BIOSYSTEMATICS AND LIFE HISTORY STRATEGIES OF SCENTLESS CHAMOMILE (Matricaria perforata Merat) IN CANADA Submitted to the Faculty of Graduate Studies and Research in Partial Fulfillment of the Requirements for the Degree of Master of Science in the Department of Crop Science and Plant Ecology University of Saskatchewan by Sheridan Lois Woo Saskatoon, Saskatchewan, Canada August 1989 The author claims copyright. Use shall not be made of the material contained herein without proper acknowledgement, as indicated on the following page. In presenting this thesis in partial fulfillment of the requirements for a Postgraduate degree from the University of Saskatchewan, I agree that the Libraries of this University may make it freely available for inspection. I further agree that permission for copying of this thesis in any manner, in whole or in part, for scholarly purposes may be granted by the professor or professors who supervised my thesis work or, in their absence, by the Head of the Department or the Dean of the College in which my thesis work was done. It is understood that any copying or publication or use of this thesis or parts thereof for financial gain shall not be allowed without my written permission. It is also understood that due recognition shall be given to me and to the University of Saskatchewan in any scholarly use which may be made of any material in my thesis. Requests for permission to copy or to make other use of material in this thesis in whole or in part should be addressed to: Head of the Department of Crop Science and Plant Ecology University of Saskatchewan Saskatoon, Saskatchewan Canada S7N OWO Abstract The objectives of this thesis were to investigate the biosystematics and life-history strategies of M~!Kl£~£l~ perforata Merat in Canada, giving emphasis to the taxon from the Prairies. Literature on the nomenclature of the Matricaria genera was surveyed and reviewed. Matricaria perforata Merat was accepted as the name for the Canadian species under study. Other M~!£l£~£l~ taxa were identified and the synonymy of these species was summarized. The taxonomic characters used to distinguish Matricaria from other genera and separate M. perforata from other species were noted. Two cytotypes of Matricaria perforata were found among 23 seed sources collected throughout Canada. Eight Atlantic populations were diploid (2n=18) and 13 populations from the Prairies and East-central Canada were tetraploid (2n=36). One population from St. John's Newfoundland had anomalous chromosome counts of 2n=27, 36 and 37. In addition to the documentation of the present distribution of M. perforata, the extent of crossability was also investigated. Greenhouse experiments were conducted on the reproductive biology of 13 populations of M. perforata. The breeding system was found to be allogamous. Achene length and achene width were good taxonomic characters for differentiating the two cytotype races; however, the number of achenes per capitulum and achene weight were not. Twenty three Canadian populations of M~!Kl£~~i~ ~erforata were grown in a common garden to examine the phenological and morphological differences. Diploid populations took a shorter period of time to bolt and f lower, and exhibi ted I ess overwintering abi I i ty than the tetraploid populations. Consequently, diploid plants were predominantly annual and the tetraploid plants were biennial and perennial in growth form. Morphologically, the tetraploid plants were larger, more erect and more spreading in growth than the diploid plants. Life cycle effects on four populations were examined by monthly planting dates in the field. Plantings from May to late-July resulted in a lower percentage of plants bolting or flowering from each population. Plantings to the field after July did not flower, whereas plantings after September did not bolt or flower during the first growing season. No plants survived from the October planting date. A morphometric study was conducted on 14 populations of M. ~~£f2£~!~ collected in Alberta, Saskatchewan and Manitoba. Multivariate analyses were performed on 32 morphological charact~rs and 13 environmental characters to determine if the sites were discriminating and if correlations could be made between the characters. Morphological differences were observed among the populations, but not correlated to the environmental characters. It was felt that these differences were not taxonomically meaningful since the main discriminating characters were vegetative and had a large amount of variability associated with them. Acknowledgements I would like to express my sincere gratitude to my advisors - Dr. Gordon Thomas and Dr. Vernon Harms, for their excellent co-supervision of my thesis. Their continued encouragement, advice and endless patience