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Relationships of South-East Australian Species of Senecio (Compositae

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Relationships of South-East Australian Species of Senecio (Compositae eq lq ðL RELATIONSHIPS OF SOUTH-EAST AUSTR.ALTAN SPECIES OF SENECIo(CoMPoSITAE)DEDUCEDFRoMSTUDIESoF MORPHOLOGY, REPRODUCTIVE BIOLOGY AND CYTOGENETICS by Margaret Elizabeth Lawrence, B.Sc. (Hons') Department of Botany, University of Adelaide ii Thesis submitted for the Degree of Doctor of Philosophy, University of Adelaide Mayr 1981 l \t l ûr¿a.,"lto( ? 4t ' "'-' 'l-" "l DECLARATION This thesis conÈains no material which has been accepted for the award of any other degree or diploma in any University. To the best of my knowledge and belief this thesis contains no material previously published or written by another persont except where due reference is rnade in the text' Margaret E. Lawrence ACKNOWTEDGEMENTS I wish to thank Dr. D. c. christophel for supervising this project which was completed. in the laboratories of the Department of Botany, university of Adelaide. I am especially grateful to professor P. G. Martin for his continuing advice and helpful criticism of the cytogenetic section of the work. Thanks are also due to Mr. A. Floyd, FOreStry cOrnmission, New South Wales, for collecting material of Senecig amygdalifolius; Dr. Th. Muller, NatiOnal llerbarium, Rhodesiarfor providing seed of S. discifolius Professor B. L. Turner, University of Texast Austin, fot advice concerning cytological techniques and Dr' R' O' Belcher, Ypsilanti, l4ichígan, for communicating techniques for cross-pollination experiments. l4y gratitude is expressed to the National Parks and wildlife services of south Australia, Victoria and New South VIaIes for providing collecting permits, and to the park rangers for their assistance and. direction. I am also grateful to the directors and curators at the Adetaide (AD) I Sydney (SYD) and Melbourne (MEt) Herbaria for enabling me to examine collections. My thanks to colleagues in t,he Department for many st,imulating discussions and for assistance with a variety of techniques' special thanks are due to Dr. D. T. Blackburn for help with computing problems and to Ms. J. Gibbs-clema for typing the manuscript. I am also extremely grateful to Dr. O. Zarucki, Mr. and Mrs. J. W. Lawrence and many others unnamed here, who have helped in a varietY of waYs. The research was financed by a commonwealth Postgraduate Research Award., which f wish to acknowledge. TABLE OF CONTENTS Volume I Summary I CHAPTER 1 General Introduction 6 1.1 Evolutionary processes and very large genera 6 L.2 ListinÇ of species and geographic range of study 9 1.3 Taxonomic history of Senecioneae and Senecio I4 1.3.1 Senecioneae t5 1. 3.I.1 General treatments 15 1.3.1.2 Genera of Senecioneae occurring in Australia I8 1.3.2 Senecio I8 1.3.2.1 General treatments 18 L.3.2.2 Species of Senecio occurring in Australia 19 CHAPTER 2 l4orphological variation and Descriptive Terminolögy 24 2.I Introduction 26 2.2 lvlaterials and methods 27 2. 3 Vegetative characteristics 28 2.4 The inflorescence 28 2.5 The involucre and recePtacle 29 2.6 Distribution of florets within capitula 31 2.7 The corolla 32 2.7 .L Shape 32 2.7 .2 Venation 32 2.7 .3 Pubescence 32 2.7.4 Ligule colour and ePidermis 33 2.8 The pappus 33 2.9 The androecium 34 2.9.I Filament collar 35 2.9.2 Ant.her base 35 2.9.3 Endothecal tissue 36 2.9.4 Apical appendage 36 2.L0 Gynoecium 37 2.10.I Style branches 37 2.L0.2 Style base and nectarY 37 2.L0.3 Ovary wall- crystals 3B 2.IL The achene 39 2.11.1 Pubescence and mucilaginous properties 39 2.Ll-.2 Surface sculPturing 39 2.LJ-.3 Polymorphisms 40 2.L2 "Senecioid" and "cacalioid" characteristícs 4L CHAPTER 3 Morphology and Systematics 43 3.1 Introduction 44 3.2 Senecioneae 45 3.2.I General description of tribe 45 3.2.2 Key to genera in Australia 48 3.3 Australian species of Senecio 5I 3.3.1 General dsecriPtign 51 3.3.2 Key to aII species 53 3.4 Descriptions and discussion of systematics of Senecio species 65 3.4.L Radiate species 66 3.4.2 Discoid species 116 3.4.3 Erechthitoid species 131 3.4.4 Exotic species 165 3.4.5 Sterile hybrids wrongly given taxonomic status L75 3.5 Species from other genera L77 3. 5.1 Arrhenechtites mixta r77 3. 5. 2 Erechtites'valerianaefolia 181 3 .5.3 Bedfordia salicina r83 3.6 Relatíonships deduced from morphological evidence 186 186 3 . 6. 