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Proquest Dissertations University of Alberta NEW PORASPIDINE HETEROSTRACANS FROM THE LOCHKOVIAN (EARLY DEVONIAN) MAN ON THE HILL LOCALITY, MACKENZIE MOUNTAINS, NORTHWEST TERRITORIES, CANADA, AND THE PHYLOGENY AND EVOLUTIONARY HISTORY OF PORASPIDINAE by Jessica Rae Hawthorn © 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 Edmonton, Alberta Spring 2009 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 Ottawa ON K1A0N4 OttawaONK1A0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-54698-7 Our file Notre r6f6rence ISBN: 978-0-494-54698-7 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 Nntemet, 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. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in this et des droits moraux qui protege cette these. Ni thesis. Neither the thesis nor la these ni des extraits substantiels de celle-ci substantial extracts from it may be ne doivent etre imprimes ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission. In compliance with the Canadian Conformement a la loi canadienne sur la Privacy Act some supporting forms protection de la vie privee, quelques may have been removed from this formulaires secondares ont ete enleves de thesis. cette these. While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. •+• Canada ABSTRACT Poraspidine heterostracans are extinct, jawless vertebrates that first appear in the Late Silurian, and were common components of Early Devonian faunas. The MOTH locality in the Mackenzie Mountains of northwestern Canada bears four new species of the genus Poraspis Kiaer, 1930, and two species previously known from Spitsbergen. These four new species are described, differing from previously described species in characteristics of ornamentation, sensory system pores, and in the dimensions of the dorsal shield. With the description of these new species, there are now seventeen species of Poraspis known. Phylogenetic analyses of the subfamily Poraspidinae and the genus Poraspis are here attempted for the first time, but few firm conclusions can be reached due to insufficient data and high levels of polymorphism. The significance of the MOTH poraspidine fauna is explored in the context of paleobiogeographic hypotheses, paleoenvironmental interpretation, and broader morphological trends in poraspidine evolution. ACKNOWLEDGEMENTS I would like to extend the my sincere gratitude to my supervisor, Dr. Mark V. H. Wilson, for providing access to the spectacular collection of specimens from MOTH, as well as for his support, encouragement, and guidance, and acting as my thesis supervisor. I am also grateful to Dr. Michael W. Caldwell and Dr. Brian D. E. Chatterton for their many insights throughout my time at the University of Alberta and for acting on my supervisory committee. Allan Lindoe is owed many, many thanks for his amazing preparation work on the MOTH specimens, and for his highly valued advice and feedback on my own preparation work. Thanks also go to Victoria and Jack Wiercinski for their assistance with translations from Polish, and to Victoria Wiercinski and Kyla Johnson for advice with graphics programs. This work would not have been possible without support from Dr. Mark V. H. Wilson, the Natural Sciences and Engineering Research Council of Canada (NSERC PGS-M), the Faculty of Graduate Studies and Research (J. Gordin Kaplan Graduate Student Award, Walter H. Johns Graduate Student Fellowship), the Province of Alberta (Graduate Student Scholarship), the Graduate Students' Association (Professional Development Grant), and the Department of Biological Sciences (Graduate Teaching Assistantship Award, Graduate Intern Tuition Supplement, Department of Biological Sciences Travel Award). Thanks go to anyone and everyone who has ever collected materials at the MOTH locality, and to all my fellow paleo graduate students, the postdoctoral researchers for their feedback, encouragement, and lively and stimulating discussions. Thanks also go to all the U of A staff who helped me out along the way. Lastly, to my friends and family who have always supported me, believed in me, and helped to keep me (mostly) sane. - J. R. H. TABLE OF CONTENTS ABSTRACT ACKNOWLEDGEMENTS LIST OF TABLES LIST