edinburgh.
   Google Scholar
• Ewer, R. F. 1965. The anatomy of the thecodont reptile Euparkeria capensis Broom. Philosophical Transactions of the Royal Society of London, Series B 248:379–435.
   Web of Science ®Google Scholar
• Furin, S., N. Preto, M. Rigo, G. Roghi, P. Gianolla, J. L. Crowley, and S. A. Bowring. 2006. High-precision U-Pb zircon age from the Triassic of Italy: implications for the Triassic time scale and the Carnian origin of calcareous nannoplankton and dinosaurs. Geology 34:1009–1012.
   Web of Science ®Google Scholar
• Gauthier, J. A. 1984. A cladistic analysis of the higher systematic categories of Diapsida. Ph.D. dissertation, University of California Berkeley, Berkeley, California, 564 pp.
   Google Scholar
• Gauthier, J., and K. Padian. 1985. Phylogenetic, functional, and aerodynamic analyses of the origin of birds and their flight; pp. 185–197 in M. K. Hecht, J. H. Ostrom, G. Viohl, and P. Wellnhofer (eds.), The Beginning of Birds. Freunde des Jura Museums, Eichstätt.
   Google Scholar
• Gauthier, J. A., A. G. Kluge, and T. Rowe. 1988a. The early evolution of the Amniota; pp. 103–155 in M. J. Benton (ed.), The Phylogeny and Classification of the Tetrapods, Volume 1: Amphibians, Reptiles, Birds. Clarendon Press, Oxford, U.K.
   Google Scholar
• Gauthier, J. A., A. G. Kluge, and T. Rowe. 1988b. Amniote phylogeny and the importance of fossils. Cladistics 4:105–209.
   PubMed Web of Science ®Google Scholar
• Goloboff, P. A., J. S. Farris, and K. Nixon. 2003. TNT: Tree Analysis Using New Technologies. Program and documentation available from the authors and at http://www.zmuc.dk/public/phylogeny.
   Google Scholar
• Goloboff, P. A., J. S. Farris, and K. Nixon. 2008. TNT: a free program for phylogenetic analysis. Cladistics 24:774–786.
   Web of Science ®Google Scholar
• Gower, D. J. 1999. The cranial and mandibular osteology of a new rauisuchian archosaur from the Middle Triassic of southern Germany. Stuttgarter Beiträge zur Naturkunde, Serie B (Geologie und Paläontologie) 280:1–49.
   Google Scholar
• Gower, D. J. 2003. Osteology of the early archosaurian reptile Erythrosuchus africanus Broom. Annals of the South African Museum 110:1–84.
   Google Scholar
• Gower, D. J., and R. Schoch. 2009. Postcranial anatomy of the rauisuchian archosaur Batrachotomus kupferzellensis. Journal of Vertebrate Paleontology 29:103–122.
   Web of Science ®Google Scholar
• Gower, D. J., and A. G. Sennikov. 1997. Sarmatosuchus and the early history of the Archosauria. Journal of Vertebrate Paleontology 17:60–73.
   Web of Science ®Google Scholar
• Gower, D. J., and A. G. Sennikov. 2000. Early archosaurs from Russia; pp. 140–159 in M. J. Benton, M. A. Shishkin, D. M. Unwin, and E. N. Kurochkin (eds.), The Age of Dinosaurs in Russia and Mongolia. Cambridge University Press, New York.
   Google Scholar
• Hancox, P. J. 2000. The continental Triassic of South Africa. Zentralblatt für Geologie und Paläontologie Teil I, Heft 11–12 1998:1285–1324.
   Google Scholar
• Hancox, P. J., K. D. Angielczyk, and B. S. Rubidge. 2013. Angonisaurus and Shansiodon, dicynodonts (Therapsida, Anomodontia) from subzone C of the Cynognathus Assemblage Zone (Middle Triassic) of South Africa. Journal of Vertebrate Paleontology 33:655–676.
