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Upper Devonian Plant Microfossils from Eastern and Arctic Canada: Their Taxonomy and Palaeoecological Significance

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Upper Devonian Plant Microfossils from Eastern and Arctic Canada: Their Taxonomy and Palaeoecological Significance ,{ ;" UPPER DEVONIAN PLANT MICROFOSSILS AND PALAEOECOLOGY UPPER DEVONIAN PLANT MICROFOSSILS FROM EASTERN AND ARCTIC CANADA: THEIR TAXONOMY AND PALAEOECOLOGICAL SIGNIFICANCE by WAYNE WILFRED BRIDEAUX, B.Sc. A Thesis Submitted to the Faculty of Graduate Studies in Partial Fulfillment of the Requirements for the degree Master of Science McMaster University May 1965 MASTER OF SCIENCE (1965) McMASTER UNIVERSITY (Biology) Hamilton, Ontario TITLE: Upper Devonian Plant Microfossils from Eastern and Arctic Canada: Their Taxonomy and Palaeoecological Significance AUTHOR: Wayne Wilfred Brideaux, B.Sc. (Carleton University) SUPERVISOR: Professor N. W. Radforth NUMBER OF PAGES: xii, 163. SCOPE AND CONTENTS: Plant microfossils are extracted from strata of Upper Devonian age occurring at two previously uninvestigated localities in Eastern Canada and in the Canadian Arctic. The extraction procedure includes a new permanent palynological mounting technique using corn syrup and Permount, a synthetic resin. In addition, a method of eliminating by-product in residues after hydrofluoric acid, is also described. A number of miospore and acritarch form species are described and figured; nine new form species of miospores are proposed. The miospore species recovered from both localities indicate a geological age of early Upper Devonian (Frasnian) for the enclosing strata. A new way of assessing qualitative, regional palaeofloristic changes is provided by tabulation of morphological characteristics of individual miospores. The use of miospore species in defining Upper Devonian microfloral provinces and Upper Devonian-Lower Carboniferous migration of parent forms is demonstrated. ii PREFACE Authors of earlier theses in the field of palynology have customarily included somewhat detailed resumes of the general history of palynology. As well, they have outlined the character­ istics of plant microfossils. Since the general treatments of these subjects are adequately set out elsewhere (McGregor, 1957; Wilkinson, 1959), the author thinks that remarks on these topics may be confined to subject matter related more directly to the present research. In view of this, historical observations will be limited to Devonian palynology and discussion of plant microfossils will be concerned with certain areas of definition of terms and nomenclature. The discussion of these facets of plant microfossils will have some bearing on the remarks of a historical nature, and therefore, the former will be the subject of this preface while the latter will be outlined in a later chapter. The concern of this preface is to make clear the meaning of several terms which have become vague or now have a different usage. 'Plant microfossil', 'microspore', 'miospore', and 'hystrichosphere' are among these terms. In this thesis, 'plant microfossil' will include generally microscopic, acid-resistant forms commonly referred to as pollen grains, spores, dispersed fragments of cuticle, sporangia, and iii vascular elements of plants, as well as organic microplankton. It will not include siliceous remains with plant characteristics or remains of calcareous algae. All spores described in this thesis are dispersed in a rock matrix and unattached to a parent plant. Under these conditions, botanical affinities of the spores are not likely to be deduced conclusively. To follow the common practice of dividing dispersed spores into megaspores and microspores (the former equal to or greater than 200µ and the latter less than 200µ in size) would be somewhat mislead­ ing. The word 'isospore' used in this context would also be mislead­ ing, because true isospores (male and female spores of equal or nearly equal size) fall within the size range of microspores. It would also suggest that the state of homospory was known to exist in the parent plant. To avoid these difficulties, the author will follow the practice of Guennel (1958) and refer to all spores and spore-like bodies under 200µ (including small megaspores, isospores, true micro­ spores, pre-pollens, and pollen grains) as miospores. The arbitrary lower size limit for megaspores (200µ) will be retained for convenience. Organic microplankton include the dinoflagellates, the hystrichospheres, and the acritarchs. Undoubted dinoflagellates have not been found in rocks older than the Triassic (Jansonius, 1962) and will not be discussed further. The terms, 'hystrichosphere' and 'acritarch' require further consideration. The term 'hystrichosphere' originally encompassed a wide variety of forms, including many of uncertain affinities. Evitt (1961) iv pointed out that the group was heterogenous, containing