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Chrysanthemoides A Taxonomic Study of Chrysanthemoides . ·Toum.r ex Medik. (Compositae) by Robert Chari Griffioen Town Cape of Submitted in Fulfilment for the Degree of Masters of Science University University of Cape To~n Botany Department December 1995 The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgementTown of the source. The thesis is to be used for private study or non- commercial research purposes only. Cape Published by the University ofof Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University Acknowledgements I thank my supervisor, Prof. H.P. Linder for direction and· helpful criticism of this thesis. Thanks are also extended to Wendy Hitchcock, Terry Trinder-Smith and Tony Verboom for their detailed comments and critical reviews of the manuscript. I am most grateful to Janet Thomas for advise with isozyme electrophoresis and Antony Hitchcock for rooting cutting material of Chrysanthemoides. My special thanks goes to Anneke Swart for helping with labelling of herbarium material and Raymond carelse for his advise on photography. The research was supported in part by the Federation for Research and Development and CSIRO, Division of Entomology, my thanks to Dr. Penny Edwards and Mr. Alan Wood. I would also like to thank the Department of Botany, University of Cape Town and their staff for the use of their facilities. Specimens were kindly loaned by BOL, NBG, NU, PRE, RUH, s, STEU and LO. Contents Page Acknowledgments Contents List of Figures and Tables Abstract 1. o. Introduction . .............................. ~ ........... 1 1.1. The Chrysanthemoides Problem .••••.•.••.•.......••• 1 1.2. Generic Delimitation of Chrysanthemoides ...•....•• 1 1. 3. Taxonomic History . ................................. 2 2.0. Materials and Methods .................................. 6 2.1. Numerical Analysis ................................ 6 2 . 1 . 1. · Proto co 1 . ....................................... 6 2. 1. 1. 1. Cluster Analysis .•.••...•........•.•••••••.••• 7 2.1.1.2. Principal Component Analysis .....•.••.•....••• 7 2.1.1.3. The C. monilifera ssp. pisifera Complex .••.••. 8 2.1.2. Justification for PCA and Cluster Analysis .••••• 8 2.2. Eco-geographical Data ••••••..•••••..•.•.....••..•. 9 2 . 2 . 1. Protoco 1 . .......· ................................ 9 2.2.1.1. Eco-geographical Data Collection ••....•.•....• 9 2.2.1.2. Geographical Data Analysis ..•...•••..••.•.••• 10 2. 2. 1. 3. Ecological Data Analysis .....••..•..•.•.•.••• 10 2. 3. Cladistics . ...................................... 11 2 . 3 . 1 . Protoco 1 . ...................................... 11 2.4. Isozyme Electrophoresis ......•••.••••.••••••••••• 12 2.4.1. Protocol ....................................... 12 2 . 4 . 1. 1. Study Sites . ................................. 12 2. 4. 1. 2. Technique . ................................... 14 2.4.1.3. Starch Preparation ••••.•.•••.••....•.•.••.•.• 14 2.4.1.4. Gel Preparation and Loading .•.•....•....••••• 15 2.4.1.5. The Setting Up of Gel and Buffer Trays ..••••. 15 2.4.1.6. Gel Scoring and Measures Used ...•.•.....••••• 16 2. 4. 2. Justification .................................. 17 2. 4. 2. 1. Available Techniques •....•........•......••.• 17 2. 4. 2. 2. Isozyme electrophoresis ....................•• 18 3 . o. Results . .............................................. 2 o 3 .1. Numerical Taxonomy ..••.•......•••....•...•..•••.. 20 3.1.1. Cluster Analysis •.••.•..•.....•....•.•.•...•••• 21 3.1.2. Principal component Analysis ................... 25 3. 2. Ecology and Distribution .•...•................... 32 3.2.1. Distribution of the Genus ...................... 32 3.2.2. Ecology ....................................... 33 3.2.3. Ecological Data Analysis Using PCA .............. 37 3.2.4. Distributional Overlap Between Taxa .... ~ ....... 43 3. 3. Cladistics . ...................................... 44 3.4. Molecular Systematics ............................ 49 3. 4 .1. Isozymic Variation ...•...•..................... 49 3.4.2. Heterozygote Deficiency .......................• 50 3.4.3. Genetic Distance and Identity .................. 57 3.4.4. Electrophoretic Analysis of Chrysanthemoides Populations from Swellendam and Infanta ............... 60 4. O. Discussion .......... " ....................... " ......... 63 4.1. Numerical Phenetics .............................. 63 4.1.1. The PCA and Cluster Analysis ................... 63 4.1.1.1. Chrysanthemoides incana ssp. incana .......... 63 4.1.1.2. C. monilifera ssp. septentrionalis and ssp. canes cens ............................................. 6 5 4.1.l.3. C. monilifera ssp. floribunda (forms 1 and 2) and ssp. rotunda ta . ... ., ............................... 65 4. 1. 1. 