   Web of Science ®Google Scholar
• Huene, F. von. 1938. Ein grosser Stagonolepide aus der jüngeren Trias Ostafrikas. Neues Jahrbuch für Geologie und Paläeontologie, Beilage-Bände Abteilung B 80:264–278.
   Google Scholar
• Huene, F. von. 1939. Ein kleiner Pseudosuchier und ein Saurischier aus den Ostafrikanischen Mandaschichten. Neues Jahrbuch für Geologie und Paläeontologie, Beilage-Bände Abteilung B 81:61–69.
   Google Scholar
• Huene, F. von. 1942. Die fossilen Reptilien des Sudamerikanischen Gondwanalandes. Ergebnisse der Sauriergrabungen in Sudbrasilien, 1928/1929. C.H. Beck'she, Munich, 332 pp.
   Google Scholar
• Hutchinson, J. R. 2001a. The evolution of pelvic osteology and soft tissues on the line to extant birds. Zoological Journal of the Linnean Society 131:123–168.
   Web of Science ®Google Scholar
• Hutchinson, J. R. 2001b. The evolution of femoral osteology and soft tissue on the line to extant birds (Neornithes). Zoological Journal of the Linnean Society 131:169–197.
   Web of Science ®Google Scholar
• Irmis, R. B., R. Mundil, J. W. Martz, and W. G. Parker. 2011. High-resolution U-Pb ages from the Upper Triassic Chinle Formation (New Mexico, USA) support a diachronous rise of dinosaurs. Earth and Planetary Science Letters 309:258–267.
   Web of Science ®Google Scholar
• Irmis, R. B., S. J. Nesbitt, K. Padian, N. D. Smith, A. H. Turner, D. Woody, and A. Downs. 2007. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science 317:358–361.
   PubMed Web of Science ®Google Scholar
• Juul, L. 1994. The phylogeny of basal archosaurs. Palaeontologia africana 31:1–38.
   Google Scholar
• Krebs, B. 1965. Die Triasfauna der Tessiner Kalkalpen. XIX. Ticinosuchus ferox, nov. gen. nov. sp. Ein neuer Pseudosuchier aus der Trias des Monte San Georgio. Schweizersiche Palaontologische, Abhandlungen 81:1–140.
   Google Scholar
• Li, C., X.-C. Wu, L.-J. Zhao, T. Sato, and L.-T. Wang. 2012. A new archosaur (Diapsida, Archosauriformes) from the marine Triassic of China. Journal of Vertebrate Paleontology 32:1064–1081.
   Web of Science ®Google Scholar
• Long, R. A., and P. A. Murry. 1995. Late Triassic (Carnian and Norian) tetrapods from the southwestern United States New Mexico Museum of Natural History and Science Bulletin 4:1–254.
   Google Scholar
• Lucas, S. G. 1998. Global Triassic tetrapod biostratigraphy and biochronology. Palaeogeography, Palaeoclimatology, Palaeoecology 143:347–384.
   Web of Science ®Google Scholar
• Maddison, W., and D. Maddison. 2003. MacClade, version 4.06. Sinauer Association, Sunderland, Massachusetts.
   Google Scholar
• Martinez, R. N., P. C. Sereno, O. A. Alcober, C. E. Colombi, P. R. Renne, I. P. Montanez, and B. S. Currie. 2011. A basal dinosaur from the dawn of the dinosaur era in southwestern Pangaea. Science 331:206–210.
   PubMed Web of Science ®Google Scholar
• Muttoni, G., D. V. Kent, P. E. Olsen, P. DiStefano, W. Lowrie, S. Bernasconi, and F. M. Hernandez. 2004. Tethyan magnetostratigraphy from Pizzi Mondello (Sicily) and correlation to the Late Triassic Newark astrochronological polarity time scale. Geological Society of America Bulletin 116:1043–1058.
