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Xerox University Microfilms, Ann Arbor, M Ichigan 48106 77-2401 GARDNER, Robert Carl, 1946- THE SYSTEMATICS OF LIPOCHAETA (COMPOSITAE:HELIANTHEAE) OF THE HAWAIIAN ISLANDS. The Ohio State University, Ph.D., 1976 Botany Xerox University Microfilms,Ann Arbor, Michigan 48106 THE SYSTEMATICS OF LIPOCHAETA (COMPOSITAE:HELIANTHEAE OF THE HAWAIIAN ISLANDS DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in the Graduate School of The Ohio State University By Robert Carl Gardner, B.A., M.Sc. ***** The Ohio State University 1976 Reading Committee: Approved By: Dr. Roland L. Seymour Dr. Ronald L. Stuckey Dr. Tod F. Stuessy Adviso Dr. Thomas N. Taylor Department otany ACKNOWLEDGMENTS This dissertation could not have been completed without the assistance of numerous people. I acknowledge especially the willingness to help, the encouragement, and direction offered by Dr. Tod F. Stuessy, my major professor. He served as principal investigator under the terms of the grant funded by the National Science Foundation. I also thank Dr. Sherwin Carlquist of the Rancho Santa Ana Botanical Garden for advice concerning the logistics of the field work and for reviewing the grant proposal. Several people assisted me during the field work by acting as guides or by allowing access to their private land. Especially helpful were: Dr. John J. Fay of the Pacific Tropical Botanical Garden, Kauai; Mr. Noah Pekelo of the Hawaii Department of Natural Resources; Mr. Akah Hodges, of Molokai Ranch, Molokai; the Hawaiian Homelands Commission, Molokai; and Mr. Robert Cushnie of the Lanai Company, Lanai. Drs. Derry1 Herbst, Lyon Arboretum, and John J. Fay provided me with living and pressed material for some of the taxa I did not locate while in the field. I wish to thank the curators of the herbaria from which specimens were borrowed (A, B, BISH, F, G, GB, GH, K, MO, NY, PH, UC, US, W ) . Without the support and fieldassistance of ii my wife, P .t, and without the ideas and suggestions of my fellow graduate students this research would have been much more difficult. Appreciation is expressed to Drs. Jere N. Brunken, Roland L. Seymour, Ronald L. Stuckey, Tod F. Stuessy, and Thomas N. Taylor for their comments on and criticisms of the dissertation, and to Carol Stuessy for preparing Figure 9, Chapter I and Figures 1 and 2, Chapter III. Financial support for chemical supplies from Sigma Xi Grants-in-Aid of Research and for the field work from National Science Foundation grant GB-41175 is greatfully acknowledged. VITA March 25, 194 6 ............ Born - Tipton, Iowa 1970 ........................ B.A., University of Wyoming, Laramie 1970-1972.. ................. Graduate Teaching Assistant, Department of Botany, University of Wyoming, Laramie 1971 ........................ Sigma Xi Grants-in-Aid of Research 1972 ..................... M.Sc., University of Wyoming, Laramie 1972-1976 ................. Graduate Teaching Associate, Department of Botany, The Ohio State University, Columbus 1974 ........................ NSF - Doctoral Dissertation Research Grant 1976 ........................ Sigma Xi Grants-in-Aid of Research 1976 ........................ Outstanding Graduate Student Award, Department of Botany, The Ohio State University, Columbus 1976 ........................ G.W. Cooley Award, AIBS Annual Meeting PUBLICATIONS Gardner, R.C. 1972. Taxonomic status of Cirsium pulcherrimum and C. subniveum. Colorado-Wyoming Acad. Sci. 7:26. (Abstr.) Gardner, R.C. 1972. Biosystematic study of Cirsium (Compositae) in Wyoming. M.Sc. Thesis, University of Wyoming, 119 pp. Gardner, R.C. 197 3. Acacetin-7-0-rutinoside and pectolina- rin from Cirsium coloradense. Phytochem. 12:223. iv Gardner, R.C. 1974. Systematics of Cirsium (Compositae) in Wyoming. Madrono 22:239-265. Crawford, D.J. & R.C. Gardner. 1974. Documented chromosome numbers. 1974:2. Sida 5:292-294. Gardner, R.C. 1975. Systematics of Lipochaeta (Compositae) of the Hawaiian Islands. Amer. J. Bot. 62 (Supplement) :54. (Abstr.) Gardner, R.C. 1976. Patterns of adaptive radiation in Lipochaeta DC:(Compositae) of the Hawaiian Islands. Amer. J. Bot. 63 (Supplement):53. (Abstr.) Gardner, R.C. 1976. Compositae: Cardueae of Panama In: Woodson, et ad. Flora of Panama. Ann. Missouri Bot. Gard. (in press). Gardner, R.C. 1977. Observations on tetramerous disc florets in the Compositae. Rhodora (in press). v TABLE OF CONTENTS Page ACKNOWLEDGMENTS ......................................... ii VITA ..................................................... iv LIST OF TA B L E S ....................... ix LIST OF FIGURES ................................... x CHAPTER I. REVISION OF LIPOCHAETA (COMPOSITAE: HELIANTHEAE) OF THE HAWAIIAN ISLANDS . 1 Introduction .................................. 1 Taxonomic History ............................. 