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Revision of Erpetosuchus (Archosauria: Pseudosuchia) and new erpetosuchid material from the Late Triassic ‘Elgin Reptile’ fauna based on μCT scanning techniques

Published online by Cambridge University Press:  06 November 2020

Davide FOFFA Open the ORCID record for Davide FOFFA [Opens in a new window] ,
Richard J. BUTLER ,
Sterling J. NESBITT ,
Stig WALSH ,
Paul M. BARRETT ,
Stephen L. BRUSATTE  and
Nicholas C. FRASER
Davide FOFFA*
Affiliation:
Department of Natural Sciences, National Museums Scotland, Chambers Street, EdinburghEH1 1JF, UK
Richard J. BUTLER
Affiliation:
School of Geography, Earth and Environmental Sciences, University of Birmingham, Edgbaston, BirminghamB15 2TT, UK
Sterling J. NESBITT
Affiliation:
Department of Geosciences, Virginia Tech, 4044 Derring Hall (MC0420) 926 West Campus Drive, Blacksburg 24061, Virginia, USA
Stig WALSH
Affiliation:
Department of Natural Sciences, National Museums Scotland, Chambers Street, EdinburghEH1 1JF, UK School of GeoSciences, University of Edinburgh, Grant Institute, Hutton Road, EdinburghEH9 3FE, UK
Paul M. BARRETT
Affiliation:
Department of Earth Sciences, Natural History Museum, Cromwell Road, LondonSW7 5BD, UK
Stephen L. BRUSATTE
Affiliation:
Department of Natural Sciences, National Museums Scotland, Chambers Street, EdinburghEH1 1JF, UK School of GeoSciences, University of Edinburgh, Grant Institute, Hutton Road, EdinburghEH9 3FE, UK
Nicholas C. FRASER
Affiliation:
Department of Natural Sciences, National Museums Scotland, Chambers Street, EdinburghEH1 1JF, UK School of GeoSciences, University of Edinburgh, Grant Institute, Hutton Road, EdinburghEH9 3FE, UK
*
*Corresponding author. Email: D.Foffa@nms.ac.uk
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Abstract

The Late Triassic fauna of the Lossiemouth Sandstone Formation (LSF) from the Elgin area, Scotland, has been pivotal in expanding our understanding of Triassic terrestrial tetrapods. Frustratingly, due to their odd preservation, interpretations of the Elgin Triassic specimens have relied on destructive moulding techniques, which only provide incomplete, and potentially distorted, information. Here, we show that micro-computed tomography (μCT) could revitalise the study of this important assemblage. We describe a long-neglected specimen that was originally identified as a pseudosuchian archosaur, Ornithosuchus woodwardi. μCT scans revealed dozens of bones belonging to at least two taxa: a small-bodied pseudosuchian and a specimen of the procolophonid Leptopleuron lacertinum. The pseudosuchian skeleton possesses a combination of characters that are unique to the clade Erpetosuchidae. As a basis for investigating the phylogenetic relationships of this new specimen, we reviewed the anatomy, taxonomy and systematics of other erpetosuchid specimens from the LSF (all previously referred to Erpetosuchus). Unfortunately, due to the differing representation of the skeleton in the available Erpetosuchus specimens, we cannot determine whether the erpetosuchid specimen we describe here belongs to Erpetosuchus granti (to which we show it is closely related) or if it represents a distinct new taxon. Nevertheless, our results shed light on rarely preserved details of erpetosuchid anatomy. Finally, the unanticipated new information extracted from both previously studied and neglected specimens suggests that fossil remains may be much more widely distributed in the Elgin quarries than previously recognised, and that the richness of the LSF might have been underestimated.