   Web of Science ®Google Scholar
• Nesbitt, S. J. 2003. Arizonasaurus and its implications for archosaur divergences. Proceedings of the Royal Society of London, Series B 270(Supplement 2):S234–S237.
   Web of Science ®Google Scholar
• Nesbitt, S. J. 2005. The osteology of the Middle Triassic pseudosuchian archosaur Arizonasaurus babbitti. Historical Biology 17:19–47.
   Google Scholar
• Nesbitt, S. J. 2007. The anatomy of Effigia okeeffeae (Archosauria, Suchia), theropod convergence, and the distribution of related taxa. Bulletin of the American Museum of Natural History 302:1–84.
   Google Scholar
• Nesbitt, S. J. 2011. The early evolution of Archosauria: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 352:1–292.
   Web of Science ®Google Scholar
• Nesbitt, S. J., and R. J. Butler. 2012. Redescription of the archosaur Parringtonia gracilis from the Middle Triassic Manda beds of Tanzania, and the antiquity of Erpetosuchidae. Geological Magazine 150:225–238.
   Web of Science ®Google Scholar
• Nesbitt, S. J., R. J. Butler, and D. J. Gower. 2013. A new archosauriform (Reptilia: Diapsida) from the Manda beds (Middle Triassic) of southwestern Tanzania. PLoS ONE . 8(9): e72753 doi:10.1371/journal.pone.0072753
   Google Scholar
• Nesbitt, S. J., J. Liu, and C. Li. 2011. A sail-backed suchian from the Heshanggou Formation (Early Triassic: Olenekian) of China. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 101:271–284.
   Web of Science ®Google Scholar
• Nesbitt, S. J., M. R. Stocker, B. Small, and A. Downs. 2009b. The osteology and relationships of Vancleavea campi (Reptilia: Archosauriformes). Zoological Journal of the Linnean Society 157:814–864.
   Web of Science ®Google Scholar
• Nesbitt, S. J., P. M. Barrett, S. Werning, C. A. Sidor, and A. J. Charig. 2012. The oldest dinosaur? A Middle Triassic dinosauriform from Tanzania. Biology Letters 9:20120949. doi:10.1098/rsbl.2012.0949.
   PubMed Web of Science ®Google Scholar
• Nesbitt, S. J., C. A. Sidor, R. B. Irmis, K. D. Angielczyk, R. M. H. Smith, and L. A. Tsuji. 2010. Ecologically distinct dinosaurian sister-group shows early diversification of Ornithodira. Nature 464:95–98.
   PubMed Web of Science ®Google Scholar
• Nesbitt, S. J., N. D. Smith, R. B. Irmis, A. H. Turner, A. Downs, and M. A. Norell. 2009a. A complete skeleton of a Late Triassic saurischian and the early evolution of dinosaurs. Science 326:1530–1533.
   PubMed Web of Science ®Google Scholar
• Novas, F. E. 1996. Dinosaur monophyly. Journal of Vertebrate Paleontology 16:723–741.
   Web of Science ®Google Scholar
• Parker, W. G. 2008. Description of new material of the aetosaur Desmatosuchus spurensis (Archosauria: Suchia) from the Chinle Formation of Arizona and a revision of the genus Desmatosuchus. PaleoBios 28:1–40.
   Google Scholar
• Parrish, J. M. 1993. Phylogeny of the Crocodylotarsi, with reference to archosaurian and crurotarsan monophyly. Journal of Vertebrate Paleontology 13:287–308.
   Google Scholar
• Peyer, K., J. G. Carter, H.-D. Sues, S. E. Novak, and P. E. Olsen. 2008. A new suchian archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 28:363–381.
   Web of Science ®Google Scholar
• Rauhut, O. W. M. 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology 69:1–214.
   Google Scholar
• Romer, A. S. 1927. The pelvic musculature of ornithischian dinosaurs. Acta Zoologica 8:225–275.
