Relationships among fragrance, phylogeny and pollination in southwestern Nyctaginaceae Item Type text; Dissertation-Reproduction (electronic) Authors Levin, Rachel Ann Publisher The University of Arizona. Rights Copyright © is held by the author. Digital access to this material is made possible by the University Libraries, University of Arizona. Further transmission, reproduction or presentation (such as public display or performance) of protected items is prohibited except with permission of the author. Download date 30/09/2021 20:23:11 Link to Item http://hdl.handle.net/10150/280128 INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter fece, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. 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Bell & Howell Information and Learning 300 North Zeeb Road, Ann Arbor, Ml 48106-1346 USA 800-521-0600 RELATIONSHIPS AMONG FRAGRANCE, PHYLOGENY AND POLLINATION IN SOUTHWESTERN NYCTAGINACEAE by Rachel Ann Levin Copyright © Rachel Ann Levin 2001 A Dissertation Submitted to the Faculty of the DEPARTMENT OF ECOLOGY AND EVOLUTIONARY BIOLOGY In Partial Fulfillment of the Requirements For the Degree of DOCTOR OF PHILOSOPHY In the Graduate College THE UNIVERSITY OF ARIZONA 200 1 UMI Number: 3002512 Copyright 2001 by Levin, Rachel Ann All rights reserved. __ (ft UMI UMI Microform 3002512 Copyright 2001 by Bell & Howell Information and Leaming Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. Bell & Howell Information and Leaming Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, Ml 48106-1346 2 THE UNIVERSITY OF ARIZONA « GRADUATE COLLEGE As members of the Final Examination Committee, we certify that ve have read the dissertation prepared by Levin entitled ilelationships among Fragrance. Phvlnppnv anri Pnii inatinn in Southwestern Nyctaginaceae and recommend that it be accepted as fulfilling the dissertation requirement for the Degree of Doctor of Pliilosophy /! yi4c 00 LUtlinda A. ..IcUadi D^e U. I^JkcLuxj^ /'^ Cd Rob^^t if.) Robichaux Date Robert A./Raguso Date Q/diOp, /. -BKmrnur^ IS her en .^^ith L. ISronstein Date Date Final approval and acceptance of this dissertation is contingent upon the candidate's submission of the final copy of the dissertation to the Graduate College. I hereby certify that I have read this dissertation prepared under my direction and recommend that it be accepted as fulfilling the dissertation requirement. Dissertation Director Date Lucinda A. :tcDade 3 STATEMENT BY AUTHOR This dissertation has been submitted in partial fulfillment of requirements for an advanced degree at The University of Arizona and is deposited in the University Library to be made available to borrowers under rules of the Library. Brief quotations from this dissertation are allowable without special permission, provided that accurate acknowledgment of source is made. Requests for permission for extended quotation from or reproduction of this manuscript in whole or in part may be granted by the copyright holder. SIGNED: 4 ACKNOWLEDGMENTS This dissertation would not have been possible without the collaboration and assistance of many people. I would especially like to thank my advisor, Lucinda McDade, for encouraging me to join her and Rob Raguso in collaborating on a project studying relationships among fragrance, phylogeny, and pollinators. Further, I greatly appreciate Lucinda's amazing support, her love of systematics, and her considerable help in preparing manuscripts for publication. I cannot thank Rob enough for his enthusiasm and for teaching me how to collect and analyze fragrance and injecting numerous samples into the GC-MS for me. Thanks also to the other members of my committee, Drs. Judie Bronstein and Rob Robichaux, for their helpful comments and criticisms. The fragrance collections from field locations that were often at least a day's drive from Tucson (and often much farther) depended on the kind assistance of Amy Boyd. Thomas Tully, Mariette Manktelow, and David Heam; I am very grateful for their help, photographic expertise, and companionship on these long field trips. I would also like to thank Alice Le Due and Mark Fishbein, who provided much needed seeds of species that I was