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; A % ;:.V ‘ \ ' , | J l National Library Blblloth6oue nattonale ■ t ' o( Canada du Canada \ \ ^ O' ' \ ' Canadian Theses Service Services des theses danadiennes V' • . , ■ Ottawa, Canada \ K1A0N4 >’ ' 11 it. V \ • ' * ■ A \ A V CANADIAN THESES THESES CANADIENNES > ■ k NOTICE AVIS The quality of this microfiche is heayjly dependent upon the La qualltS de cette microfiche d6pend grandement de )S qualitd quality of the original thesis submitted for microfilming. Every de la thSse soumlse au mlcrofllmage. 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Previously copyrighted'materials (journal articles, published Les documents qui font d6jd I'objet d'un droit d'auteur (articles tests. 6tc.) are not filmed. de revue, examens publlds, etc.) ne sont pas mlcrofilmds. ^ , > s Reproduction In full or in part of this film Is governed by the La reproduction, mfime partlelle, de ce microfilm est soumlse Canadian Copyright Act, R.S.C. 1970, c. C-30. d la Lot canadienne sur le droit dliuteur, SRC 1970, c. C-30. THIS DISSERTATION lath£s e a £t£ HAS BEEN MICROFILMED MICROFILMIi E TELLE QUE EXACTLY AS RECEIVED NOUS L’AVONS REQUE Nl-339 (r.86/06) Canada Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. \ \ ' . v ' THE UNIVERSITY OF ALBERTA i A biosystematic study of Arnica subgenus Arctfca bY Stephen Roy Downie v(c. A THESIS SUBMITTED TO THE FACULTY OF GRADUATE STUDIES AND RESEARCH IN PARTIAL FULFILMENT OF THE REQUIREMENT? FOR THE DEGREE OF Doctor of Philosophy IN Plant Taxonomy Department of Botany EDMONTON. .ALBERTA Spring 1987 s s' / J , Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Permission has been granted L'autorisati'on a ete accordee to the National Library of a la Biblioth&que nationale Canada to microfilm this du Canad,a de m ic ro fllm e r thesis and to lend or sell cette these "et de preter ou copies of the film. de vehdre des exemplaires du film . The author (copyright owner) L'auteur (titulaire du droit h a s reserved’ other d'auteur) se reserve les publication rights, autres droits de publication; neither the thesis fnor ni la these ni de longs extensive extracts fronji it extraits de celle-ci ne may be printed or oth^j'wise d o iv e n t etre imprimes ou reproduced without his/her autrement reproduits sans son written permission, f aqtorisation ecrite. ■ \ \ ■ ISB10-315/37617-1 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. THE UNIVERSITY OF ALBERTA RELEASE FORM NAME OF AUTHOR Stephen Roy Downie • ‘ \ ■ o ’’ TITLE OF THESIS ' A biosystematic study of Arnica subgepus Arct/ca ' DEGREE FOR WHICH THESIS W AS PRESENTED Doctor of Philosophy > * YEAR THIS DEGREE GRANTED Spring 1987 . - . - i Permission is hereby granted to THE UNIVERSITY OF ALBERTA LIBRARY to ' 1 ■ reproduce single copies of ttys thesis and fo lend or sell such copies for private, scholarly or scientific research purposes only. The author reserves other publication .rights, and neither the thesis nor ' o extensive extracts from it may be printed or otherwise reproduced without the author’s written permission. ^ (SIGNED) !?/:.Ty PERMANENT ADDRESS: o *o f c r / c > .......................... , DATED . Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. THE UNIVERSITY OF ALBERTA vV ■.V FACULTY OF GRADUATE STUDIES AND RESEARCH i\ The undersigned certify that they have read, and recommend to the 1 i'll i'. Faculty of Graduate Studies and Research, for acceptance, a thesis entitled A biosystematic study of Arnica subgenus Arctica submitted Stephen Roy Downie in • • ' ' • . ' ' , ■' V' "* ■ partial fulfilment of the reqyirements for the degree of Doctor of,Philosophy in Plant ^ Supervisor External Examiner Date Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 'S Dedication To my parents, Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ' Abstract. ’ • ■■ ■ ■ ' “ ■ Evaluation of systematic relationships of Arnica subgerrUs Arctica show that the ■ ’ ' '• ' . “ . v' ■ . complex, as circumscribed in the present study, consists of seven species. No new taxa have been proposed; however/names of seven taxa previously recognized are treated ' 1 ' /■ as synonyms. Arnica angustifotia, A. frigida and A ionchophyi{a, each consisting of tw o subspecies, are polymorphic and varied in ploidy level and