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Information to Users INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, som e thesis and dissertation copies are in typewriter face, while others may b e from any type of computer printer. Tfie quality of this reproduction is dependent upon the quality of the copy submitted. Broken or irxJistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand comer and continuing from left to right in equal sections with small overlaps. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6" x 9” black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. Bell & Howell Information and Leaming 300 North Zeeb Road, Ann Arbor, Ml 48106-1346 USA 800-521-0600 UMI NOTE TO USERS This reproduction Is the best copy available. UMI Systematic and Biogeographic Study of a Plant Species Complex; Aster Section Bucephalus by Elizabeth Anne Zamluk BSc., University of British Columbia, 1983 M Sc, Dalhousie University, 1992 A Dissertation Submitted in Partial Fulfilment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY in the Department o f Biology We accept this dissertation as conforming to the required standard Dr. Geraldine A Allen, Supervisor Dr. Patrick T. G re g ^ , Departmental (Department /of Biology) er (Department of Biology) t, Dep Dr. David Dufifus, CmMme Member (Department of Biology) (Department of Geography) Dr. Susan Aiken, External Member (Research Scientist, Vascular Plants, Research Division, Canadian Museum o f Nature) ©Elizabeth Anne Zamluk, 1999 University of Victoria All rights reserved. This dissertation may not be reproduced in whole or in part, by photocopying or other means, without the permission of the author. 11 Supervisor; Dr. Geraldine Allen ABSTRACT Aster section Eucephalus (Nutt.) Munz & Keck is comprised of about 16 taxa in North America (some rare and localized, others abundant and widespread) which appear to form a homogeneous, and probably monophyletic, group. I used a stratified sampling design to select 141 specimens from 1,669 loaned herbarium sheets. I chose morphological characters for analysis on the basis of character descriptions and the taxonomic history of Aster section Bucephalus species derived fi*om published scientific papers and floras. I calculated similarity indexes based on 33 to 36 characters, using Gower’s general similarity coefficient. Thirty-one phenetic groups were found by clustering specimens with UPGMA. The cluster memberships were adjusted by evaluating changes in the eigen values generated during discriminant analysis of “before” and “after” cluster memberships. An axis with increased value indicated that discrimination between the groups had increased relative to the axis, and a decrease showed that the groups were less separated relative to the axis. Those characters that could not be used in discriminant analyses were assessed for gaps or overlaps among groups by applying t-tests and visual inspection of box plots. Twenty-five phenetic groups remained after the iterative adjustment process. Taxonomic names were assigned to the phenetic groups based on published descriptions. Aster eastwoodiae Zaml. comb. nov. (Aster bicolor was not an available name for this taxon) reinstates a morphologically distinct taxon (Eucephalus bicolor Eastwood), previously included in A. brickellioides (Greene) Greene, that is endemic to the Klamath region of Oregon and California. Aster engelmannii Gray is divided into var. engelmannii and var. monticola Zaml. var. nov. based on size, number of phyllary rows on the involucre, and trichome characteristics. Aster wasatchensis (Jones) Blake is separated into var. wasatchensis and var. grandifolius Zaml. var. nov. based on plant size, phyllary colours, and leaf trichome characteristics. Phenetic groups were used as the bases for cladistic analyses and a hypothesis of m descent was developed from a cladogram derived by coding taxon character means as multi-state characters. Ancestral conditions were inferred from multiple outgroups including Aster turbinellus Lindel. ex. Hook. Aster wasatchensis was hypothesised to be the basal species. Locality information gathered from herbarium labels was used to produce distribution maps. Biogeographic distribution information combined with cladistic results, and an assumption of a founding taxon from Mexico (Noyes and Rieseberg (1999) hypothesised that New World asters were derived from southern taxa) suggested several biogeographical hypotheses for Aster section Eucephalus. Four lines of descent were hypothesised to give rise to I) an ancestral form in the Sierra Nevada; 2) an ancestral form in the Siskiyou Mountains of Oregon, together with Aster giaucodes Blake in the Great Basin and the Rocky Mountains; 3) a widespread group including Aster engelmannii Gray, A. vialis (Bradshaw) Blake, A. perelegans Nels. & Macbr., and A. glaucescens (Gray) Blake; and 4) A. wasatchensis (Jones) Blake in Utah. Taxa could then have developed through the processes of range expansion, isolation and vicariance. Aster wasatchensis is probably a palaeoendemic, whereas A. eastwoodiae, A. gormanii (Piper) Blake, A. vialis, A. glaucescens, and A. paucicapitatus (Robins.) Robins, are probably neoendemics. The current distribution of taxa likely reflects range modifications resulting from climatic changes caused by glaciation, and probably does not indicate the original relative positions of the taxa. Oregon and northern California form one area of species richness and Utah forms another. For these taxa, the coastal ranges exhibit more diversity and a higher rate of endemism. Rarity in Aster section Eucephalus is probably due to limited habitats and recent origin rather than any particular character trait. IV Examiners; Dr. Geraldine A Allen, SuDervis^(Department of Biology) Dr. Patri k T Gregory, D epartm ^al Member (Department of Biology) epartment of Biology) Dr. David Duffiis, Outside/M ^ber (Department of Geography) Dr. Susan Aiken, External Member (Research Scientist, Vascular Plants, Research Division, Canadian Museum of Nature) TABLE OF CONTENTS ABSTRACT............................................................................................................................. ii TABLE OF CONTENTS ....................................................................................................... v LIST OF TABLES .............................................................................................................. ix LIST OF FIGURES, ILLUSTRATIONS, AND MAPS ................................................ xi ACKNOWLEDGMENTS................................................................................................ xiv DEDICATION .......................................................................................................................xv I INTRODUCTION TO TFŒSIS.......................................................................................... 1 Modem approaches in Systematics ................................................................................. 3 Characters and character analysis ................................................................................... 9 Aster section Eucephalus (Nutt.) Munz & K e c k..........................................................13 Taxonomic history of Aster section E u cep h a lu s ....................................................... 16 Aster floral m orphology .................................................................................................17 Description of morphological characters of A ster section E ucephalus .......................18 Differences in morphological characters among species of Aster section Eucephalus reported in literature ............................................................................................ 19 II PHENETIC ANALYSIS.................................................................................................34 INTRODUCTION......................................................................................................... 34 Character choices ...................................................................................................34 Assessing ability of dendrogram to represent original data matrix using cophenetic correlations ................................................................................. 35 VI MATERIALS AND METHODS ..................................................................................35 Specimen and character choices for analysis .......................................................35 Analytical tools ..................................................................................................... 37 Leaf shape calculations .......................................................................................... 43 Analysis of characters .............................................................................................43
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