sign in sign up
<<
Home , Doswelliidae, Loricata

Spine = 33mm SP379 cover art 2 amended 19-7-13_SP246 finalart/5.05 19/07/2013 14:26 Page 1 A A P a r n c l a a h t e o o o s m b a y u i o ,

Anatomy, Phylogeny and r

l s P o

h a g

y Anatomy, Phylogeny and

Palaeobiology of Early n y l d

o o

g t and their Kin f h e

E e n Palaeobiology of Early a i y r r

Edited by

l a K y n

S. J. Nesbitt, J. B Desojo and R. B. Irmis i n d Archosaurs and their Kin

Archosaurs, an important group that includes today’s and , arose Edited by during the in the aftermath of the greatest mass of all time. In the last S. J. Nesbitt, J. B Desojo and R. B. Irmis 20 years, our understanding of the early evolution of the group has improved substantially with the discovery of new fossils and species of early archosaurs and their closest relatives, a better understanding of the relationships of these , and Geological new insights into their palaeobiology. In to synthesize Society Geological Society these new data, researchers of early archosaurs from around Special the world met at the first symposium of early Publication Special Publication 379 evolution at the IV Congreso Latinoamericano de Paleontología 379 de Vertebrados (September 20 11) in San Juan, . This symposium facilitated collaboration and strove to paint a better understanding of these E extraordinary animals. The resultant body of work is a state-of-the-art examination of S d . J i

early archosaur groups and their close relatives including historical, anatomical, t . e N biogeographical, evolutionary and palaeobiological data. This contribution furthers our d

e b s y knowledge of the anatomy, relationships, and palaeobiology of species-level taxa as well b

i t

as more global patterns of archosaur evolution during the Triassic. t ,

J . B

Visit our online bookshop: http ://www.geolsoc.org.uk/bookshop D e s o

Geological Society web site: http ://www.geolsoc.org.uk j o

a n d

R . B . I r m i s

Cover illustration: , an early stem relative of crocodylians, from the Middle Triassic of Brazil. Illustration by Voltaire Paes Neto (20 11, graphite and digital art).

The Geological Society

Downloaded from http://pubs.geoscienceworld.org/books/book/chapter-pdf/3911064/9781862396395_frontmatter.pdf by guest on 01 October 2021 Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and their Kin

Downloaded from http://pubs.geoscienceworld.org/books/book/chapter-pdf/3911064/9781862396395_frontmatter.pdf by guest on 01 October 2021 The Geological Society of London Books Editorial Committee

Chief Editor Rick Law (USA)

Society Books Editors Jim Griffiths (UK) Dave Hodgson (UK) Howard Johnson (UK) Phil Leat (UK) Daniela Schmidt (UK) Randell Stephenson (UK) Rob Strachan (UK) Mark Whiteman (UK)

Society Books Advisors Ghulam Bhat (India) Marie-Franc¸oise Brunet (France) James Goff (Australia) Mario Parise (Italy) Satish-Kumar (Japan) Marco Vecoli (Saudi Arabia) Gonzalo Veiga (Argentina) Maarten de Wit (South Africa)

Geological Society books refereeing procedures

The Society makes every effort to ensure that the scientific and production quality of its books matches that of its journals. Since 1997, all book proposals have been refereed by specialist reviewers as well as by the Society’s Books Editorial Committee. If the referees identify weaknesses in the proposal, these must be addressed before the proposal is accepted. Once the book is accepted, the Society Book Editors ensure that the volume editors follow strict guidelines on refereeing and quality control. We insist that individual papers can only be accepted after satisfactory review by two independent referees. The questions on the review forms are similar to those for Journal of the Geological Society. The referees’ forms and comments must be available to the Society’s Book Editors on request. Although many of the books result from meetings, the editors are expected to commission papers that were not presented at the meeting to ensure that the book provides a balanced coverage of the subject. Being accepted for presentation at the meeting does not guarantee inclusion in the book. More information about submitting a proposal and producing a book for the Society can be found on its website: www.geolsoc.org.uk.

It is recommended that reference to all or part of this book should be made in one of the following ways:

Nesbitt, S. J., Desojo,J.B.&Irmis, R. B. (eds) 2013. Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and their Kin. Geological Society, London, Special Publications, 379.

Trotteyn, M. J., Arcucci,A.B.&Raugust, T. 2013. Proterochampsia: an endemic archosauriform from . In:Nesbitt, S. J., Desojo,J.B&Irmis, R. B. (eds) Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and their Kin. Geological Society, London, Special Publications, 379, 59–90. First published online June 11, 2013, http://dx.doi.org/10.1144/SP379.23.

