University of Alberta Lower Palaeozoic arthropod traces: Clarity in the mud by Stacey Gibb A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements for the degree of Doctor of Philosophy Earth and Atmospheric Sciences ©Stacey Gibb Spring 2011 Edmonton, Alberta Permission is hereby granted to the University of Alberta Libranes to reproduce single copies of this thesis and to lend or sell such copies for private, scholarly or scientific research purposes only Where the thesis is converted to, or otherwise made available in digital form, the University of Alberta will advise potential users of the thesis of these terms The author reserves all other publication and other nghts in association with the copyright in the thesis and, except as herein before provided, neither the thesis nor any substantial portion thereof may be pnnted or otherwise reproduced in any material form whatsoever without the author's prior wntten penmssion Library and Archives Bibliotheque et 1*1 Canada Archives Canada Published Heritage Direction du Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington OttawaONK1A0N4 OttawaONK1A0N4 Canada Canada Your Tile Votre reference ISBN: 978-0-494-80955-6 Our file Notre r6f6rence ISBN: 978-0-494-80955-6 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library and permettant a la Bibliotheque et Archives Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par I'lnternet, preter, telecommunication or on the Internet, distribuer et vendre des theses partout dans le loan, distribute and sell theses monde, a des fins commerciales ou autres, sur worldwide, for commercial or non­ support microforme, papier, electronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats. 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May she forever rest in peace. ABSTRACT Ichnology is the study of trace fossils and therefore the inferred behaviour of organisms that may have created the ichnofossils. Through analyses of traces from numerous different localities in the southern Rocky Mountains of Canada, the coast of Wales, and the deserts of Morocco and central Australia, from the lower Cambrian to the Upper Devonian, numerous conclusions were made. From the lower Cambrian Gog Group of Canada, it is evident that the 'Cambrian Explosion' was indeed an explosion of epic portions, not only in the advent of hard exo- and endoskeletons, but also of novel behaviours. The Cambrian of Morocco provided evidence for a relationship between sea floor current directions and the orientations of the trace Selenichnites in a shallow clastic palaeoenvironment. The Ordovician of Morocco provided direct evidence of a putative tracemaker, Asaphellus aff.fezouataensis, in very close physical association with the trace that it made Rusophycus carleyi, and allowed me to speculate on the form of the, yet to be discovered, ventral appendages of that trilobite. This locality also provided further insights on the importance of Microbially Induced Sedimentary Structures in the formation and preservation of ichnofossils. The Devonian of Wales showed that trace fossils can provide the first evidence of the migration of an organism into a new environment (xiphosurans into fresh water); it provides additional evidence of preferential orientations of Selenichnites in relation to currents occurring at the surface of the substrate; and shows that turbulent hydrodynamic influences assisted in the creation of the traces and perhaps caused the trace maker to burrow into the substrate. The synergy of these apparently quite different chapters is that the ichnofossils were all plausibly made by arthropods. Preliminary neoichnological research provided a behavioural reason to justify the morphological distinctions between the ichnogenera Cruziana and Rusophycus. Further research is only going to provide more insight into the 'other dimension' of palaeontology - ichnology: therefore, the ethology of organisms that cannot be as definitely acquired from a static body fossil. ACKNOWLEDGEMENTS The academic exchange, support, and friendship of my supervisors, Brian Chatterton and George Pemberton, was invaluable. I also acknowledge