Wild mountain thunder Echoes my quest My body aches but I'll not rest Quartz light to guide me Till sunrise leads My passion screams, my heart it bleeds Desert Plains by Judas Priest University of Alberta Description, paleoenvironmental and paleobiogeographical implications of Miocene fish faunas from Jabal Zaltan and Sahabi (Libya). by Thodoris Argyriou A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements for the degree of Master of Science in Systematics and Evolution Department of Biological Sciences ©Thodoris Argyriou Spring 2014 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. The author reserves all other publication and other rights in association with the copyright in the thesis and, except as herein before provided, neither the thesis nor any substantial portion thereof may be printed or otherwise reproduced in any material form whatsoever without the author's prior written permission I dedicate this thesis to both my biological as well as my expanded academic family. I would have not made it this far without their support. ABSTRACT The Libyan fossiliferous sites of Jabal Zaltan (early to middle Miocene) and Sahabi (late Miocene) have been excavated for decades and have produced an immense diversity of vertebrate fossils including fish. Fish remains from a recent field campaign at the two sites are described here in detail. The new, greatly expanded, ichthyofaunal lists add considerably to our knowledge of the fossil fish diversity present at the two sites. Several first occurrences of fish taxa in the African continent or the fossil record are also recorded. The increased knowledge of the two Libyan ichthyofaunas allows for accurate reconstructions of the paleoenvironments present in their vicinity at the time of the deposition of their sediments. The biogeographical affinities of the two faunas are also discussed in detail. Paleoichthyological data from Sahabi and other penecontemporaneous African ichthyofaunas are used to discuss various proposed scenarios for the paleohydrology of the Sahara during the Messinian. ACKNOWLEDGEMENTS This work was carried out between September 2011 and January 2014, at the Department of Biological Sciences, University of Alberta. The completion of this project would not have been possible without the help of many people I would like to thank below. Above all, I want to express my immense gratitude to my supervisor Dr. Alison Murray for believing in me and patiently guiding and supporting me through the whole process of my M.Sc. program in Systematics and Evolution. Her ideas, mentoring and feedback were indispensable to my academic advancement, a significant part of which is reflected in this thesis. It is a great honor for me to have been her student. My sincere gratefulness is extended to the members of my committee, Dr. Michael Caldwell and Dr. Robert Losey, as well as the “arm’s length” examiner, Dr. Cynthia Paszkowski, for accepting to be a part of my committee and for their patience in reviewing my project and thesis particularly at so difficult a time as the Christmas break. I would especially like to thank Drs. Noel Boaz and Paris Pavlakis for their vital support and contributions at various stages of my academic career and especially for giving me the opportunity to participate in the 2008 and 2010 field missions to the Libyan sites of Sahabi and Jabal Zaltan. I am also indebted to them for entrusting me with the study of the Libyan fish fossils that formed the basis of this project. It is difficult to express how thankful I am to Dr. Pavlakis for his unofficial supervision… via Skype. Drs. Ahmed Muftah and Wilma Wessels provided me with invaluable information on the two fossil sites that produced the faunal diversities presented here and I am very thankful for that. I am much obliged to Dr. Todd Cook for aiding me significantly with the study of the Libyan elasmobranchs and teaching me all I know about fossil cartilaginous fishes. I am also grateful to Dr. Robert Holmes for the useful conversations we had. This project would not have been feasible without the help of the E.L.N.R.P. members and especially the 2010 E.L.N.R.P. field crew (Dr. Noel Boaz, Dr. Paris Pavlakis, Dr. Ahmed Muftah, Dr. Wilma Wessels, Dr. Efthimia Pavlakis, Dr. Dimitrios Michailidis and Mr. Tzouma Bohlaigha) who assisted me in collecting the sample presented in this project. It is an honor for me to extend my deepest gratitude to Drs. Olga Otero and Aurélie Pinton (iPHEP) for contributing to this project in numerous ways like providing comments and ideas, arranging my stay in Poitiers