forms definitely related to the dinoflagellates and forms of uncertain or unknown affinities. ~vitt (1963) then proposed that 'hystrichosphere' be restricted to those forms showing an affinity to the dinoflagellates and placed the remaining organic microplankton of uncertain affinities in a new group, the Acritarcha. Downie, Evitt, and Sarjeant (1963) expanded this classification and regrouped genera to conform to this system. Since this seems to be the most satisfactory solution yet proposed to this problem, the author accepts this system and will classify appropriate forms in accordance with it. The question of which nomenclatural code should be followed for taxonomy involving acritarchs is not so easily resolved. Some workers (as in Eisenack, 1963) follow the International Code of Zoological Nomenclature when dealing with these forms, while others (Downie, Evitt, and Sarjeant, 1963) treat them as plants and use the International Code of Botanical Nomenclature. The author chooses to follow the latter practice in view of the reasonable arguments advanced by Downie, et !!!• (1963) and the fact that the author's definition of plant microfossils includes forms usually assigned to the Acritarcha. v ACKNOWLEDGEMENTS The author wishes to express his sincere gratitude to Dr. N. w. Radforth for his interest and encouragement throughout this investigation. The author is especially grateful for being allowed to use material, often collected with much difficulty, by Dr. Radforth, Dr. H. N. Andrews of the University of Connecticut, and Dr. T. L. Phillips. To Dr. Geoffrey Norris, post-doctoral fellow in palynology, the author extends his deepest appreciation for many hours of comment and discussion. The author also wishes to thru'l;k Mr. J. A. Wilfong and Mr. K. Ashdown for assistance in early stages of photomicrography, and Miss Joan Fink for undertaking the many long hours of typing. The financial assistance of a McMaster University Graduate Scholarship and Summer Assistantship and an Ontario Graduate Fellowship is gratefully acknowledged. vi TABLE OF CONTENTS SCOPE AND CONTENTS ii PREFACE iii ACKNOWLEDGEMENTS vi TABLE OF CONTENTS vii LIST OF TABLES xi LIST OF TEXT-FIGURES xii INTRODUCTION 1 DEVONIAN PALYNOLOGY 1 THE CONTRIBUTION OF EARLIER WORK 4 THE OBJECTIVES OF THE PRESENT RESEARCH 6 DEVONIAN PLANT PALAEOECOLOGY: CONCEPTS AND SOURCES OF INFORMATION 7 INTRODUCTION 7 ECOLOGY AND PALAEOECOLOGY 8 PALAEOECOLOGY AND FOSSILIZATION 9 THE CONCEPT FOR DEVONIAN PLANT PALAEOECOLOGY 10 PLANT MICROFOSSILS AND PALA:EX>ECOLOGICAL DATA 13 GEOLOGICAL AND GEOGRAPHICAL INFORMATION 19 INTRODUCTION 19 STRATIGRAPHY OF THE OKSE BAY FORMATION 19 STRATIGRAPHY OF THE ESCUMINAC FORMATION 23 STRATIGRAPHIC POSITION, LITHOLOGY, AND GEOGRAPHICAL LOCATION OF THE SAMPLE SITES 24 Goose Fiord, Ellesmere Island 24 Escuminac Bay, Quebec 27 vii DESCRIPTION AND DISCUSSION OF LABORATORY TECHNIQUES 28 EXTRACTION OF THE PLANT MICROFOSSILS 28 Introduction 28 Extraction Procedure: Description and Discussion 28 Crushing- 28 Acid Treatment­ 30 By-Product Removal­ 30 Oxidation- 32 Alkali Treatment­ 33 OTHER LABORATORY PROCEDURES 35 Photography 35 Marking of Slides 36 OBSERVATIONS ON THE DISCLOSED PLANT MICROFOSSILS AND TAXONOMY OF THE MIOSPORES AND ACRITARCHS 37 GENERAL RESULTS 37 Goose Fiord Samples 37 Escuminac Bay Samples 38 RESULTS OF THE MORPHOLOGICAL CHARAC~ER STUDY 39 Quantitative Results 39 RESULTS OF OTHER QUANTITATIVE STUDIES 46 TAXONOMY OF THE MIOSPORES AND ACRITARCHS 48 Method of Classification 48 Criteria for New Species 51 Relocation of Specimens 51 Explanation of Descriptive Terms and Headings 52 SYSTEMATICS 52 Leiotriletes 52 viii Punctatisnorites 54 Punctatasporites 58 Calamospora 58 Retusotriletes 60 Granulatisporites 66 Verrucosisporites 66 Planisporites 67 Apiculatasporites 68 Apiculatisporis 69 Dictyotriletes 70 Convolutispora 71 Perotrilites 74 Lycospora 75 Murospora 75 Stenozonotriletes 76 Lophozonotriletes 78 Cirratriradites 82 Cingulizonates 83 Emphanisporites 85 Ancyrospora 86 Endosporites 92 Rhabdosporites 97 Leiosphaeridia 100 DISCUSSION 102 MORPHOLOGICAL CHARACTERISTICS AND MIOSPORE ASSEMBLAGES 102 Valuation of Hypotheses 102 Morphological Tabulation and Devonian Plant Palaeoecology 112 Objections to Palaeoecological Interpretations 117 The Morphological Tabulatio~ Method: General Conclusions 120 MIOSPORE SPECIES AND DEVONIAN PALAEOFLORAS 121 Microfloras and Parent Floras 121 Comparison of Microfloral Assemblages 122 Goose Fiord Assemblage 122 Escuminac Bay Assemblage 125 Miospore Species and Microfloral Provinces 126 Miospore Species and Floral Migration 130 Miospore Species: General Observations 132 ix STRATIGRAPHIC POSITION OF THE ASSEMBLAGES 132 Geological Age of the Lower Okse Bay Formation 132 Geological Age of the Escuminac Formation 133 Extension of Stratigraphic Ranges 134 BOTANICAL AFFINITIES OF THE MIOSPORES 134 BIBLIOGRAPHY 137 APPENDIX I - GLOSSARY OF DESCRIPTIVE
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