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CYTOTAXONO?C STUDIES in the GENUS TULBAGHIA by ROBERT

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CYTOTAXONO?C STUDIES in the GENUS TULBAGHIA by ROBERT CYTOTAXONO?C STUDIES IN THE GENUS TULBAGHIA by ROBERT BRIN SLEY BIJRBI DGE Thesis presented for the Degree of Doctor of Philosophy at the University of Edinburgh in the Faculty of Science. October 1970. (0 ACKNOWLEDGEMENTS I wish to acknowledge with gratitude the financial assistance accorded to me by the Science Research Council., without which this work would have been impossible. I am particularly indebted to my supervisors Dr P.H,Dafls and Dr A.P.Dyer for continued support, encouragement and valuable discuesics: and also to Mr B..L.Burtt and Dr J.A.Ratter for advice in times of need. I wish also to thank the Regina Keeper and staff of the Royal Botanic Garden, Edinburgh for the use of herbarium, library and garden facilities. The taxonomic revision of Tulbaghia is based on material from the following herbaria: British Museum (Natural History), London (BM); Royal Botanic Garden, Edinburgh (E); Royal Botanic Gardens, Kew (K); Botanical Museum, Lund (Li)); Botanisohe Staatssammlung, tMnchen (tO; 3otany Department, University of Natal (VU); Botanical Research institute, National Herbarium, Pretoria (PRE); School of Botany, Dublin (TaD); Institute of Systematic Botany, University of Uppsala (UPS). I wish to express my gratitude to the Directors and staff of these institutions for their co-operation and help. -i- CONTENTS INTRODUCTION AND OBJECTIVES 1 a. Introduction 2 be Objectives 3 c. A definition of Tu].baghia and a short account of related genera 4 NUMERICAL STUDIES ON PRESUMED RELATED GENERA 9 a. Introduction 10 b. Methods and choice of characters 10 c Results 15 Discussion of method - 19 Discussion of results 22 VESSELS IN TULBAGHIA AND RELATED GENERA 32 THE a. Introduction 33 Materials and methods 33 Results and discussion 34 A DISCUSSION OF THE DISTRIBUTION OF THE GENERA RELATED TO TULBAGHIA -10 A SUMMARY OF SECTIONS 1-4 51 THE MORPHOI:OGY AND ANATOMY OF TULBAGHIA SPECIES 52 A. Introduction 53 a. Roots 54 b, Rootstock 54 c. Leaves - 55 1. Scape 56 -11- Involucral bracts 57 Inflorescence 57 Pedicels 57 Perianth 58 Corona J. Anthers 67 k. Gynaeciuxn 67 1. Capsule 68 68 M, Seed Embryology 68 Pollen 68 70 7. STUDIES IN THE KARYOTYPE Materials and methods 71 Pre-treatments 72 Fixation 72 Hydrolysis and staining 74 Heterochromatic segments 76 Results 76 Discussion 89 8. CHIASMA LOCALISATION 98 Introduction 99 Materials and methods 99 c • Results 100 9. DISCUSSION : THE GENETIC AND BREEDING SYSTEM • IN TULBAGHIA 103 -iii- 10. THE TAXONOMY OF TULBAGHIA 109 Introduction 110 Early nomenclatural problems in Tulbaghia 111 Species limits in Tulbaghia 114 d, A note on the illustrations accompanying the species descriptions 117 e. Synopsis of classification 120 1. Key to species 122 Generic and specific descriptions 125 Species of uncertain affinities 207 Species insufficiently known 208 CONCLUSIONS 209 BIBLIOGRAPHY INDEX -iv.- LIST OF ILLUSTRATIONS PLATES. 1,. Vessels of Tulbaghia violacea Somatic chromosomes of T. ludwigiana S. S - Somatic chromosomes of T. dieterlenil S. FIGURES. Miersia and Gilliesia floral dissections 24 Vessels from root metaxylem of Tulbaghia and related genera 35 Diagram showing measurements of perianth and corona • 59 Camera lucida drawings of serial sections through a flower bud of Tulbaghia n. sp. 1 63-66 Karyotype of T. cápensis 78 Karyotype of T. alliacea 78 Karyotype of T. n. s. 1 79 Karyotype of T. ñatalenais 80 & 10. Karyotype of T. ludwigiana 80 Karyotype of T. n. sp. 2 81 Karyotype of T. leucantha 81-62 Karyotype of T. dieterlenii 82 Karyotype of T. acutiloba 83 Karyotype of T. fragrans 84 Karyotype of T. simmleri 85 17 Karyotype of T. violacea var. violacea 86 18.-20. Karyotype of T. violacea var. obtusa 86-87 21. Nucleo&ar organiser chromosome'types in Tulbaghia 91 22. T. capensis 130 23. T. alliacea 135 24. T. sp. nov. 1 138 25. T. ludwigiana 141 26. T. campanulata 143 27. T. dregeana 147 28. T. natalensia iso 29. T. friesii 153 30. T. en. nov. 2 156 31. T. aeou.inoctalis 160 32. T. tenuior 163 33. T. calcarea 165 34. T. leucantha 171 35. & 36. Scatter diagrams (of T. leucantha) 172 37. & 38. T. cameronii 176-177 T. dieterlenhi 182 T. acutilàba 185 T. fragrans 189 T. rhodesica 192 T. si1eri 197 T. galpinii 199 T. violacea 205 -vi- MAPS. 1. Total distribution of the genus Tulbaghia in southern Africa 40 Distribution of the subfamily Allioieae . 