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Asteraceae) in British Columbia THE SYSTEMATICS OF INDIGENOUS SPECIES OF HIERACIUM (ASTERACEAE) IN BRITISH COLUMBIA by GERALDINE ANNE ALLEN GUPPY B.Sc, University of British Columbia, 1972 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in the Department of Botany We accept this thesis as conforming to the required standard The University of British Columbia September, 1975 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of The University of British Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 - 11 - ABSTRACT Hieracium L., a large and taxonomically complex genus in the family Asteraceae, is represented in British Columbia by seven species. The present study was undertaken in order to gain insight into the relationships between these species. Data was gathered from various sources. Patterns of morphological variation were studied, as well as the type of reproduction in each taxon and its ability to hybridize with other taxa. Chromosome number, karyotype, and meiotic behaviour were also investigated. Five of the species were examined chemically and their flavonoids identified. The species of Hieracium in British Columbia fall into two groups. The subgenus Euhieracium (a predominantly European group) contains one species, Hieracium umbel!a turn, which is triploid, apomictic and morphologically variable. The other six species belong to the subgenus Stenotheca (a mainly New World group). Within the subgenus they form three clusters of one or more species each, all of which are diploid and sexually reproducing. The clusters are morphologically and chemically distinct from one another, and differ in reproductive strategy. However, they have very similar chromosomes and are interfertile. The hybrids between species of different clusters, though they have lowered pollen viability, are all meiotically regular (indicating close relationships between the parents). Within the clusters the species intergrade morphologically and reproductively, and are chemically similar (with the exception of Hieracium triste and H_. gracile, which differ chemically). A synopsis of the seven Hieracium species is - iii - TABLE OF CONTENTS page ABSTRACT ii ACKNOWLEDGEMENTS x I. INTRODUCTION 1 II. MORPHOLOGY 5 Species of the Subgenus Euhieracium 6 The Hieracium scouleri Complex and Hieracium albiflorum . 24 (1) Variation within the Hieracium scouleri complex . 26 (2) Variation in Hieracium albiflorum 30 (3) The relationship between the Hieracium scouleri complex and Hieracium albiflorum ... 32 The Hieracium triste - Hieracium gracile Complex .... 39 III. REPRODUCTIVE BIOLOGY 47 Breeding System 48 Hybridization Experiments 56 Pollen Viability 66 IV. CYTOGENETICS 69 Methods 69 Results 76 (1) Chromosome number 76 (2) Karyotype 77 (3) Meiosis 83 - iv - page V. FLAVONOID CHEMISTRY 92 Methods 93 Results 98 VI. DISCUSSION 122 Theoretical Considerations 122 The Species of Hieracium: Biology and Interrelationships . 123 (1) The North American Hieracium umbel!atum complex . 123 (2) The Hieracium scouleri complex 128 (3) Hieracium albiflorum 133 (4) The Hieracium triste - Hieracium gracile complex . 136 (5) Relationships between sections 142 (6) The position of the genus within the Cichorieae . 143 A Synopsis of Hieracium Species in British Columbia . ... 146 LITERATURE CITED 148 - V - LIST OF TABLES page I. Morphological features of some North American species of Hieracium subgenus Euhieracium, based on published descriptions 8 II. Collection sites of the plant material used for breeding experiments 49 III. Results of experiments to determine breeding system . 54 IV. Statistical comparisons of germination rates 55 V. Morphological features of the hybrids obtained from crossing experiments 60 VI. Pollen viability 67 VII. Collection sites of plant material used for chromosome studies 71 VIII. Sources of plant material used for chemical analyses . 94 IX. Rf values and colour reactions of Hieracium flavonoids . 99 X. Distribution of the flavonoid compounds in species of Hieracium 103 - vi - LIST OF FIGURES page 1. Species of Hieracium found in British Columbia, according to Cronquist (1955a) 3 2. The Hieracium umbel latum complex: collection sites of all material examined from western North America 15 3. The Hieracium umbellatum complex: collection sites of all material examined from other regions 16 4. Leaf length-to-width ratios in the Hieracium umbellatum complex in western North America 17 5. Leaf length-to-width ratios in the Hieracium umbellatum complex in Europe, Asia and eastern North America 18 6. Morphological variation in the Hieracium umbellatum complex in western North America 20 7. Morphological variation in the Hieracium umbellatum complex in eastern North America, Europe and Asia 21 8. Numbers of flower heads in the Hieracium umbellatum complex . 