215280299.Pdf

215280299.Pdf

UNIVERSITY OF OKLAHOMA GRADUATE COLLEGE SYSTEMATIC STUDIES IN PHYSALIS (SOLANACEAE) A DISSERTATION SUBMITTED TO THE GRADUATE FACULTY in partial fulfillment of the requirements for the DOCTOR OF PHILOSOPHY oy JANET R. SULLIVAN Norman, Oklahoma 1984 SYSTEMATIC STUDIES IN PHYSALIS (SOLANACEAE) APPPOOTOD BY lu^ksa DISSERTATION COMMITTEE PREFACE This dissertation consists of four manuscripts that have been organized according to the format specified for publication in the American Journal of Botany (chapters 1 and 3), Systematic Botany (chapter 2), and the Vascular Flora of the Southeastern United States (chapter 4). The tables and figures are numbered independently in each chapter. ACKNOWLEDGEMENTS This work was supported in part by grants from the National Science Foundation (Dissertation Improvement Grant DEB-8205749 to J. R. Estes and myself), and Sigma Xi, and by funds from the George H. M. Lawrence Award from the Hunt Institute for Botanical Documentation. There are several people to be thanked for their technical assistance with this research: Drs V, Tepedino and T. Griswold of the USDA Bee Biology and Systematical Lab in Logan, Utah, and Dr. G. C. Eickwort of Cornell University provided the insect identifications; Dr. D. A. Young welcomed me into his lab and taught me the basics of flavonoid analysis; 17. Chissoe assisted with the SEM analysis of pollen; D. Brandenburg was the first to wrestle with my keys to species; and D . Hough straightened out the snags in the computer programs. I am also grateful to both D. Hudson and M. Menzel for openly sharing their knowledge of and ideas on Phvsalis. There are also several people who are much-appreciated for their non­ technical support, especially Barb Ripley, Chuck Daghlian, and Cecil & Bill Hoisington. I am especially grateful to my family, who have always provided a strong network of friendship, love, and encouragement, and to my husband's family, who have provided an extention of that network. I thank my committee members, Drs C. P. Daghlian, J. J. Skvarla, R. B. Phillips, and G. D. Schnell, for their advice and discussion during the course of this study, and for their constructive comments on these manuscripts. Dr. George Goodman provided many hours of instructive and enjoyable discussion; I appreciate his always-open door and eagerness to listen and to help. I am most grateful to Dr. James Estes for the past four years of guidance, encouragement, and friendship. I can hardly begin to measure all that I have learned under his tutelage, but hope his influence will always be reflected in my work. Finally, my husband, Dr. Tom Laue (the real "Phvsalis Phynder"!), deserves a special thanks for the many ways he has contributed to this thesis and to the overall quality of my life. TABLE OF CONTENTS List of T ables....................................................viii List of Figures...................................................... ix Pollination biology of Phvsalis viscosa var. cineraseens (Solanaceae) Abstract ........................................................ 1 Introduction......................................................2 Materials and Methods.............................................3 R e s u l t s .......................................... 6 D i s c u s s i o n ..................................................... 11 Literature Cited ............................................. 15 Tables........................................................... 18 F i g u r e s ........................................................ 19 Systematica of the Phvsalis vi scosa complex (Solanaceae) Abstract........................................................ 21 Introduction...................................... 23 Materials and Methods. ..... 26 R e s u l t s ........................................................ 29 D i s c u s s i o n ..................................................... 37 Taxonomic Treatment .......................................... 45 Literature Cited .............................. ..... 58 Tables........................................................... 60 F i g u r e s ........................................................ 55 Palynology and systematical of Phvsalis and Quincula (Solanaceae) Abstract........................................................ 77 Introduction.................................................... 78 Materials and Methods............................................ 79 R e s u l t s ........................................................80 D i s c u s s i o n .....................................................83 Literature Cited ............................................. 86 F i g u r e s ........................................................89 Phvsalis (for the Vascular Flora of the Southeastern United States) ........................................................95 Appendices. A. Color-filter photography of Phvsalis flowers............... 