Keywords

anatomyErpetosuchidaeErpetosuchus grantiLeptopleuronsystematics
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Earth and Environmental Science Transactions of The Royal Society of Edinburgh , Volume 111 , Issue 4 , December 2020 , pp. 209 - 233
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Copyright © The Author(s) 2020. Published by Cambridge University Press on behalf of The Royal Society of Edinburgh

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References

8. References

Bennett, S. C. 2020. Reassessment of the Triassic archosauriform Scleromochlus taylori: neither runner nor biped, but hopper. PeerJ 8, e8418.CrossRefGoogle ScholarPubMed
Benton, M. J. & Walker, A. D. 1985. Palaeoecology, taphonomy, and dating of Permo-Triassic reptiles from Elgin, north-east Scotland. Palaeontology 28, 207–34.Google Scholar
Benton, M. J. & Walker, A. D. 2002. Erpetosuchus, a crocodile-like basal archosaur from the Late Triassic of Elgin, Scotland. Zoological Journal of the Linnean Society 136, 25–47.CrossRefGoogle Scholar
Benton, M. J. & Walker, A. D. 2011. Saltopus, a dinosauriform from the Upper Triassic of Scotland. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 101, 285–99.CrossRefGoogle Scholar
Butler, R. J., Sullivan, C., Ezcurra, M. D., Liu, J., Lecuona, A. & Sookias, R. B. 2014. New clade of enigmatic early archosaurs yields insights into early pseudosuchian phylogeny and the biogeography of the archosaur radiation. BMC Evolutionary Biology 14, 128.CrossRefGoogle ScholarPubMed
Butler, R. J., Nesbitt, S. J., Charig, A. J., Gower, D. J. & Barrett, P. M. 2018. Mandasuchus tanyauchen, gen. et sp. nov., a pseudosuchian archosaur from the Manda Beds (?Middle Triassic) of Tanzania. In Sidor, C. A. & Nesbitt, S. J. (eds) Vertebrate and climatic evolution in the Triassic rift basins of Tanzania and Zambia, 17, 96–121. Society of Vertebrate Paleontology Memoir.Google Scholar
Butler, R. J., Sennikov, A. G., Dunne, E. M., Ezcurra, M. D., Hedrick, B. P., Maidment, S. C. R., Meade, L. E., Raven, T. J. & Gower, D. J. 2019a. Cranial anatomy and taxonomy of the erythrosuchid archosauriform ‘Vjushkovia triplicostata’ Huene, 1960, from the Early Triassic of European Russia. Royal Society Open Science 6, 191289.CrossRefGoogle Scholar
Butler, R. J., Sennikov, A. G., Ezcurra, M. D. & Gower, D. J. 2019b. The last erythrosuchid? A revision of Chalishevia cothurnata Ochev, 1980 from the late Middle Triassic of European Russia. Acta Palaeontologica Polonica 64, 757–74.CrossRefGoogle Scholar
Cabreira, S. F., Kellner, A., Dias-da-Silva, S., Roberto da Silva, L., Bronzati, M., Marsola, J., Müller, R. T., Bittencourt, J. S., Batista, B. J., Raugust, T., Carrilho, R., Brodt, A. & & Langer, M. C. 2016. A unique Late Triassic dinosauromorph assemblage reveals dinosaur ancestral anatomy and diet. Current Biology 26, 3090–95.CrossRefGoogle ScholarPubMed
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, Biological Sciences 309, 395–460.Google Scholar
Clark, J. M., Sues, H.-D. & Berman, D. S. 2000. A new specimen of Hesperosuchus agilis from the Upper Triassic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology 20, 683–704.CrossRefGoogle Scholar
Cope, E. D. 1869. Synopsis of the extinct Batrachia, Reptilia, and Aves of North America. Transactions of the American Philosophical Society 14, 1–252.CrossRefGoogle Scholar
Cunningham, J. A., Rahman, I. A., Lautenschlager, S., Rayfield, E. J. & Donoghue, P. C. J. 2014. A virtual world of paleontology. Trends in Ecology and Evolution 29, 347–57.CrossRefGoogle ScholarPubMed
