University of Alberta

University of Alberta Select Devonian Proetid Trilobites of southern Morocco by Darrin Molinaro A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements for the degree of Master of Science Department of Earth and Atmospheric Sciences © Darrin Molinaro Spring 2012 Edmonton, Alberta Permission is hereby granted to the University of Alberta Libraries 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. 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Canada Abstract The Tropidocoryphinae and Cornuproetinae represent two subfamilies of proetid trilobites (Class Trilobita, Order Proetida) found throughout the Lower and Middle Devonian of southern Morocco's Anti-Atlas region. Surveys of the various trilobite-bearing horizons within three separate Moroccan basins have yielded a diverse and well preserved collection of genera from both subfamilies. Given the exceptional (complete and articulated) nature of the specimens, systematic description and cladistic analysis, in conjunction with previously described Moroccan and Central European species of the two subfamilies, was undertaken. In total, one new genus (Pontoproetus) and 11 new species (Astycoryphe ditropidia, Tropidocoryphe taharajatensis, T. lahfirensis, T. sculptaridgiata, Diademaproetus rudimentus, D. issoumourensis, D. corrugatus, D. auxiliairus, D. langus, Pontoproetus truncatus, and P. granulosus) are described and the relationships of Tropidocoryphe-Astycoryphe and Cornuproetus-Diademaproetus- Pontoproetus are discussed. Acknowledgements Many special thanks are due to my supervisors Dr. Brian Chatterton and Dr. Lindsey Leighton, for giving me the opportunity to work with them, providing much needed guidance and support, and putting up with my "writing" style. Both of them have provided me with numerous invaluable skills, making me a much better scientist than when I first began this work. Funding for this work was provided by a National Science and Engineering Research Council of Canada Discovery Grant to B. D. E. Chatterton. I would also like to thank the other member of my committee, Dr. Felix Sperling, for his time and comments. Thanks are also due to: Stacey Gibb, for support and guidance beyond measure; Ryan McKellar, for help with PAUP and figure guidelines; Frank Forcino, Emily Stafford, Ben Collins, and Michael Burns for intellectual discussions and support; Allan Lindoe, for the preparation of some specimens, and Kevin Brett, for fieldwork and some original stratigraphic data. I am also in debt to the Department of Earth and Atmospheric Sciences, for their financial support via teaching assistantships and office space. I would also like to thank our Moroccan guide and preparator Hammi Ait H'ssaine, his preparator brother Amur Ait H'ssaine, and their families for their hospitality, support and preparation work on some of the examined specimens; the Berber miners encountered during this work for locating and working the trilobite horizons to produce the bulk of specimens in this study; and the Ministere de l'Energie et des Mines in Rabat, for providing the permits necessary for this work. Lastly, great thanks is also due to my family, in particular my parents for their never ending support and encouragement. Without them I would have never been able to do this work and accomplish what I have. To my grandparents, many thanks for putting up with me not coming home that often and for the many long and always interesting phone conversations. Table of Contents Chapter 1: Introduction Introduction 1 Location and geology 2 Examined localities 5 Tindouf Basin 5 Ma'der Basin 6 Tafilalt Basin 8 Purpose and scope 8 Previous research 11 Format 12 Literature cited 18 Chapter 2: Systematic Palaeontology I Introduction 23 Previous work 25 Materials and methods 27 Stratigraphy 28 Tindouf basin 29 Ma'der basin 30 Terminology 31 Phylogenetic analysis 32 Taxa examined in the analysis 35 Characters and character states 37 Cephalon 42 Glabella 54 Thorax 56 Pygidium 57 Characters not included in the analysis 63 Results 63 Discussion and conclusions 65 Systematic palaeontology 70 Order Proetida Fortey and Owens, 1975 70 Astycoryphe Richter and Richter, 1919 70 Astycoryphe ditropidia new species 72 Tropidocoryphe Novak, 1890 78 Tropidocoryphe amuri Chatterton et al., 2006 79 Tropidocoryphe taharajatensis new species 80 Tropidocoryphe lahfirensis new species 86 Tropidocoryphe sculptaridiata new species 91 Literature Cited 136 Chapter 3: Systematic Palaeontology II Introduction 142 Previous work 144 Stratigraphy and geology 146 Tindouf basin 146 Ma'der basin 148 Tafilalt basin 151 Terminology 152 Materials and methods 152 Phylogenetic analysis 157 Taxa examined in the analysis 159 Characters and character states 164 Cephalon 165 Thorax 180 Pygidium 181 Characters excluded from the analysis 186 Results 188 Discussion and conclusions 190 Systematic Palaeontology 196 Order Proetida Fortey & Owens, 1975 196 Cornuproetus Richter & Ricter, 1949 196 Diademaproetus Alberti, 1964 199 Diademaproetus praecursor Alberta, 1969 202 Diademaproetus mohamedi Chatterton et al., 2006 204 Diademaproetus rudimentus new species 206 Diademaproetus issoumourensis new species 213 Diademaproetus corrugatus new species 218 Diademaproetus auxiliarus new species 223 Diademaproetus langus new species 229 Pontoproetus new genus 234 Pontoproetus truncatus new species 235 Pontoproetus granulosus new species 240 Literature Cited 304 Chapter 4: Conclusions Synthesis 309 Future work 312 Literature Cited 314 List of Tables Chapter 2 Table 2-1. Table outlining included and omitted taxa for the phylogenetic analysis. Images used in coding of specimens are indicated as the source, while material indicates the completeness of specimens being coded. Reasoning provides explanation as to why the taxon was either included or excluded from the analysis. All remaining species not outlined in the table were omitted either based on lack of material (less than 50% of characters could be coded), poor documentation, or not being located in or around the Rheic Ocean of the Devonian. 130 Table 2-2. Results of character state statistical tests on a character by character, state by state basis. Use of Mann-Whittney U, ANOVA, and Hoteling's t-test were dependent on data type. All tests were measured against a 95% confidence interval. When two or more character states are present, a Bonferroni correction adjusted p-value was used to test at the 95% confidence interval. Grey shaded cells represent results which should be used with caution as some of their populations (character states) do not have large enough sample sizes to be statistically accurate. 134 Chapter 3 Table 3-1. Table outlining included and omitted taxa for the phylogenetic analysis. Images used in coding of specimens are indicated as the source, while material indicates the completeness of specimens being coded. Reasoning provides explanation as to why the taxon was either included or excluded from the analysis. All remaining species not outlined in the table were omitted either based on their lack of material (less

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