4. C. monilifera ssp. pisif era complex and ssp. monilifera . ........................................... 66 4.2. Phytogeography and Ecology ••.•..•................ 67 4.2.1. The PCA and Ecological Variables ............... 67 4.3. Cladistic Analysis ........................•...... 70 4.3.1. Cladistic Analysis and Bootstrap Values ........ 70 4. 4. Molecular Techniques ........................•.... 7 3 4.4.1. Genetic Identity and Geography .....•..•......•. 73 4.4.2. Genetic Identities Compared to Other Studies ... 75 4.4.3. Fit of Electrophoretic Results to Numerical Phenetics and Cladistics ...•.......................... 76 4.4.4. Population Structure ........................... 77 4.4.5. Adequacy of Isozyme Electrophoretic Results .... 79 4.5 .. Criteria for Assignment of Taxonomic Rank ..•..... 80 4.5.1. Species Concepts ............................... 80 4. 5. 2. Species . ............................... e ••••••• 84 4.5.3. Infraspecific Classification ................... 86 4. 5. 3 . 1. Subspecies . .................................. 8 7 4. 5. 3. 2. Varieties ..•................................. 8 8 4 . 5 . 3 • 3 . Forms ........................................ 9 o 5.0. Taxonomic Treatment of Chrysanthemoides ............... 91 5.1. Key to the species, subspecies and varieties of Chrysanthemoides ..................................... e 93 5.2. Numerical Key .................................... 96 5 . 2 . 1 . Methods . ...............................•...•... 9 6 5.2.2. Morphological Codes •....•••.................... 96 5.2.3. Distribution Codes ...........•................. 97 5.3. Populations Showing Phenotypic Plasticity ....... 132 References . .......................................... 13 4 Appendices . .......................................... 148 1. Characters used in PCA and Cluster Analysis •.•..•. 148 2. Characters used in Ecological PCA .........•••..... 158 3. Characters used in the Phylogenetic Analysis .....• 159 4 .1. Enzymes . ........................................ 164 4.2. Grinding Buffers ..............•................. 165 4.2.1. Tris-HCl Buffer ............................... 165 4.2.2. Tris-Malate Buffer ............................ 165 4.3. Gel and Tray Buffer Formulations ................ 166 4.4. Stock Solutions for Staining Components ......... 167 4.5. Staining Buffer Formulations •..•................ 168 4.6. Stain Recipes .............•..................... 169 5. List of Exsiccatae .........•...................... 173 List of Figures and Tables Tla Pre-Linnaean descriptions of Chrysanthemoides Tlb Abbreviations for taxon names T2 Taxa recognized in Chrysanthemoides Fl & F2 Phenograms 1 and 2 F3 & F4 PCA 1 and 2 T3 & 4 Cumulative variation percentages 1 and 2 T5 & 6 Eigenvalues 1 and 2 T7 Vegetation and habitat types T8 Soil types T9 Rainfall and altitude ranges TlO Eigenvalues for ecological data F5 PCA of ecological data F6, 7 & 8 Distribution range outlines maps 1, 2 and 3 Tll Distribution overlap proportions T12 Character weightings F9 Strict consensus tree using nelsen option FlO Successively weighted tree T13 Cladistic data matrix T14 Polymorphism, alleles and heterozygosity T15 Observed and expected genotype frequencies - Still Bay T16 Observed and expected genotype frequencies - Buff els Bay T17 Observed and expected genotype frequencies - Hoekville T18 Observed and expected genotype frequencies - Saldanha T19 Observed and expected genotype frequencies - Skoenmakerskop T20 Coefficient of heterozygote deficiency or excess T21 Genetic similarity (Nei 1978) T22 Genetic similarity (Nei 1972) T23 Overall genetic similarity Fll Localities and genetic similarity F12 Cluster analysis using genetic distance T24 Allele frequencies for three populations T25 Genetic similarity (Nei 1972) F18 Zymogram T26 Morphological and distribtional codes Mapl Distribution - C. monilifera ssp. monilifera Map2 Distribution - c. monilifera ssp. floribunda (form 1) Map3 Distribution - c. monilifera ssp. floribunda (form 2) Map4 Distribution - c. monilifera ssp. pisifera var. pisifera (form l} Map5 Distribution - C. monilifera ssp. pisifera var. pisifera (form 2} Map6 Distribution - c. monilifera ssp. pisifera var. borealis Map7 Distribution - C. monilifera ssp. pisifera var. angustifolia Maps Distribution - c. monilifera ssp. canescens Map9 Distribution - c. monilifera ssp. septentrionalis MaplO Distribution - C. monilifera ssp. rotundata Mapll Distribution - c. incana ssp. incana Mapl2 Distribution - c. incana ssp. incana Mapl3 Distribution - C. incana ssp. subcanescens F14 Chrysanthemoides drupe shape F15, 16 & 17 Three plates of leaf variation in Chrysanthemoides F18 Chrysanthemoides leaf margin types Ta Chromosome number T4.3 Gel and tray buffer formulations T4.4
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