Davies, T. G., Rahman, I. A., Lautenschlager, S., Cunningham, J. A., Asher, R. J., Barrett, P. M., Bates, K. T., Bengtson, S., Benson, R. B. J., Boyer, D. M., Braga, J., Bright, J. A., Claessens, L. P. A. M., Cox, P. G., Dong, X.-P., Evans, A. R., Falkingham, P. L., Friedman, M., Garwood, R. J., Goswami, A., Hutchinson, J. R., Jeffery, N. S., Johanson, Z., Lebrun, R., Martínez-Pérez, C., Marugán-Lobón, J., O'Higgins, P. M., Metscher, B., Orliac, M., Rowe, T. B., Rücklin, M., Sánchez-Villagra, M. R., Shubin, N. H., Smith, S. Y., Starck, J. M., Stringer, C., Summers, A. P., Sutton, M. D., Walsh, S. A., Weisbecker, V., Witmer, L. M., Wroe, S., Yin, Z., Rayfield, E. J. & Donoghue, P. C. J. 2017. Open data and digital morphology. Proceedings of the Royal Society B 284, 20170194.CrossRefGoogle ScholarPubMed
Desojo, J. B., Heckert, A. B., Martz, J. W., Parker, W. G., Schoch, R. R., Small, B. J. & Sulej, T. 2013. Aetosauria: a clade of armoured pseudosuchians from the Upper Triassic continental beds. In Nesbitt, S. J., Desojo, J. B. & Irmis, R. B. (eds) Anatomy, phylogeny and palaeobiology of early archosaurs and their kin, 379, 203–39. Geological Society of London, Special Publication.Google Scholar
Dilkes, D. W. & Arcucci, A. B. 2012. Proterochampsa barrionuevoi (Archosauriformes: Proterochampsia) from the Late Triassic (Carnian) of Argentina and a phylogenetic analysis of Proterochampsia. Palaeontology 55, 853–85.CrossRefGoogle Scholar
Ezcurra, M. D. 2016. The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriforms. PeerJ 4, e1778.CrossRefGoogle ScholarPubMed
Ezcurra, M. D., Fiorelli, L. E., Martinelli, A. G., Rocher, S. M., Von Baczko, B., Ezpeleta, M., Taborda, J. R. A., Hechenleitner, E. M., Trotteyn, M. J. & Desojo, J. B. 2017. Deep faunistic changes preceded the raise of dinosaurs in southwestern Pangaea. Nature Ecology & Evolution 1, 1477–83.CrossRefGoogle Scholar
Frostick, L., Reid, I., Jarvis, J. & Eardley, H. 1988. Triassic sediments of the Inner Moray Firth, Scotland: early rift deposits. Journal of the Geological Society 145, 235–48.CrossRefGoogle Scholar
Gauthier, J. & 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. (eds) The beginning of birds, 185–97. Eichstatt: Freunde des Jura-Museums.Google Scholar
Goloboff, P. A., Farris, J. S. & Nixon, K. C. 2008. TNT: a free program for phylogenetic analysis. Cladistics 24, 774–86.CrossRefGoogle Scholar
Gordon, G. 1859. On the geology of the lower and western part of the province of Moray: its history, present state of enquiry and points for future examination. Edinburgh New Philosophical Journal, New Series 9, 14–58.Google Scholar
Gower, D. J. 1999. Cranial osteology of a new rauisuchian archosaur from the Middle Triassic of southern Germany. Stuttgarter Beiträge zur Naturkunde B 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
Irmis, R. B., Nesbitt, S. J. & Sues, H.-D. 2013. Early Crocodylomorpha. In Nesbitt, S. J., Desojo, J. B. & Irmis, R. B. (eds) Anatomy, phylogeny and palaeobiology of early archosaurs and their kin, 379, 275–302. Geological Society, London, Special Publications.Google Scholar
Jirak, D. & Janacek, J. 2017 Volume of the crocodilian brain and endocast during ontogeny. PLoS ONE 12, e0178491.CrossRefGoogle ScholarPubMed
Jones, A. S. & Butler, R. J. 2018. A new phylogenetic analysis of Phytosauria (Archosauria: Pseudosuchia) with the application of continuous and geometric morphometric character coding. PeerJ 6, e5901.CrossRefGoogle ScholarPubMed