unable to visit in the field. A hearty thank you to both present and past graduate students of the University of Arizona Department of Ecology and Evolutionary Biology for discussions, support, relaxation, and friendship. I want to especially thank Jill Miller, Juliann Aukema, Henar Alonso-Pimentel, Maria Clauss, Margrit Mcintosh. Mark Fishbein, Betsy Arnold. Margaret Evans, Amy Boyd, Ken Moriuchi, Wendy Goodfriend, Francisco Omelas, Sandy Adondakis, Susan Masta, and Amy Faivre. A number of undergraduates associated with the McDade lab were also very helpful: these include Katy Riley. Alexandra Chetochine, Amanda Fox, and Alissa Martin. I want to thank the staff at Big Bend National Park, TX and the Jornada LTER. NM for permission to collect fragrance and plant materials, and their assistance in locating species within these areas. For help in locating Mirabilis macfarlanei. I thank the staff at the Bureau of Land Management in Idaho, and Joe Duft for allowing fragrance collection of this species from a plant growing in his personal garden. Many people have helped me locate plants in the field; I would especially like to thank Richard Spellenberg and Jackie Poole. I am grateful to Mats Thulin for providing me with leaf material of Selinocarpus sontalensis; I was able to extract DNA from this material and, thus, include this species in my phylogenetic analysis of the genus. I am very thankful for the support of my parents, Ellen and Joe Levin, my sister. Sara Levin, and my invaluable friends, Jill Miller, Henar Alonso-Pimentel. and Juliann Aukema; all of these people were indispensable in the completion of this dissertation. A research assistantship from a National Science Foundation grant to Lucinda McDade and Rob Raguso provided me with the flexibility of time and sufficient resources to conduct my dissertation research. I was also supported by the American Society of Plant Taxonomists, Sigma Xi, the Department of Ecology and Evolutionary Biology at the University of Arizona, and the University of Arizona Research Training Group in the Analysis of Biological Diversification. 5 TABLE OF CONTENTS ABSTRACT 6 CHAPTER 1: INTRODUCTION 8 Frcifirance 9 Hawkmoth Pollinators 10 Phylogenetic History 11 Evaluating Ecological and Evolutionary Effects on Fragrance 13 This Study 13 Explanation of Dissertation Format 13 CHAPTER 2: PRESENT STUDY 16 APPENDIX A: PHYLOGENETIC RELATIONSHIPS WITHIN NYCTAGINACEAE TRIBE NYCTAGINEAE: EVIDENCE FROM NUCLEAR AND CHLOROPLAST GENOMES 20 APPENDIX B: TAXONOMIC STATUS OF ACLEISANTHES, SELINOCARPUS, AND AMMOCODON (NYCTAGINACEAE) 36 APPENDIX C: FRAGRANCE CHEMISTRY AND POLLINATOR AFRNITIES IN NYCTAGINACEAE 51 APPENDIX D: RELATIONSHIPS AMONG FRAGRANCE, PHYLOGENY, AND POLLINATION IN THREE GENERA OF NYCTAGINACEAE 87 REFERENCES 151 6 ABSTRACT Fragrances appear to act in conjunction with visual cues to attract specific pollinators. Besides the ecological influence of pollinator type on fragrance, as with many other attributes of organisms, phylogenetic history may also affect fragrance composition. In this dissertation I examine the quality and quantity of floral and vegetative fragrance, and explore the relationships among fragrance, pollinators, and phylogenetic history in the plant family Nyctaginaceae. Using DNA sequence data, I inferred phylogenetic relationships among and within the Nyctaginaceae genera. Adeisanthes, Selinocarpus, and Mirabilis. There is a high incidence of hawkmoth pollination within these genera, in addition to multiple pollinator transitions. Results suggest that neither Acleisanthes nor Selinocarpus are monophyletic, but that together they comprise a monophyletic lineage. Because of this finding, I have taxonomically combined these two genera into a single genus. Analyses of floral and vegetative fragrance from Acleisanthes. Selinocarpus, and Mirabilis species included in the phylogenetic study show that each species has a unique fragrance profile. Further, although there is substantial variation among individuals within species, intraspecific variation is significantly lower than interspecific variation in fragrance profiles. Fragrances are composed of 5-108 different compounds from at least seven different