OF FIGURES LIST OF APPENDICES I. GENERAL INTRODUCTION 1 Study Objectives 6 Abbreviations 6 Anatomical Abbreviations 6 Systematic History 7 Suprageneric Taxonomy 7 Poraspis 9 Locality and Age 11 Heterostracans from the MOTH Locality 16 Literature Cited 18 II. REVIEW OF PORASPIS KIAER (CYATHASPIDIDAE, PORASPIDINAE) AND A DESCRIPTION OF NEW PORASPIDINE TAXA FROM THE LOWER DEVONIAN MOTH LOCALITY, NORTHWEST TERRITORIES, CANADA Introduction 28 Anatomical Terminology 28 Dorsal Shield Measurements 30 Locality and Age 32 Materials and Methods 36 Fossil Materials 36 Preparation of Specimens 36 Photography, Illustration, and Measurement 37 Abbreviations 37 Institutional and Locality Abbreviations 37 Measurement Abbreviations 37 Systematic Paleontology 38 Poraspidinae 38 Poraspis 39 Poraspis sericea 42 Poraspis rostrata 43 Poraspis cf. P. rostrata 46 Poraspis sturi 51 Poraspis pompeckji 54 Poraspis siemiradzkii 55 Poraspis simplex 56 Poraspis barroisi 56 Poraspis polaris 58 Poraspis cf. P. polaris 62 Poraspis heintzae 65 Poraspis brevis 65 Poraspis cracens 67 Poraspis thules 67 Poraspis parmula 68 Poraspis sp. nov. A 69 Poraspis sp. nov. B 72 Poraspis sp. nov. C 75 Poraspis sp. nov. D 77 Poraspis sp. indet. 80 Discussion 82 Growth and Variation in Size 82 Dorsal Shield Measurements as Diagnostic Characters 85 Ornamentation 88 Conclusions 90 Literature Cited 91 PHYLOGENETIC ANALYSIS OF PORASPIDINAE Introduction 106 Materials and Methods 108 Abbreviations 109 Institutional and Locality Abbreviations 109 Results 109 Phylogenetic Analysis of Genus Poraspis 109 Phylogenetic Analysis of Subfamily Poraspidinae 114 Discussion 117 Phylogenetic Analysis of Genus Poraspis 117 Phylogenetic Analysis of Subfamily Poraspidinae 118 General Discussion 119 Conclusions 120 Literature Cited 121 EVOLUTIONARY HISTORY OF PORASPIDINAE Introduction 130 Abbreviations 131 Institutional and Locality Abbreviations 131 Regional Abbreviations 131 Species Abbreviations 131 Geographic and Temporal Distribution of Poraspidines 132 Trends in Poraspidine Evolution 139 Paleoenvironment of Poraspidines 144 Conclusions 147 Literature Cited 148 CONCLUSIONS Conclusions 154 Literature Cited 156 LIST OF TABLES Pa2e Table Caption (31) TABLE 2.1. Terminology of measurements of the dorsal and ventral shields used in the current and previous studies of poraspidines. (34) TABLE 2.2. Measurements (in mm) of the dorsal shields of specimens of Poraspis from the MOTH locality used in the current study. (35) TABLE 2.3. Indices derived from measurements (in mm) of the dorsal shields of specimens of Poraspis from the MOTH locality. (52) TABLE 2.4. Measurements (in mm) of the ventral shields of specimens of Poraspis from the MOTH locality used in the current study. (142) TABLE 4.1. Median and pineal lengths of the dorsal shield of specimens of Poraspis polaris from Spitsbergen and the MOTH locality. LIST OF FIGURES Page Figure Caption (4) FIGURE 1.1. Reconstruction of Poraspis showing the nomenclature of the different components of the dermoskeleton. (5) FIGURE 1.2. Morphology of the dorsal shield of Poraspis. (12) FIGURE 1.3. Map indicating general location of the MOTH locality in the Mackenzie Mountains of the Northwest Territories, Canada, UALVP 129. Equivalent to GSC locality 69014 and lies in Unit 10 of Section 43 of Gabrielse et al. (1973). (33) FIGURE 2.1. Measurements used on the dorsal shields of poraspidines. (44) FIGURE 2.2. Poraspis rostrata. A, UALVP 41398; B, UALVP 41876. (47) FIGURE 2.3. Poraspis cf. P. rostrata. A, UALVP 32744; B, UALVP 47059. (48) FIGURE 2.4. Poraspis cf. P. rostrata. A, UALVP 32881; B, UALVP 41359. (59) FIGURE 2.5. Poraspispolaris. A, UALVP 23394; B, UALVP 23436; C, UALVP 32783. (60) FIGURE 2.6. Poraspis polaris. A, UALVP 32785; B, UALVP 41423; C, UALVP 47060. (63) FIGURE 2.7. Poraspis cf. P. polaris. A, UALVP 41382; B, UALVP 43053; C, UALVP 43054; D, UALVP 47064. (70) FIGURE 2.8. Poraspis sp. nov. A. UALVP 49532. A, dorsal view; B, ventral view. (73) FIGURE 2.9. Poraspis sp. nov. B. A, UALVP 32886; B, UALVP 45941; C, UALVP 47062. (76) FIGURE 2.10. Poraspis sp. nov. C. UALVP 43232. (78) FIGURE 2.11. Poraspis sp. nov. D. UALVP 41426. (81) FIGURE 2.12. Poraspis sp. indet. UALVP 32820.
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