Keeble, E. & Benton, M. J. 2020. Three-dimensional tomographic study of dermal armour from the tail of the Triassic aetosaur Stagonolepis robertsoni. Scottish Journal of Geology 56, 55–62. doi: 10.1144/sjg2019-026.CrossRefGoogle Scholar
Krebs, B. 1974. Die Archosaurier. Naturwissenschaften 61, 17–24.CrossRefGoogle Scholar
Lacerda, M. B., de Franca, M. A. G. & Schultz, C. L. 2018. A new erpetosuchid (Pseudosuchia, Archosauria) from the Middle–Late Triassic of southern Brazil. Zoological Journal of the Linnean Society 184, 804–24.CrossRefGoogle Scholar
Langer, M. C. & Benton, M. J. 2006. Early dinosaurs: a phylogenetic study. Journal of Systematic Palaeontology 4, 309–58.CrossRefGoogle Scholar
Lessner, E. J., Stocker, M. R., Smith, N. D., Turner, A. H., Irmis, R. B. & Nesbitt, S. J. 2016. A new rauisuchid (Archosauria, Pseudosuchia) from the Upper Triassic (Norian) of New Mexico increases the diversity and temporal range of the clade. PeerJ 4, e2336.CrossRefGoogle ScholarPubMed
Lucas, S. G., Wild, R. & Hunt, A. T. 1998. Dyoplax O. Fraas, a Triassic sphenosuchian from Germany. Stuttgarter Beiträge zur Naturkunde B 263, 1–13.Google Scholar
Maisch, M. W., Matzke, A. T. & Rathgeber, T. 2013. Re-evaluation of the enigmatic archosaur Dyoplax arenaceus O. Fraas, 1867 from the Schilfsandstein (Stuttgart Formation, lower Carnian, Upper Triassic) of Stuttgart, Germany. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 267, 353–62.CrossRefGoogle Scholar
Martin, J. C. 1860. A ramble among the fossiliferous beds of Moray. 9 pp. Unpublished MS held by Elgin Museum.Google Scholar
Mastrantonio, B. M., Von Baczko, M. B., Desojo, J. B. & Schultz, C. L. 2019. The skull anatomy and cranial endocast of the pseudosuchid archosaur Prestosuchus chiniquensis from the Triassic of Brazil. Acta Palaeontologica Polonica 64, 171–98.CrossRefGoogle Scholar
Müller, R. T., Langer, M. C. & Dias-da-Silva, S. 2018. An exceptionally preserved association of complete dinosaur skeletons reveals the oldest long-necked sauropodomorphs. Biology Letters 14, 20180633.CrossRefGoogle ScholarPubMed
Müller, R. T., Von Baczko, M. B., Desojo, J. B. & Nesbitt, S. J. 2020. The first ornithosuchid from Brazil and its macroevolutionary and phylogenetic implications for Late Triassic faunas in Gondwana. Acta Palaeontologica Polonica 65, 1–10.CrossRefGoogle Scholar
Murchison, R. I. 1859. On the sandstones of Morayshire (Elgin, &c.) containing reptilian remains; and on their relations to the Old Red Sandstone of that country. Quarterly Journal of the Geological Society 15, 419–39.CrossRefGoogle Scholar
Nesbitt, S. J. 2007. The anatomy of Effigia okeeffeae (Archosauria, Suchia), theropod-like convergence, and the distribution of related taxa. Bulletin of the American Museum of Natural History 302, 1–84.CrossRefGoogle Scholar
Nesbitt, S. J. 2011. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 352, 1–292.CrossRefGoogle Scholar
Nesbitt, S. J., Brusatte, S. J., Desojo, J. B., Liparini, A., De Franca, M. A. G., Weinbaum, J. C. & Gower, D. J. 2013. Rauisuchia. In Nesbitt, S. J., Desojo, J. B. & Irmis, R. B. (eds) Anatomy, phylogeny and palaeobiology of early archosaurs and their kin, 379, 241–74. Geological Society, London, Special Publications.Google Scholar
Nesbitt, S. J., Stocker, M. R., Parker, W. G., Wood, T. A., Sidor, C. A. & Angielczyk, K. D. 2018. The braincase and endocast of Parringtonia gracilis, a Middle Triassic suchian (Archosauria: Pseudosuchia). In Sidor, C. A. & Nesbitt, S. J. (eds) Vertebrate and climatic evolution in the Triassic rift basins of Tanzania and Zambia, 17, 122–41. Society of Vertebrate Paleontology Memoir.Google Scholar