fpHar flav.onoid chemistry. Cluster (TAXMAP and UPGMA) and principal component analyses of A angustifot ia (s.f.) revealed this circumpolar aggregate to be best represented as two subspecies: A. angustifofia subsp. angustifotia (a combination! of the previously, _ ■ recognized subspecies angustifot ia, attenuata, sornborgeri, intermedia; iijin ii, aipina and A. piantaginea ); and A. angustifoiia subsp. tomentosa. Subspecies iouiseana and ; frig id a o f A. iouiseana are now ranked as species, and a new combination, A. frig id a subsp. griscom ii, is pffesented. The subspecies name chionopappa of A. fonchophylia is synonymized with the latter; A. ionchophyiia suhsn~Jrfnogiossa is maintained. Studies oh - . • 7 , . ' '• "• ' :■ the reproductive behaviour of A: fuigens ar^JiJ sororia revealed them to be amphimictic and ee|f-incompatible, with^rhfjfcial hybridization experiments being , unsuccessful. Both are maintained as separate species. .. ■ ■ » : / ' ■ ‘ ' . ■ r ■ . ■ ■ V ’ ' ■■ The basic chromosome ,number fo r the subgenus is x= 19. with A. angustifofia and A frig id a 2n=38, 57, 76 and 95, A. fohchophytia 2 n = 3 8 ,5 7 and 76, A Iouiseana 2 n -7 6 arid 95, A .r>rydjbergii'% n=38 and 76; and both A. fufgens and A sororia 2n=38. Twelve flavonoids 'w4re‘isolated, and eleven identifiedrfrom Arn /ca subgenuS Aifcf/ca.' Simple mono-glycosides o f quercetin and kaempf erol were ubiquitous* thus indicative of ' a somewhat primitive biochemical profile. Flavonoid profiles are relatively simple,-With .tw o tb s ik compounds identifiedpiec population. FlavonOid diversity appearsto have '•*< ' v ' ; > * ' ' ■*./..' ; y ; f- !■" ; : ■ -y,, .> - i.- ■;.1 . l y , acbornpanied high morphological variability.: Amphimictic species o f subgenus Arctica are i ' ■ ■ V -'S. ! • 1i ; closely correlated withpon-glaciatqd regions. Disjunct distributions are probably the result o f survival in refugie during tfie fate .Wiscohsinan, with apomictic phases being] Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. responsible for the recologization of previously glaciated aj^as. Phylogenetic relations amongst all taxa, using structural and flav^>noid syhapomorphies, revealed A. angustifolia to be the progenitor of all taxa within the subgenus. Arnica subgenus Arctica probably represents the most ancestral group, giving rise to ail other Arniba •' ' : ■■ ■ ' ■ ’ • ■ • : ; ' > .■ subgenera. •. , • i;.‘ _ A taxonomic revision of the subgenus includes descriptions, keys, synonymies, * . i1 , ■ -\ 1 ' ' f distribution maps, and listings of representative specimens.', " V. # t * > . ' v--: ■■ .y ■ ■ t> * . ■ . ■ - > * ■ ■ , : ■" v • ',‘1 , ' v #.. ■ . ■ y / . ^ : >■ ■ .y ,, ^ *■' • ’ I"- Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Acknowledgments , • , . s ** ■ ' ’ , \ My appreciation and thanks first extends to Dr. Keith E. Denford, my thesis advisor, who for the last four years had contributed significantly to this thesis. From , , ■ suggesting the project ahd providing the facilities arid generous financial assistance to , carry it but, to being a never ending source of stimulating ideas and moral support, his irtput is greatly acknowledged. ) vyould also like to thank Dr, Guy Brassard for acting as my external examiner and, yes, I will never again underestimate the importance of french accentuation, Dr. Mark R'T. Dale, on many occasions; provided critical reviews of papers and helpful suggestions on the preparation of this thesis. His iriterest“in''this project, and his home brew, will always be appreciated. I would also like tg. thank the remainder of my '■ ' ' ' ’ ’ . , . ■ A ' \ : A ' *■' ■ ; dissertation committee. Dr. George E. Ball and Dr. Dale H. Vitt, for their valuable A \ •. ■> ' ’ editorial criticisims. ’ \ ‘ ■■ ■ ■ \ • , ' : A, ■ ■■■• ,;A„ ,. ;■ ’ . A'number of people provided assistance1 and companionship both in the field and .the laboratory. IN^y appreciation goes to: Jim Denford, Brian Husband, William Gruezo, ' • )' I * A ' ■' -> , ’ ‘ , , f'C h r i s Pisko^Dubienski, Zuzu Gadallah land heF laugh), Agnes Vanende,'Barry Storeshaw, Ann Smreciu and Brfj Kbhli. Appreciation also (extends'to the girls in the office who ; ' •' . ,. ' A' (' ■ ■ ’. , ' , ' A incessantly’put up with me and my demands; and the Department o f Botany