2 Generic Relationships ........................ 4 Morphological and Taxonomic Criteria . 5 Floral Features ............................... 7 Hybridization .................................. 10 Leaf Flavonoid Chemistry ................... 11 P h y l o g e n y ................................. 17 Taxonomic Treatment .......................... 19 Key to the T a x a .......................... 25 Lipochaeta Section Lipochaeta ................. 3 0 Lipochaeta succulenta (Hook. & Arn.)DC. 3 0 Lipochaeta degeneri Sherff ............ 38 Lipochaeta rockii Sherff .............. 39 Lipochaeta heterophylla A. Gray . 46 vi CHAPTER Page Lipochaeta lobata (Gaud.) DC............. 49 Lipochaeta lobata var. lobata ......... 53 Lipochaeta lobata var. hastulatoides Degener & Sherff ................ 57 Lipochaeta lobata var. leptophylla Degener & Sherff .................. 58 Lipochaeta connata (Gaud.) DC............ 58 Lipochaeta connata var. connata .... 62 Lipochaeta connata var. acris (Sherff) Gardner ........................ 64 Lipochaeta Section Aphanopappus (Endl.) Benth. & Hook........................... 67 Lipochaeta tenuifolia A. Gray ...... 67 Lipochaeta dubia Degener & Sherff . 71 Lipochaeta tenuis Degener & Sherff . 72 Lipochaeta remyi A. G r a y ................. 7 4 Lipochaeta ovata Gardner ................. 75 Lipochaeta waimeaensis St.John .... 78 Lipochaeta deltoidea St. J o h n ......... 79 Lipochaeta fauriei Leveille ............ 82 Lipochaeta micrantha (Nutt.) A. Gray . 83 Lipochaeta micrantha var. micrantha . 8 4 Lipochaeta micrantha var. exigua (Degener & Sherff) Gardner ...... 85 Lipochaeta subcordata A. Gray............ 86 Lipochaeta venosa Sherff ................. 91 vii CHAPTER Page Lipochaeta populifolia (Sherff) Gardner . 92 Lipochaeta kamolensis Degener & Sherff . 9 3 Lipochaeta bryanii Sherff ................. 94 Lipochaeta lavarum (Gaud,) DC............ 9 5 Lipochaeta kawaihoaensis St. John .... 100 Lipochaeta integrifolia (Nutt.) A. Gray . 102 Doubtful and Excluded Names ................... 108 Literature Cited ................................. 113 II. CHROMOSOME STUDIES OF LIPOCHAETA (COMPOSITAE: HELIANTHEAE) 117 Materials and Methods .......................... 118 Results and Discussion .......................... 119 Phyletic interpretations........ 119 Generic relationships ..................... 126 Maturity of the complex ................. 12 6 Taxonomic implications ................... 127 Literature Cited ................................. 12 9 III. EVOLUTION AND ADAPTIVE RADIATION IN LIPOCHAETA (COMPOSITAE) OF THE HAWAIIAN ISLANDS .... 131 O r i g i n ........................................... 134 Evolution and Adaptive Radiation .............. 13 8 P o l y p l o i d y ............................... 139 Distribution and ecology ............... 141 Modes of evolution ...................... 14 4 Flavonoid chemistry ........................ 148 Literature Cited ................................. 155 viii LIST OF TABLES TABLE CHAPTER I Page 1. Major leaf flavonoid constituents in L i p o c h a e t a ......................................... 15 CHAPTER II 1. Chromosome numbers in Lipochaeta .............. 12 0 CHAPTER III 1. Vegetationzones inhabited by Lipochaeta in the Hawaiian Islands ............, 14 5 2. Major leaf flavonoid constituents in L i p o c h a e t a .............. ........... .............. 151 ix LIST OF FIGURES FIGURE CHAPTER I Page 1. Drawing of two dimensional paper chromatograms of Lipochaeta leaf extracts ..................... 13 2. Phylogenetic relationships among Lipochaeta sect. A p h a n o p a p p u s ............................... 20 3. Phylogenetic relationships among Lipochaeta sect. L i p o c h a e t a ................................. 22 4. Distribution of L. integrifolia, L. kawai- hoaensis, and L. succulenta ....................... 33 5. Distribution of L. degeneri, L. heterophylla, and L. rockii ...................................... 4 0 6. Distribution of L. lobata var. hastulatoides, var. leptophylla, and var. lo b a t a ...................51 7. Distribution of L. connata var. acris and var. c o n n a t a ...................................... 6 0 8. Distribution of L. dubia, L. ovata, L. remyi, L. tenuifolia, and L. t e n u i s ................... 69 9. Lipochaeta ovata. A, h&bit; B, ray floret and achene; C, disc floret and achene ..........7 6 10. Distribution of L. deltoidea, L. fauriei, L. micrantha var. exigua and var. micrantha, and L. waimeaensis ............................... 8 0 x FIGURE Page 11. Distribution of L . brvanii, L. kamolensis, L. lavarum, L. populifolia, L. subcordata, and L. venosa ......................................... 8 8 CHAPTER II 1. & 2. Camera lucida drawings of meiotic chromosomes in Lipochaeta .....................................
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