Nesbitt, S. J. & Butler, R. J. 2013. Redescription of the archosaur Parringtonia gracilis From the Middle Triassic Manda Beds of Tanzania, and the antiquity of Erpetosuchidae. Geological Magazine 150, 225–38.CrossRefGoogle Scholar
Newton, E. T. 1894. Reptiles from the Elgin Sandstone. Description of two new genera Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences 185, 573–607.Google Scholar
Olsen, P. E., Sues, H.-D. & Norell, M. A. 2001. First record of Erpetosuchus (Reptilia: Archosauria) from the Late Triassic of North America. Journal of Vertebrate Paleontology 20, 633–36.CrossRefGoogle Scholar
Parker, W. G. 2018. Anatomical notes and discussion of the first described aetosaur Stagonolepis robertsoni (Archosauria: Suchia) from the Upper Triassic of Europe, and the use of plesiomorphies in aetosaur biochronology. PeerJ 6, e5455.CrossRefGoogle ScholarPubMed
Parker, W. G., Irmis, R. B., Nesbitt, S. J., Martz, J. W. & Browne, L. S. 2005. The Late Triassic pseudosuchian Revueltosaurus callenderi and its implications for the diversity of early ornithischian dinosaurs. Proceedings of the Royal Society B: Biological Sciences 272, 963–69.CrossRefGoogle ScholarPubMed
Parrish, J. M. 1993. Phylogeny of the Crocodylotarsi, with reference to archosaurian and crurotarsan monophyly. Journal of Vertebrate Paleontology 12, 287–308.CrossRefGoogle Scholar
Peacock, J. D., Berridge, N. G., Harris, A. L. & May, F. 1968. The Geology of the Elgin District (Sheet 95). Memoirs of the Geological Survey of Scotland, HMSO, Edinburgh, 155 pp.Google Scholar
Roberto-Da-Silva, L., França, M. A. G., Cabreira, S. F., Müller, R. T. & Dias-Da-Silva, S. 2016. On the presence of the subnarial foramen in Prestosuchus chiniquensis (Pseudosuchia: Loricata) with remarks on its phylogenetic distribution. Anais da Academia Brasileira de Ciências 88, 1309–23.CrossRefGoogle ScholarPubMed
Säilä, L. 2010. Osteology of Leptopleuron lacertinum Owen, a procolophonoid parareptile from the Upper Triassic of Scotland, with remarks on ontogeny, ecology and affinities. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 101, 1–25.CrossRefGoogle Scholar
Schachner, E. R., Irmis, R. B., Huttenlocker, A. K., Sanders, K., Cieri, R. L. & Nesbitt, S. J. 2020. Osteology of the Late Triassic bipedal archosaur Poposaurus gracilis (Archosauria: Pseudosuchia) from Western North America. Anatomical Record 303, 874–917.CrossRefGoogle ScholarPubMed
Sereno, P. C. 1991. Basal archosaurs: phylogenetic relationships and functional implications. Society of Vertebrate Paleontology Memoir 2, 1–53.CrossRefGoogle Scholar
Sereno, P. C., Forster, C. A., Rogers, R. R. & Moneta, A. M. 1993. Primitive dinosaur skeleton from Argentina and the early evolution of the Dinosauria. Nature 361, 64–66.CrossRefGoogle Scholar
Sereno, P. C., McAllister, S. & Brusatte, S. L. 2005. Taxonsearch: a relational database for suprageneric taxa and phylogenetic definitions. PhyloInformatics 8, 1–21.Google Scholar
Sereno, P. C., Martínez, R. N. & Alcober, O. A. 2013. Osteology of Eoraptor lunensis (Dinosauria, Sauropodomorpha). Basal sauropodomorphs and the vertebrate fossil record of the Ischigualasto Formation (Late Triassic: Carnian-Norian) of Argentina. Journal of Vertebrate Paleontology Memoir 12, 83–179.Google Scholar
Stocker, M. R., Zhao, L. J., Nesbitt, S. J., Wu, X. C. & Li, C. 2017. A short-snouted, Middle Triassic phytosaur and its implications for the morphological evolution and biogeography of Phytosauria. Scientific Reports 7, 46028.CrossRefGoogle ScholarPubMed