   Google Scholar
• Romer, A. S. 1956. The Osteology of the Reptiles. University of Chicago Press, Chicago, Illinois, 772 pp.
   Google Scholar
• Romer, A. S. 1971. The Chañares (Argentina) Triassic reptile fauna. XI. Two new long-snouted thecodonts, Chanaresuchus and Gualosuchus. Breviora 379:1–22.
   Google Scholar
• Rubidge, B. S. 2005. Re-uniting lost continents—fossil reptiles from the ancient Karoo and their wanderlust. South African Journal of Geology 108:135–172.
   Web of Science ®Google Scholar
• Schachner, E. R., P. L. Manning, and P. Dodson. 2011. Pelvic and hindlimb myology of the basal archosaur Poposaurus gracilis (Archosauria: Poposauroidea). Journal of Morphology 272:1464–1491.
   PubMed Web of Science ®Google Scholar
• Sen, K. 2005. A new rauisuchian archosaur from the Middle Triassic of India. Palaeontology 48:185–196.
   Web of Science ®Google Scholar
• Sereno, P. C. 1991. Basal archosaurs: phylogenetic relationships and functional implications. Society of Vertebrate Paleontology Memoir 2:1–53.
   Google Scholar
• Sereno, P. C., and A. B. Arcucci. 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.
   Google Scholar
• Sidor, C. A., D. A. Vilhena, K. D. Angielczyk, A. K. Huttenlocker, S. J. Nesbitt, B. R. Peecook, J. S. Steyer, R. M. H. Smith, and L. A. Tsuji. 2013. Provincialization of terrestrial faunas following the end-Permian mass extinction. Proceedings of the National Academy of Sciences of the United States of America 110:8129–8133.
   PubMed Web of Science ®Google Scholar
• Sookias, R. B., and Butler, R. J. 2013. Euparkeriidae; pp. 35–48 in S. J. Nesbitt, J. B. Desojo, and R. B. Irmis (eds.), The Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and Their Kin. Special Publications of the Geological Society of London 379. doi:10.1144/SP379.6.
   Google Scholar
• Stocker, M. R. 2010. A new taxon of phytosaur (Archosauria: Pseudosuchia) from the Late Triassic (Norian) Sonsela Member (Chinle Formation) in Arizona, and a critical reevaluation of Leptosuchus Case, 1922. Palaeontology 53:997–1022.
   Web of Science ®Google Scholar
• Stockley, G. M. 1932. The geology of the Ruhuhu coalfields, Tanganyika Territory. Quarterly Journal of the Geological Society of London 88:610–622.
   Google Scholar
• Swofford, D. L. 2002. PAUP* (Phylogenetic Analysis Using Parsimony and Other Methods), version 4.10b. Sinauer Association, Sunderland, Massachusetts.
   Google Scholar
• Walker, A. D. 1964. Triassic reptiles from the Elgin area: Ornithosuchus and the origin of carnosaurs. Philosophical Transactions of the Royal Society of London, Series B 248:53–134.
   Web of Science ®Google Scholar
• Welman, J. 1998. The taxonomy of the South African proterosuchids (Reptilia, Archosauromorpha). Journal of Vertebrate Paleontology 18:340–347.
   Web of Science ®Google Scholar
• Wilson, J. A. 1999. A nomenclature for vertebral laminae in sauropods and other saurischian dinosaurs. Journal of Vertebrate Paleontology 19:639–653.
   Web of Science ®Google Scholar
• Wu, X.-C., and A. P. Russell. 2001. Redescription of Turfanosuchus dabanensis (Archosauriformes) and new information on its phylogenetic relationships. Journal of Vertebrate Paleontology 21:40–50.
   Web of Science ®Google Scholar
• Submitted May 11, 2013; revisions received September 14, 2013; accepted October 20, 2013.
   Google Scholar
• Handling editor: Hailu You.
   Google Scholar