25 9. Collection sites of all material examined of the Hieracium scouleri complex 27 10. Morphological variation in the Hieracium scouleri complex . 29 11. Numbers of flower heads in the species of the Hlteracium scouleri complex. 31 12. Collection sites of all material examined of Hieracium albiflorum 33 13. Morphological variation in Hieracium albiflorum 34 14. Variation in numbers of flower heads in Hieracium albiflorum . 35 15. Relative abundance of stellate hairs in Hieracium albiflorum and in the Hieracium scouleri complex 37 16. Collection sites of all material examined of Hieracium triste and Hieracium gracile 40 - vii - page 17. Morphological variation in the Hieracium triste - Hieracium gracile complex 42 18. Numbers of flower heads in the Hieracium triste - Hieracium gracile complex 45 19. Infrared spectra of apigenin 7-0-B-D-glucuronide and its y-lactone 101 20. The probable biosynthesis of the flavonoid aglycones found in Hieracium 107 21. Possible glycosylating enzymes in the different species of Hieracium 109 22. Hypothetical relationships among the western North American species of Hieracium subgenus Stenotheca 144 - vi i i - LIST OF PLATES page 1. Hieracium gracile on a mountainside southwest of Hedley, British Columbia xii 2. Pressed specimens of Hieracium cynoglossoides, H. gracile, and the-artificially produced first-generation hybrid between them 62 3. Flower heads of Hieracium scouleri, H_. albiflorum and a natural hybrid between them 64 4. A natural hybrid between Hieracium albiflorum and H_. gracile . 65 5. Karyotype of Hieracium umbellatum .... 78 6. Karyotype of Hieracium umbellatum 78 7. Karyotype of Hieracium albertinum 79 8. Karyotype of Hieracium cynoglossoides 79 9. Karyotype of Hieracium scouleri 80 10. Karyotype of Hieracium albiflorum 80 11. Karyotype of Hieracium triste 81 12. Karyotype of Hieracium gracile 81 13. Meiosis (diakinesis) in pollen mother cells of Hieracium umbellatum 85 14. Metaphase I in Hieracium umbellatum 85 15. Metaphase I in Hieracium umbellatum 85 16. Anaphase I in Hieracium umbellatum 85 17. Diakinesis in Hieracium albertinum 87 18. Metaphase I in Hieracium albertinum 87 19. Diakinesis in Hieracium cynoglossoides 87 20. Metaphase I in Hieracium cynoglossoides 87 21. Diakinesis in Hieracium scouleri 88 - ix - page 22. Metaphase I in Hieracium scouleri 88 23. Diakinesis in Hieracium albiflorum ..... 88 24. Metaphase I in Hieracium albiflorum 88 25. Metaphase I in Hieracium gracile 89 26. Anaphase I in Hieracium gracile 89 27. Diakinesis in Hieracium gracile 89 28. Metaphase I in Hieracium gracile 89 29. Diakinesis in pollen mother cells of Hieracium albiflorum X H. albertinum 91 30. Metaphase I in Hieracium albiflorum X H_. al bertinum .... 91 31. Metaphase I in Hieracium gracile X H_. scouleri 91 32. Metaphase I in Hieracium gracile X H_. albiflorum 91 33. A two-dimensional chromatogram of Hieracium umbellatum . 113 34. A two-dimensional chromatogram of Hieracium albertinum . 113 35. A two-dimensional chromatogram of Hieracium cynoglossoides . 114 36. A two-dimensional chromatogram of Hieracium scouleri .... 114 37. A two-dimensional chromatogram of Hieracium albiflorum . 115 38. A two-dimensional chromatogram of Hieracium triste .... 115 39. A two-dimensional chromatogram of Hieracium gracile .... 116 40. A two-dimensional chromatogram of Hieracium gracile X H. albiflorum 116 41. A two-dimensional chromatogram of Hieracium gracile X H_. cynoglossoides 117 42. A two-dimensional chromatogram of Hieracium albiflorum X H. scouleri 117 - x - ACKNOWLEDGEMENTS I would like to express my appreciation to my graduate adviser, Dr. Roy L. Taylor of the University of British Columbia Botanical Garden, for his help and guidance throughout the course of this research and during the preparation of the thesis. Dr. Bruce A. Bohm of the Department of Botany, University of British Columbia, provided me with laboratory space for the chemical research, and was generous with assistance and advice regarding this portion of the work. Dr. F. William Collins (now of the Department of Botany, University of Toronto) provided much useful advice and discussion concerning chemical techniques. Dr. Christopher J. Marchant (Research Scientist, University of British Columbia Botanical Garden) assisted
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