117 B. Insect collections. ........................... 119 C. Phenstic analysis.......................................... 120 D. Flavonoid analysis............................. .127 E. Pollen analysis vouchers................. 130 LIST OF TABLES Pollination biology of Phvsalis viscosa var. cineraseens (Solanaceae) 1. Study populations .......................................... 18 Systematica of the Phvsalis viscosa complex (Solanaceae) 1. Voucher collections used in analyses......................... 60 2. Characters used in phenetic analyses......................... 64 APPENDIX B 1. Insects collected on Phvsalis s p e c i e s ..................... 119 APPENDIX C 1. Vouchers of Phvsalis species other than the members of the P.. viscosa complex used in phenetic analyses.................. 120 2. Similarity values for the minimum spanning network . .122 APPENDIX D 1. Vouchers of Phvsalis species other than the members of the P.. viscosa complex used in flavonoid analyses.................. 127 APPENDIX E 1. Vouchers used for pollen analysis...........................130 LIST OF FIGURES Pollination biology of Phvsalis viscisa var. cineraseens (Solanaceae) 1-6. Flowers of Phvsalis species included in study ............ 19 Systematics of the Phvsalis viscosa complex (Solanaceae) 1. Phonogram for the Phvsalis viscosa complex................. 65 2. Minimum spanning network for the Phvsalis viscosa complex. 67 3-6. Representative levels of SIMGRA analysis for the Phvsalis viscosa complex............................................ 69 7-8. Representative levels of SIMGRA analysis for the Phvsalis viscosa complex............................................ 71 9. Composite chromatogram and distribution of flavonoids for the Phvsalis viscosa complex................................... 73 10. Hybridization among the members of the Phvsalis viscosa c o m p l e x .................................................. 75 Palynology and systematics of Phvsalis and Quincula (Solanaceae) 1. Size range, smaple mean, and P/E ratio for Phvsalis and Quincula pollen..................................................... 89 2-7. SEM of Phvsalis pollen................................... 91 8-11. SEM of Phvsalis and Quincula pollen.................... 93 APPENDIX A 1. Flower of Phvsalis viscosa var. cineraseens photographed with color f i l t e r s .......................................... .117 APPENDIX C 1. Phenogram for the members of the Phvsalis viscosa complex and ix representatives of other species in the genus............ 123 2. Minimum spanning network for the members of the Phvsalis viscosa complex and representatives of other species in the genus. .125 APPENDIX D 1. Flavonoid profiles for Phvsalis species other than the members of the P.. viscosa c o m p l e x .................................... 128 POLLINATION BIOLOGY OF PHYSALIS VISCOSA VAR. CINERASCENS (SOLANACEAE) Janet R. Sullivan Department of Botany and Microbiology, University of Oklahoma, Norman, OK 73019 ABSTRACT Phvsalis viscosa var. cineraseens is a self-incompatible, herbaceous perennial which occurs in disturbed or open habitats in the south- central United States and eastern Mexico. The plants are low-growing and the pendant blossoms are often obscured by the foliage. In Oklahoma the species blooms from April through October. The flowers are yellow with dark purple or brown maculations in the center and a mat of white hairs in the throat. Nectar is produced in small quantities at the base of the ovary and pollen is present in copious amounts. The pollinators are solitary bees which visit the flowers for nectar and pollen. Perdita halictoides is the primary pollinator and is oligolectic on Phvsalis. Phvsalis is an American genus of about 90 species. Generic boundaries are clearly marked by the pendant flowers and inflated fruiting calyx which encloses the berry. The corolla also exhibits an uncommon color combination for the family — the limb in all but two or three species is yellow, typically with a darker color in the center, and a mat of white hairs in the throat (Fig. 1). The pendant blossoms are often obscured by the foliage and many of the flowers hang just above the ground. In addition, the blossoms have no detectable odor. Consequently, it is difficult to envision how pollinator attraction, and hence pollination, might be accomplished. Pendant flowers are uncommon but distributed among a wide variety of families of dicots and monocots (e.g., Ervthronium. Aquilagia, Podecatneon). Host, however, have brightly colored floral parts and/or are borne on erect stems above the foliage. Specific boundaries in Phvsalis are poorly

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