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Uhm Ms 3970 R.Pdf EXPERI:MENTAL HYBRIDIZATION BETWEEN MOUNTAIN AND COASTAL FORMS OF SIDA FALLAXWALP., AND BETWEEN S. FALLAXAND S. RHOMBIFOLIA L. (MALVACEAE) A THESIS SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAI'I IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE IN BOTANY (ECOLOGY, EVOLUTION, AND CONSERVATION BIOLOGY) MAY 2005 By Mitsuko Yorkston Thesis Committee: Curtis C. Daehler, Chairperson Gerald D. Carr Clifford W. Morden ACKNOWLEDGEMENTS I would like to thank my thesis committee members, Dr. Curt Daehler, Dr. Gerald Carr, and Dr. Clifford Morden for their guidance and support throughout the project. I am also grateful for the support and help provided for this project by my family and friends for working in the greenhouse, collecting the data, and editing my writing. 111 ABSTRACT Natural hybridization has been a significant evolutionary process in the development ofthe Hawaiian flora. In this research, I documented the outcome of experimental hybridization in Sida species in order to determine the degree ofcrossability and the morphological characteristics and fitness ofhybrids. I examined hybridization between two morphologically distinct mountain and coastal forms ofindigenous Sida fallax (Malvaceae), and between these and the alien species S. rhombifolia. This study could not confirm any hybridization between S. fallax and S. rhombifolia. In contrast, pollination experiments indicated that beach and mountain ecotypes ofS. fallax were highly crossable, and the hybrids were viable. Investigation ofmorphological variation among reciprocal hybrids between beach and mountain ecotypes ofS. fallax indicated that hybrids do not express uniformly intermediate characteristics. Out of 17 morphological traits examined in hybrids, four traits were intermediate, three traits were dominant (inherited from one parent), one trait was extreme relative to both parents. Nine traits did not statistically differ among beach, mountain and hybrid plants. The overall growth, flowering, and survival rate ofhybrids, under well-watered and drought treatments, as well as pollen stainability, suggest that the reciprocal hybrids between beach and mountain S. fallax are equally fit relative to the parental ecotype. These morphological analyses and fitness comparisons ofdivergent ecotypes ofS. fallax and their hybrids provide a baseline study on the heritability ofmorphological characters while also raising questions about the mechanisms that have maintained divergent forms ofS. fallax in the Hawaiian Islands. IV TABLE OF CONTENTS Acknowledgements iii Abstract iv Table ofContents v List ofTables x List ofFigures xii Chapter 1: Literature survey and thesis proposal.. 1 Introduction 1 The role ofhybridization in evolution 1 Definition ofhybridization 2 Hybridization and native Hawaiian plants 3 Physical and ecological background 4 Sidafallax Walp 6 Sida rhombijolia L. 10 Morphological variations within the species over the wide ecological range 12 Hybridization between divergent forms 13 Plant hybridization and invasion 14 Hybridization, restoration and conservation 17 Thesis proposal 18 Statement ofpurpose 18 Hypotheses 19 Materials and methods 21 v Seed collections 21 Experimental crosses 21 Morphology and fitness measurements 22 Fitness studies 23 Drought treatments 24 Chapter 2: Reproductive biology and hybridization ofmountain and beach Sidafallax Walp. (Malvacea) 26 Introduction 26 Materials and methods 28 Source ofstudy plants 28 Experimental pollination 28 Breeding system ofSida fallax 29 Flower bagging and self-pollination experiments 31 Artificial crossability 31 Seed (mericarp) fertility and characteristics 31 Germination velocity 32 Results 33 Flower bagging and self-pollination experiments 33 Artificial crossability 34 Seed measurements 35 Controls 35 Seed set following self-pollination 36 Crosses between beach and mountain Sidafallax 36 VI Germination velocity 36 Seeds derived from self-pollination and crossing within ecotypes ................................................................................................... 36 Seeds derived from cross pollinations ofbeach and mountain Sida fallax 38 Discussion 39 Breeding systems ofSidfallax 39 Artificial crossability, seed measurements and germination velocity 40 Hybrids between beach and mountain Sidafallax 40 Conclusions 42 Chapter 3: Morphological assessment and pollen stainability ofhybrids between mountain and coastal ecotypes ofSidafallax Walp. (Malvaceae) 43 Introduction 43 Intermediacy ofhybrid morphology 45 Materials and methods 46 Plant materials 46 Experimental pollination 48 Morphological traits 49 Digital images 53 Voucher specimens 53 Pollen stainability 53 Results 54 Morphological traits 54 Vll Digital images 57 Pollen stainability 64 Discussion 65 Morphological traits 65 Pollen stainability 70 Conclusions " 71 Chapter 4: Growth, flowering and survival rates ofhybrids between coastal forms ofSida fallax Walp. (Malvaceae) 72 Introduction 72 Materials and methods 74 Plant materials 74 Hybrids and control crosses 76 Drought treatments 77 Leafand stem growth rate, flowering rate and survival.. 