during the summer of 2013 and allowing me to access their collections and unpublished data. Dr. Otero is also warmly thanked for her very insightful comments on this thesis and her priceless contributions to the advancement of my academic career. Dr. Georgios Theodorou, as my undergraduate supervisor, played an instrumental role in fueling my interest in paleontology and in my decision to focus on paleoichthyology. I very much appreciated his support and ideas over the years. Special thanks go to all the people of the University of Alberta Laboratory for Vertebrate Paleontology and the Museum of Geology and Paleontology of the University of Athens for the interesting discussions we had and, most of all, for putting up with me throughout my years as a university student. My friends are also warmly thanked for sustaining my moral high throughout the long Edmontonian winters. The identification of the rich fossil fish assemblages from Libya would have been impossible without access to extensive comparative material and literature. Along those lines I want to acknowledge the following staff and institutions for lending specimens or providing access to their collections: Dr. Philippe Béarez (MNHN), Dr. Kathlyn M. Stewart (CMN), Dr. John P. Friel and Charles M. Dardia (CUMV), Dr. Douglas Nelson (UMMZ) and of course the staff at the University of Alberta Libraries. Funding for this research was provided by the Department of Biological Sciences of the University of Alberta, the University of Alberta Graduate Students Association and the Natural Sciences and Engineering Research Council of Canada Discovery Grant 327448 to Alison M. Murray. Last but not least, the unconditional moral and financial support of my family played a significant role in the completion of this work. Words cannot describe how grateful I am to them. TABLE OF CONTENTS Chapter 1: General Introduction………...………………………………………..1 Chapter 2: Fish fossils from the early to middle Miocene of Jabal Zaltan, Libya……………………………………………………………………………..15 Chapter 3: Fish diversity and paleoenvironments from the late Miocene of Sahabi, Libya. Implications for tracing Miocene Saharan Rivers……………...139 Chapter 4: General discussion and conclusions………………...……………..298 Appendix……………………………………………………………………….307 LIST OF TABLES TABLE 2-1. Fossil fish taxa present in each locality sampled in Jabal Zaltan...........................................................................................................111, 112 TABLE 3-1. Aspects of ecology and paleoenvironmental preferences of the Sahabi actinopterygians………….………………………………………..257-259 Appendix. Table of recent comparative material examined……………...308-309 LIST OF FIGURES FIGURE 2-1. Map of Libya and Jabal Zaltan with the position of the localities that produced fish fossils...………………………………………………….…..114 FIGURE 2-2. Elasmobranch teeth recovered from Jabal Zaltan. Odontaspididae – Carcharhinidae……………………………………………………………….....116 FIGURE 2-3. Elasmobranch teeth recovered from Jabal Zaltan. Carcharhinidae – Myliobatidae……………………………………………………………………118 FIGURE 2-4. Elasmobranch teeth recovered from Jabal Zaltan. Myliobatidae – Rhinopteridae………………………………………………………...................120 FIGURE 2-5. Polypterid and osteoglossid fossils from Jabal Zaltan..…...........122 FIGURE 2-6. Unidentified teleost teeth from Jabal Zaltan…….……………...124 FIGURE 2-7. Characiform teeth from Jabal Zaltan. Characiformes indet. – Alestes/Brycinus.………………………………………………………………..126 FIGURE 2-8. Characiform teeth from Jabal Zaltan. Sindacharax sp ......……..128 FIGURE 2-9. Siluriform fossils from Jabal Zaltan. Siluriformes indet. A – cf. Synodontis sp………………………………………………………………..130 FIGURE 2-10. Siluriform fossils from Jabal Zaltan. Claroteidae………..……132 FIGURE 2-11. Clariid and unidentified perciform fossils from Jabal Zaltan…134 FIGURE 2-12. Perciform fossils from Jabal Zaltan. cf. Semlikiichthys sp. – Sparidae…………………………………………………………………………136 FIGURE 2-13. Perciform fossils from Jabal Zaltan. Latidae………………….138 FIGURE 3-1. Map of Libya and Sahabi……………………………….………261 FIGURE 3-2. Simplified stratigraphic log showing the succession of sediments exposed at Sahabi……………………………………………………………….263 FIGURE 3-3. Polypterid, cyprinid and alestid fossils from Sahabi……………265 FIGURE 3-4. Alestid and siluriform fossils from Sahabi. Alestes/Brycinus teeth……………………………………………………………………………..267 FIGURE 3-5. Siluriform fossils from Sahabi……………………………….....269 FIGURE 3-6. Bagrid fossils from Sahabi. Bagrus sp………………………….271 FIGURE 3-7. Claroteid fossils from Sahabi…………………………………...273 FIGURE 3-8. Claroteid fossils from Sahabi.