Downloaded from http://pubs.geoscienceworld.org/books/book/chapter-pdf/3911064/9781862396395_frontmatter.pdf by guest on 01 October 2021 GEOLOGICAL SOCIETY SPECIAL PUBLICATION NO. 379

Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and their Kin

EDITED BY

S. J. NESBITT University of Washington, USA

J. B. DESOJO Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’, Argentina

and

R. B. IRMIS Natural History Museum of Utah, USA

2013 Published by The Geological Society London

Downloaded from http://pubs.geoscienceworld.org/books/book/chapter-pdf/3911064/9781862396395_frontmatter.pdf by guest on 01 October 2021 THE GEOLOGICAL SOCIETY

The Geological Society of London (GSL) was founded in 1807. It is the oldest national geological society in the world and the largest in . It was incorporated under Royal Charter in 1825 and is Registered Charity 210161. The Society is the UK national learned and professional society for geology with a worldwide Fellowship (FGS) of over 10 000. The Society has the power to confer Chartered status on suitably qualified Fellows, and about 2000 of the Fellowship carry the title (CGeol). Chartered Geologists may also obtain the equivalent European title, European Geologist (EurGeol). One fifth of the Society’s fellowship resides outside the UK. To find out more about the Society, log on to www.geolsoc.org.uk. The Geological Society Publishing House (Bath, UK) produces the Society’s international journals and books, and acts as European distributor for selected publications of the American Association of Petroleum Geologists (AAPG), the Indonesian Petroleum Association (IPA), the Geological Society of America (GSA), the Society for Sedimentary Geology (SEPM) and the Geologists’ Association (GA). Joint marketing agreements ensure that GSL Fellows may purchase these societies’ publications at a discount. The Society’s online bookshop (accessible from www.geolsoc. org.uk) offers secure book purchasing with your credit or debit card. To find out about joining the Society and benefiting from substantial discounts on publications of GSL and other societies worldwide, consult www.geolsoc.org.uk, or contact the Fellowship Department at: The Geological Society, Burlington House, Piccadilly, London W1J 0BG: Tel. +44 (0)20 7434 9944; Fax +44 (0)20 7439 8975; E-mail: [email protected]. For information about the Society’s meetings, consult Events on www.geolsoc.org.uk. To find out more about the Society’s Corporate Affiliates Scheme, write to [email protected].

Published by The Geological Society from: The Geological Society Publishing House, Unit 7, Brassmill Enterprise Centre, Brassmill Lane, Bath BA1 3JN, UK The Lyell Collection: www.lyellcollection.org Online bookshop: www.geolsoc.org.uk/bookshop Orders: Tel. +44 (0)1225 445046, Fax +44 (0)1225 442836 The publishers make no representation, express or implied, with regard to the accuracy of the information contained in this book and cannot accept any legal responsibility for any errors or omissions that may be made. # The Geological Society of London 2013. No reproduction, copy or transmission of all or part of this publication may be made without the prior written permission of the publisher. In the UK, users may clear copying permissions and make payment to The Copyright Licensing Agency Ltd, Saffron House, 6–10 Kirby Street, London EC1N 8TS UK, and in the USA to the Copyright Clearance Center, 222 Rosewood Drive, Danvers, MA 01923, USA. Other countries may have a local reproduction rights agency for such payments. Full information on the Society’s permissions policy can be found at: www.geolsoc.org.uk/permissions

British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library. ISBN 978-1-86239-361-5 ISSN 0305-8719

Distributors For details of international agents and distributors see: www.geolsoc.org.uk/agentsdistributors Typeset by Techset Composition India (P) Ltd., Bangalore and Chennai, India. Printed by Berforts Information Press Ltd, Oxford, UK

Downloaded from http://pubs.geoscienceworld.org/books/book/chapter-pdf/3911064/9781862396395_frontmatter.pdf by guest on 01 October 2021 Participants from the First Symposium of Early Archosaur Evolution

Participants from the First Symposium of Early Archosaur Evolution at the IV Congreso Latinoamericano de Paleonto- logı´a de Vertebrados (September 2011) in front of a life-sized reconstruction of Fasolasuchus outside the Instituto y Museo de Ciencias Naturales, Universidad Nacional de San Juan, San Juan, Argentina. Participants from left to right: Tiago Raugust, Randall B. Irmis, David J. Gower, Bryan J. Small, Julia B. Desojo, M. Bele´n von Baczko, Andrea B. Arcucci, Jeremı´as R. A. Taborda, Martı´n D. Ezcurra, Corwin K. Sullivan, Sterling J. Nesbitt and Marcel Lacerda. Not pictured: Jonathas S. Bittencourt, Stephen L. Brusatte, Ignacio A. Cerda, Thomas Garner, Casey M. Holliday, Max C. Langer, Alexandre Liparini, Martins Bianca Mastrantonio, William G. Parker, Cesar L. Schultz, Andrey G. Sennikov, Nathan D. Smith, Michelle R. Stocker, Marı´a Jimena Trotteyn, Alan H. Turner, Jonathan C. Weinbaum and Sarah Werning.