77 Results 78 Leafand stem growth 78 Flowering rate 78 Survival rate 79 Discussion 84 Leafand stem growth 84 Flowering rate 84 Survival rate 86 Conclusions 86 Vill Chapter 5: Experimental hybridization between Sidafallax Walp. and Sida rhombifolia L. (Malvaceae) 88 Introduction 88 Materials and methods 89 Plant materials 89 Experimental pollination 91 Breeding system ofSida rhombifolia 92 Flower bagging and self-pollination experiments 92 Artificial crossability 93 Seed (mericarp) fertility and characteristics 93 Germination velocity 93 Results 94 Flower bagging and self-pollination experiments 94 Artificial crossability 95 Seed production and quality 96 Germination velocity , 97 Discussion 98 Conclusions 99 Chapter 6: Hypotheses revisited and conclusions 100 Literature Cited 103 IX LIST OF TABLES TABLE PAGE 1.1 Taxa with unusually complex variation patterns 13 2.1 Collection sites of original maternal plants 30 2.2 Results offlower bagging and self-pollination experiments 33 2.3 Seed production as a result ofpollination within and between beach and mountain Sida fallax ecotypes 34 2.4 Seed parameter means 35 2.5 Mean germination velocity (aV) ofSida seeds derived from various pollination treatments 37 3.1 Collection sites oforiginal maternal plants 47 3.2 Sidafallax putative cross pollination matrix 48 3.3 Traits assessed in the morphological analysis ofSidafallax 50 3.4 . Mean values for each of 17 traits of putative hybrids between beach (B) and mountain (M) forms ofSida fallax and controls grown together in a greenhouse 56 3.5 Summary ofmorphological trait expression in hybrids between beach and mountain Sida fallax 58 3.6 Mean pollen stainabilility ofSidafallax parental plants (B and M), and crosses (BxB MxM, BxM and MxB) 64 4.1 Conceptual models ofthe natural hybridization outcome 72 4.2 Collection sites oforiginal maternal plants. Rainfall data ofthese sites is from Giambelluca et al. (1996) 75 x 4.3 Sida cross pollination matrix 76 4.4 Mean flowering age (days) and flowering frequency (%) in Sidafallax hybrids under well-watered and drought treatments 82 5.1 Collection sites oforiginal maternal plants. Rainfall data ofthese sites is from Giambelluca et al. (1996) 90 5.2 Sida cross pollination matrix 91 5.3 Results offlower bagging and self-pollination experiments 95 5.4 Seed production as a result ofpollination between Sidafallax and S. rhonlbifolia 96 5.5 Seed parameter means 96 5.6 Mean germination velocity (GV) ofSida seeds derived from various pollination treatments 97 Xl LIST OF FIGURES FIGURE PAGE 1.1 The beach ecotype Sida fallax 7 1.2 The mountain ecotype Sida fallax 8 1.3 Typical leaves (A), a flower (B) and a habit (C) ofSida rhombifolia 11 2.1 Collection sites oforiginal maternal plants 30 2.2 Seed Germination (%) seeds produced by crossing between beach (B) and mountain (M) forms ofSida fallax and seeds from crosses within the two ecotypes 37 3.1 Collection sites oforiginal maternal plants 47 3.2 Leaves of selected Sida fallax from the study 51 3.3 A flower (a) and a petal (b) ofselected Sidafallax from the study 52 3.4 Typical leaves ofselected mountain (A), beach (B) and hybrid between beach and mountain forms (C) ofSidafallax 59 3.5 Habits of selected mountain (left), hybrids (center) and beach (right) Sidafallax. 60 3.6 Sidafallax seeds (mericarps) in the fruit (schizocarp) (A) and selected seeds of Sidafallax (beach, mountain and hybrid) (B) 61 3.7 Selected flowers ofSida fallax 62 3.8 A selected cultivated flower ofBlack Coral 'Ilima 69 4.1 Collection sites oforiginal maternal plants 75 4.2 Stem length (cm) of seedlings ofSida fallax hybrids under well-watered and drought treatments 80 4.3 Total leafnumber ofseedlings ofSidafallax hybrids under well-watered and drought XII treatments 81 4.4 Mean flowering frequency (%) among hybrids ofSidafallax under well-watered and drought conditions 82 4.5 Survival rate (%) ofSidafallax hybrids under well-watered and drought treatments 83 5.1. Collection sites oforiginal plants 90 Xl11 Chapter 1 Literature Survey and Thesis Proposal Introduction The role of hybridization in evolution Hybridization as an important evolutionary process has long been recognized by plant biologists (Allen 1949, Anderson ]949, Heiser ]949, Stebbins 1950, Baker 1951, Anderson and Stebbins 1954, Gillett ]972b, Heiser 1973, Stace 1975, Grant 1981, Ganders 1989, Kaneshiro 1990, Harrison 1993b, Arnold and Hodges
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