Murray Gingras who was a valuable member of my supervisory committee. My examiners, both in the final and candidacy examinations, provided challenging, though entertaining and thought provoking experiences, for this I thank: Nigel Hughes, Rich Palmer, Lindsey Leighton, Alex Wolfe and Heather Proctor. Funding from the University of Alberta and Department of Earth & Atmospheric Sciences, in the form of scholarships, teaching assistantships, research assistantships, office space, and administrative assistance, and financial support from the NSERC Discovery Grants of B. Chatterton and G. Pemberton. The supervisory assistance of Derek Siveter while I was a 'visiting scholar' at the University of Oxford is greatly appreciated. I thank the University of Oxford, the Oxford University Museum of Natural History and Wolfson College for acceptance and hospitality. The following individuals, whose help time and effort was invaluable in the field: B. Chatterton, H. ait H'ssaine, R. McKellar, D. Molinaro, D. Nordheimer, and A. Madden. Institutions which either allowed and/or provided assistance for research include: Parks Canada (granting Research permits JNP- 2005-2178 and LL-2010-6392); Royaume du Maroc, Ministere de I'Energie, des Mines, de l'Eau et de l'Environnement (collection and export permits); in Australia, the Australian Museum, the Maryvale Station, and the Idracowra Station; Oxford University Museum of Natural History; and the University of Alberta and more specifically the Department of Earth & Atmospheric Sciences. I would like to thank the following for their support and assistance through most, if not all of my dissertation: my Mom and Dad, my sister and her family, my grandma, grandpa and other extended family members (you know who you are), R. McKellar, B. Frizzell, R. Madden, A. Lindoe, J. Bohun, J. Duffy, S. Hesse, K. Hibbard, and the many sports teams/clubs I have been involved with. Finally, it would not have been possible without the love and support of Art Madden. TABLE OF CONTENTS CHAPTER 1: INTRODUCTION Introduction 1 Localities 2 Cambrian southern Rocky Mountains of Canada 2 Cambrian of Morocco 3 Ordovician of Australia 3 Ordovician of Morocco 4 Devonian of Wales 4 Previous Research 5 Methodology 6 Format 7 References 9 CHAPTER 2: CAMBRIAN OF CANADA Introduction 11 Geographical Location 12 Geological Background 13 Systematic Palaeontology (Ichnology) 16 Cruziana d'Orbigny, 1842 17 Cruziana billingsi Fillion & Pickerill, 1990 18 Cruziana dispar (Linnarsson, 1869) 19 Cruziana irregularis Fenton & Fenton, 1937 20 Cruziana Jenningsi Fenton & Fenton, 1937 21 Cruziana mesodelta isp. nov. 23 Cruziana navicella Fenton & Fenton, 1937 24 Cruziana omanica Seilacher, 1970 26 Cruziana penicillata Gibb, Chatterton & Pemberton, 2009 26 Cruzianaplicata Crimes, Legg, Marcos & Arboleya, 1977 27 Cruzianaproblematica (Schindewolf, 1928) 29 Cruziana ramellensis (Legg, 1985) 30 Cruziana rugosa d'Orbigny, 1842 31 Diplichnites Dawson, 1873 32 Diplichnites obliquus isp. nov. 33 Diplichnites twelvetreesi (Chapman, 1928) 34 Monomorphichnus Crimes, 1970b 36 Monomorphichnus bilinearis Crimes, 1970b 36 Marcos & Arboleya, 1977 37 Monomorphichnus trilinearis isp. nov. 37 Rusophycus Hall, 1852 38 Rusophycus arizonensis (Seilacher, 1970) 39 Rusophycus eutendorfensis (Linck, 1942) 41 Rusophycus latus Webby, 1983 43 Rusophycus mesodeltus isp. nov. 45 Rusophycus radwanskii Alpert, 1976 47 Rusophycus subnotous isp. nov. 48 Rusophycus unilobus (Seilacher, 1970) 49 Rusophycus vie torus isp. nov. 50 Conclusions 52 References 57 CHAPTER 3: CAMBRIAN OF MOROCCO Introduction 98 Geological and Geographical Setting 99 Material 100 Systematic Ichnology 101 Selenichnites (Romano and Whyte, 1987) 101 Selenichnites tacfihtus n. isp. 101 Interpretation 105 References 110 CHAPTER 4: ORDOVICIAN OF MOROCCO Introduction 122 Regional Setting 123 Geographical Location 124 Geological Setting 124 Palaeoenvironmental Reconstruction 124 Smoking Gun Scenario 127 Hard Evidence 128 Soft Evidence 130 Systematic Palaeontology 131 Ichnogenus Rusophycus Hall, 1852 131 Rusophycus carleyi (James, 1885) 131 Family Asaphidae Burmeister, 1843