43 Gondwanaland, late Jurassic. After King (1967) 48 Southern Africa - political boundaries 119 Distribution of T. capensis 131 Distribution of T. alliacea; T. n. sp. 1; T. ludwigiana & T. campanu.lata 144 Distribution of T. dregeana; P. natalensis; T. frtesii & P. sp. nov. 2 157 6. Distribution of T. aequinoctalis; T. tenuior & T. calcarea 166 Distribution of "forms" of T. cameronli 176 Distribution of T. leucantha; aff. T. leucantha & T. cameronii 179 Distribution of T. dieterlenii & T. acutiloba 186 Distribution of T. fragrans & T. rhodesica 193 Distribution of T. sinunleri; P. galpinii & P. violacea 206 -1- 1. INTRODUCTION and OBJECTIVES. -2- la. INTRODUCTION. The genus Thlbaghia Linn., a native of southern Africa, occupies a somewhat intermediate position between the Liliaceae and Arnaryliidaceae as classically constituted. A email number of clones of Tulbaghia species have been cultiva- ted in Britain since the earlier half of the nineteenth century, but only in the last decade have more than two or three species been available at any one place. Tulba,ghia species possess some of the largest chromosomes in the plert kingdom and consequently studies on 'icarybtype morphology can be made with an accuracy not possible in most genera. Hybrids between a few species have been rsde (Vosa, 1966b) and a study of meiosis in one of these hybrids has indicated tae structural alterations which have occurred in the evolution of the two species from a remote common ancestor. Taxonomically Thlbaghia has not been subject to any intensive study, and is in great, need of a taxonomic revision, It was also suitable material for a study In which the value of data provided by an investigation of the chromosomes could be assessed. At the same time as investigations within the g'rus, studies were carried out into the relationship of Tuibaghia to r. other genera within the Ltltaoeae and AmaryflIdaeeae,, -3-- and investigations were made to determine the affinities of Tuijaghia to these families. lb. Objectives. The objectives of the present study were: to provide a basic taxonomic revision of Tulbaghia using not only "traditional" morphological characters, but incorporating cytological and anatomical data. Where data from different sources suggest conflicting taxonomic treatments, an evaluation of the relative value of these data in assessing the affinities between species is made. To determine the relationship of Tulbaghia to other genera. Several authors (Hutchinson 1959; Uphof 1943; Smith & Flory 1965) considered Tulbaghia to be closely related to Agapanthus, and to be primitive within the Arnaryllidaceae. More recent work suggested closer affinity with the Liliaceae. It was intended, using similar methods to those mentioned above, to investigate the relationship of Tulbaghia to these, and other probably related genera. ME ic. A DEFINITION OF TULBAGHIA AND A SHORT ACCOUNT OF flELAIED 4ENERA.. (A detailed description appears on p.126). A cursory examination of any species of Tulbaghia suggests a gainopetalous kill-urn with the addition of a corona, either in flhindrical form or as a number (3 or 6) of scales mounted at the throat of the corolla tube. Also suggesting affinity with Allium Is the garlic or onion smell which emanates from the crashed parts of ?ulbaghia species; even material cofleàted in the middle of the eighteenth c€rntury retains this smell. The rootstock can be a rhizome, a cant or a bulb, though traditionally it has been described as rhizomatous only, an error partly accounting for the association of Tulbaghia and Agapantlius in the tribe Agapantheac. tifibaghia. is one of about thirty genera in the Liliaceae (in the sense of Bentham and looker) which possess both a superior ovary (the single character traditionally used to distinguish between the Liiiaceae and the Amaryllidaceae) and an umbellate inflorescence which is characteristic of the Ama- ryllidaceae. These genera were placed by Bentham and Hooker (i88)Jr a single tribe, the Allieae, which wqfurther divided into four subtribes: Agapanthaae, Euailieae, Giflieseae and Massonieae -5- Hutchinson (1943) transferred three of these subtribeo, the A.gapantheae, £llieae and Gillieseas, to the Amaryflidaceae, itising them to tribal rank but left the Massouieae as a tribe of the Lilir;ceae, noting that it was a "climax group showing a close approximation to (the) Amaryflidaoeae". In edition 12 of Bugler's Syllabus (1964) the Agapantheae, Gillieseae and Ailieae appear together, constituting the subfamily Aflioideae of the Liiiaceae. In all the accounts dealing with the Liliales there is a general agreement that wherever these three tribes are put, they should be put together, and except for Hutchinson (1934) 0 Traub (1963) and Huber (1969) they are placed in the Lfliaceae, despite the 'umbellate inflorescence. Data on the Liliales, particularly in the fields of phytoctemistry and karyology, is increasing rapidly, and a reassessment of the whole order is needed (the last comprehensive treatment being Dy Krause, 1930) though Traub (1963) has assembled much data on the Amaryilidaceae. Hither (1oc.oit)in a very complex and detailed paper attempted a classification of the whole of the liltoid monocotyledons (in the broadest sense). Using to a great extent characters from the seeds only, he suggested that the .kmaryllidaoeae should become a "family group" (Featliengruppe) comprising the Hemero calli daceae (one genus: Hemerocaflis), the Alliaceae (with the tribes Brodiaeae, 41liaeae and Glfliesieae) the .Agapanthaceae (with the genera - -6- Agapanthus and Tulbaghia) and the Hyacir.thaceae (with the tribes Chlorogaleae, Bowieae and Sciflieaè).
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