1 Primit.ive character states 3.6.2 Groups of sPecies 189 3.6.3 Degree of evolutionary advancement 193 3. 6. 4 Phylogenetic sPeculations 198 3.7 Index to species descriptions and líst of species not included 203 TABLE OF CONTENTS Volume 2 CHAPTER 4 ReProductive BiologY 205 4.I Introduction 206 4.2 Materials and methods 208 4.2.I Glasshouse trials 209 4.2.2 Pollen-ovule ratios 2LL 4.2.3 Seed size and number 2II 4.2.4 Seedling establishment 2L2 4.2.5 Longevity 2L2 4.3 Results and observations 2I2 4.3.1 Direct and indirect evidence of breeding systems 2L2 4.3.2 Observations of floral biology 2L8 4.3.3 Pollen vectors 2L9 4.4 Discussion 220 4.4.1 lr{ode of reproduction 220 4.4.2 Breeding systems 220 4.4.3 Breeding systems and generation length 22L 4.4.4 Seed síze and number 223 4.4.-5 Dispersal potential 225 4.4. 6 Seedling establishment 226 4.4.7 Combinations of reproductive traits: r- and K-selection 228 4.5 Conclusions 235 CHAPTER 5 Recombination in Senecio 237 5.1 Introduction 238 5.2 Materials and method 240 5.3 Results and discussion 24L 5. 3. 1 Chromosome numbers 24r 5.3.1.1 Ploidy distributions in Senecio 24] 5.3.1.2 Polyploidy and recombination 249 5.3.1.3 Polyploidy and speciation 25r 5.3.2 Effects of chiasma frequency and position 253 5.3.3 Effects of breeding sYstems 255 5.3.4 Effects of generation lengths 257 5.3.5 Pair-wise associations of regulatory factors 258 5.3.5.1 Breeding system and generation length 259 5.3.5.2 Breeding system and chromosome number 26I 5.3.5.3 Breeding system and chiasma frequency 26L 5.3.5.4 Generation length and chrOmosome number 264 5.3.5.5 Generation length and chiasma frequency 264 5.3.5.6 Chiasma frequency and chromosome number the "Recombination Index" 265 5.3.6 Recombination systems in Senecio 267 5.4 Conclusions 272 CHAPTER 6 Nuclear DNA Amounts 277 6.1 Introduction 278 6.2 Maberials and methods 28O 6.2.I Source of material 280 6.2.2 Feulgen stain and SOZ water 28]. 6.2.3 Cultivation 28r 6.2.4 Preparation of slides 28r 6.2.5 Measurement of DNA amount 283 6.2.6 Analysis of results 284 6.2.7 Hydrolysis'times and root tip sizes 284 6.2.8 Selection of a calibration standard 286 6.2 .9 CeIl volumes 287 6.3 Results and discussion 287 6.3.1 Terminology 287 6.3.2 Compar,ison of calibration standards 288 6.3.3 Reliabitity of DNA estimates and size of s ignif icant dif f erences 290 6.3.4 Interspecific differences in DNA amounts 292 6.3.5 Intraspecific differences in DNA amounts 296 6.3. 6 Nucleotypic effects 298 6.3.6.1 Size of structures 299 6.3.6.2 Ce1l cycle times 3 00 6. 3 . 6. 3 Minimum generation times 302 6.3.7 Nature of changes in DNA amounts 304 6.3.8 Speculations on the direction of changes in DNA amounts 3 06 6.4 Conclusions 3r2 CHAPTER 7 Karyotypes 314 7 .L Introduct.ion 315 7 .2 MaLerials and methods 316 7 .2.L Karyotype construction 316 7 .2.2 Interspecific comparisons 318 7.3 Results and discussion 325 7.3.1 Illustration of karyotyPes 325 7.3.2 Comparison of karyotypes 326 7.3.2.L Size of significant diff erences 326 7.3.2.2 Interpretation of data 326 7 .3.2.3 Relationships deduced from percentage similarities 327 7 .3.2.4 Satellite chromosomes 331 7 -3-3 Karyotype symmetry and' evolutionary advancement 334 7.3.4 Changes in absolute chromosome si-ze 339 7.3.5 The basic chromosome number of Seriecio 34 0 7.3.5.1 Karyotype symmetry and absolute chromosome size 342 7.3.5.2 The number of satellite chromosomes 343 7.4 ConclusÍons 345 CHAPTER B Natural and Synthetíc Hybrids 347 8.1 Introduction 348 8.2 Materials and methods 349 8.3 Results and discussion 350 8.3.1 Natural hybrids 350 8.3.1.1 Characteristics of hybrid' and parent plants 350 8.3.I.2 Evidence used to determine parent species 361 8.3.1.3 Pollen and seed development 363 8. 3. 2 Crossing programs 365 8.3.2. I Program I 367 8.3.2.2 Program 2 370 8. 3. 3 Extended studies of S. pt'erophorus x S. hypoleucus 370 8.3.3.1 Frequency of natural hybridization 37L 8.3.3.2 Likelihood of fertile hybrid formation 373 8.3.3.3 Evidence of additive gene effects 374 8.3.4 The rayed gene complex in Senecio 375 8.3.5 origins of decaploid species 377 8.3.6 Hybridization and polyploidy in Senecio and Senecioneae 379 CHAPTER 9 General Conclusions 383 9.1 Systematics of Senecioneae in Australia 384 9.1.I The generic status of species examined 384 9.L.2 Subdivisions of Australian spec ies of Senecio 387 9.2 The application of current evolutionary theories 38 9 9.2.L r- and K-selection 38 9 9.2.2 Recombination sYstems 390 9.2.3 The C-value Paradox 392 9.2.
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