Downloaded from http://pubs.geoscienceworld.org/books/book/chapter-pdf/3911064/9781862396395_frontmatter.pdf by guest on 01 October 2021 Contents

Participants from the First Symposium of Early Archosaur Evolution vii

NESBITT, S. J., DESOJO,J.B.&IRMIS, R. B. Anatomy, phylogeny and palaeobiology of early 1 archosaurs and their kin

EZCURRA, M. D., BUTLER,R.J.&GOWER, D. J. ‘’: the origin and early history of 9

SOOKIAS,R.B.&BUTLER, R. J. Euparkeriidae 35

SUES, H.-D., DESOJO,J.B.&EZCURRA, M. D. Doswelliidae: a clade of unusual armoured 49 archosauriforms from the Middle and

TROTTEYN, M. J., ARCUCCI,A.B.&RAUGUST, T. Proterochampsia: an endemic archosauriform 59 clade from South America

STOCKER,M.R.&BUTLER, R. J. Phytosauria 91

DALLA VECCHIA, F. M. Triassic pterosaurs 119

LANGER, M. C., NESBITT, S. J., BITTENCOURT,J.S.&IRMIS, R. B. Non-dinosaurian 157 Dinosauromorpha

VON BACZKO,M.B.&EZCURRA, M. D. : a group of Triassic archosaurs with a 187 unique ankle joint

DESOJO, J. B., HECKERT, A. B., MARTZ, J. W., PARKER, W. G., SCHOCH, R. R., 203 SMALL,B.J.&SULEJ, T. Aetosauria: a clade of armoured pseudosuchians from the Upper Triassic continental beds

NESBITT, S. J., BRUSATTE, S. L., DESOJO, J. B., LIPARINI, A., DE FRANc¸a, M. A. G., 241 WEINBAUM,J.C.&GOWER, D. J.

IRMIS, R. B., NESBITT,S.J.&SUES, H.-D. Early 275

RAUGUST, T., LACERDA,M.&SCHULTZ, C. L. The first occurrence of bonapartei 303 Romer 1971 (Archosauriformes, Proterochampsia) of the Middle Triassic of Brazil from the Santacruzodon Assemblage Zone, (Parana´ Basin)

NIEDZ´ WIEDZKI, G., BRUSATTE,S.L.&BUTLER,R.J.Prorotodactylus and Rotodactylus tracks: an 319 ichnological record of dinosauromorphs from the Early–Middle Triassic of Poland

LANGER,M.C.&FERIGOLO, J. The Late Triassic dinosauromorph Sacisaurus agudoensis 353 (Caturrita Formation; Rio Grande do Sul, Brazil): anatomy and affinities

SMALL,B.J.&MARTZ, J. W. A new from the Upper Triassic Chinle Formation of the 393 Eagle Basin, Colorado, USA

TABORDA, J. R. A., CERDA,I.A.&DESOJO, J. B. Growth curve of scagliai 413 Casamiquela 1960 (: Aetosauria) inferred from histology

Downloaded from http://pubs.geoscienceworld.org/books/book/chapter-pdf/3911064/9781862396395_frontmatter.pdf by guest on 01 October 2021 vi CONTENTS

MASTRANTONIO, B. M., SCHULTZ, C. L., DESOJO,J.B.&GARCIA, J. B. The braincase of 425 Prestosuchus chiniquensis (Archosauria: )

LIPARINI,A.&SCHULTZ, C. L. A reconstruction of the thigh musculature of the extinct 441 pseudosuchian Prestosuchus chiniquensis from the Dinodontosaurus Assemblage Zone (Middle Triassic Epoch), Santa Maria 1 Sequence, southern Brazil

DE FRANc¸a, M. A. G., LANGER,M.C.&FERIGOLO, J. The skull anatomy of 469 quartacolonia (Pseudosuchia: Suchia: Loricata) from the middle Triassic of Brazil

PARKER,W.G.&NESBITT, S. J. Cranial remains of gracilis (Pseudosuchia: 503 Poposauroidea) from the Upper Triassic, the distribution of the taxon, and its implications for poposauroid evolution

WEINBAUM, J. C. Postcranial skeleton of kirkpatricki (Archosauria: 525 ), from the Upper Triassic of the United States

HOLLIDAY,C.M.&NESBITT, S. J. Morphology and diversity of the mandibular symphysis of 555 archosauriforms

TURNER,A.H.&NESBITT, S. J. Body size evolution during the Triassic archosauriform radiation 573

Index 599

Downloaded from http://pubs.geoscienceworld.org/books/book/chapter-pdf/3911064/9781862396395_frontmatter.pdf by guest on 01 October 2021