Stocker, M. R. & Butler, R. J. 2013. Phytosauria. In Nesbitt, S. J., Desojo, J. B. & Irmis, R. B. (eds) Anatomy, phylogeny and palaeobiology of early archosaurs and their kin, 379, 91–117. Geological Society, London, Special Publications.Google Scholar
Sues, H.-D., Olsen, P. E., Carter, J. G. & Scott, D. M. 2003. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 23, 329–43.CrossRefGoogle Scholar
Tolchard, F., Nesbitt, S. J., Desojo, J. B., Viglietti, P., Butler, R. J. & Choiniere, J. N. 2019. ‘Rauisuchian’ material from the lower Elliot Formation of South Africa and Lesotho: Implications for Late Triassic biogeography and biostratigraphy. Journal of African Earth Sciences 160, 103610.CrossRefGoogle Scholar
Trotteyn, M. J., Arcucci, A. B. & Raugust, T. 2013. Proterochampsia: an endemic archosauriform clade from South America. In Nesbitt, S. J., Desojo, J. B. & Irmis, R. B. (eds) Anatomy, phylogeny and palaeobiology of early archosaurs and their kin, 379, 59–90. Geological Society, London, Special Publications.Google Scholar
Von Baczko, M. B. 2018. Rediscovered cranial material of Venaticosuchus rusconii Enables the first jaw biomechanics in Ornithosuchidae (Archosauria: Pseudosuchia). Ameghiniana 55, 365–80.CrossRefGoogle Scholar
Von Baczko, M. B. & Desojo, J. B. 2016. Cranial anatomy and palaeoneurology of the archosaur Riojasuchus tenuisceps from the Los Colorados Formation, La Rioja, Argentina. PLoS ONE 11, e0148575.CrossRefGoogle ScholarPubMed
Von Baczko, M. B. & Ezcurra, M. D. 2013. Ornithosuchidae: a group of Triassic archosaurs with a unique ankle joint. In Nesbitt, S. J., Desojo, J. B. & Irmis, R. B. (eds) Anatomy, phylogeny and palaeobiology of early archosaurs and their kin, 379, 187–202. Geological Society of London, Special Publication.Google Scholar
Von Baczko, M. B., Desojo, J. B. & Pol, D. 2014. Anatomy and phylogenetic position of Venaticosuchus rusconii Bonaparte, 1970 (Archosauria, Pseudosuchia), from the Ischigualasto Formation (Late Triassic), La Rioja, Argentina. Journal of Vertebrate Paleontology 34, 1342–1356.CrossRefGoogle Scholar
Von Baczko, M. B. & Ezcurra, M. D. 2016. Taxonomy of the archosaur Ornithosuchus: reassessing Ornithosuchus woodwardi Newton, 1894 and Dasygnathoides longidens (Huxley 1877). Earth and Environmental Science Transactions of the Royal Society of Edinburgh 106, 199–205.CrossRefGoogle 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: Biological Sciences 248, 53–134.Google Scholar
Watson, D. M. S. 1917. A sketch-classification of the pre-Jurassic tetrapod vertebrates. Proceedings of the Zoological Society of London 1917, 167–86.Google Scholar
Weinbaum, J. C. 2011. The skull of Postosuchus kirkpatricki (Archosauria: Paracrocodyliformes) from the Upper Triassic of the United States. PaleoBios 30, 18–44.Google Scholar
Witmer, L. M., Ridgely, R. C., Dufeau, D. L. & Semones, M. C. 2008. Using CT to peer into the past: 3D visualisation of the brain and ear regions of birds, crocodiles and nonavian dinosaurs. In Endo, H. & Frey, R. (eds) Anatomical imaging: towards a new morphology, 67–87. Tokyo: Springer Verlag.CrossRefGoogle Scholar
Zaher, M., Coram, R. A. & Benton, M. J. 2019. The Middle Triassic procolophonid Kapes bentoni: computed tomography of the skull and skeleton. Papers in Palaeontology 5, 111–38.CrossRefGoogle Scholar
Zittel, K. A. 1887–1890. Handbuch der Palaeontologie. 1. Abteilung: Palaeozoologie, 3. Munich and Leipzig. 900 pp.Google Scholar
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