during the research and the arduous writing process is greatly appreciated. I would also like to thank Dr. Ross Ashford, Dr. Malcolm Devine and Dr. Bryan Harvey for their support as members of my thesis advisory committee. Financial support was provided by the Hooke Research Account, plus the Rene Vandeveld, L.H. Hantelman and University of Saskatchewan graduate scholarships. I am grateful for the use of the facilities and land at the Agriculture Canada, Regina Research Station, as well as the facilities at the University for the duration of this study. My appreciation is extended to the valuable technical assistance provided by Robin Wise, Faiz Ahmad, Carol Bubar, Don Harron, Cathy Harron and Bill Wallace. I would like to thank Dr. John Sheard for his advice and review of the multivariate analyses. I am indebted to the generosity of Martin Reaney for the loan of his personal computer. The never-ending rallying by Peggy, Carol, the Steve's, Manjula, Li plus many others must be thanked, and recognized as being paramount to making this thesis a worthwhile adventure! A special thanks goes to Rene for his criticisms, humour, unlimited patience and support which made the achievement of this thesis possible. i TABLE OF CONTENTS Acknow1 edgements.. i Table of contents.................................................... ii List of Tables..... .................................................. v List of FiQ\lI'es........... ........................................... vii 1. General Introduction............................................. 1 2. The Nomenclature and Ta.xonany of Matricaria perforata Merat. 5 2 .1 • Introduction. .............................................. 5 2.2. History of the nomenclature................................ 6 2.3. Nomenclature of the Matricaria species in canada. 10 2.3.1. Matricaria maritima ssp. maritima L................. 11 2.3.2. Matricaria maritima ssp. phaeocephala (Rupr.) Ra.'lJSchert. .......................................... 12 2.3.3. Matricaria perforata Merat. 13 2.4. Distinguishing Matricaria fram similar taxa................ 15 2.4.1. Matricaria versus Chamomilla............... 15 2.4.2. Matricaria versus other genera...................... 17 2.4.3. The Canadian species of Matricaria.................. 18 3. The Cytogenetics of the Chromoscme Ra.ces of M. perforata......... 23 3.1. Introduction..... .......................................... 23 3.2. Distribution of the M. perforata cytotypes................. 27 3.3. Materials and methods................................. ..... 29 ii 3.3 . 1. Chranosome CO\lIlts. ................................. 29 3.3.2. Interpopulation crosses . 32 3.3.2.1. Crossing technique . 32 3.3.2.2. Viability of hybrid seed . 34 3.4. Results . 35 3.5. Discussion . 40 4. Breeding biology in the Canadian M. perforata conplex. 47 4.1. Introduction . 47 4.2. 1'1clterials and methods . 49 4.2.1. Breeding system experiments . 49 4.2.2. Reproductive biology . 52 4.3. Results . 52 4.3.1. Breeding system of M. perforata . 52 4.3.2. Reproductive biology . 54 4.4. DisC\1Ssion. ............................•................... 56 5. Phenological and Morphological Variation in canadian M. perforata 65 5.1. Introduction . 65 5.2. ~terials aI1d methods . 69 5.2.1. Growth stages in M. perforata . 69 5.2.2. Common garden study . 70 5.2.3. Study of different planting dates....•............. 75 5.2.4. Statistical analyses . 75 5.3. Results . 75 5.3.1. Growth and development . 75 5.3.2. Common garden study (1985-1988) . 80 iii 5 . 3 . 2 . 1 . PhenolOQ'Y. ............................... 80 5 . 3 . 2 . 2 . Morpl1.olOQ'Y................ 94 5.3.3. Effects of different planting times (1986-1988).... 101 5.4. DisC'USsion....... .......................................... 113 5 . 4 . 1 . CaIDllon garden. ..................................... 113 5.4.1.1. Phenol<Jgy'................................ 113 5.4.1.2. Morphol<Jgy'.............. 119 5.4.2. Effects of different planting times................ 122 5.5. Sunrnary.................................................... 128 6. Morphological variation of M. perforata from the Prairies Provinces. ....................................................... 131 6.1. Introduction............................................... 131 6.2. to1a.terials arld methods. ..................................... 137 6.2.1. Collection sites and specimens............... 137 6.2.2. Statistical analyses............................... 147 6.3. Results. ................................................... 148 6.4 . DisC'llSsion.. ............................................... 168 7. General Sunrnary and Conclusions.... 176 References Cited.... ................................................. 183 Appendix A..........................................................