A Taxonomic Study of the Thesium Goetzeanum Species Complex (Santalaceae)
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A taxonomic study of the Thesium goetzeanum species complex (Santalaceae) By Natasha Visser Dissertation Submitted in fulfilment of the requirements for the degree Master in Botany at the University of Johannesburg Supervisor: Prof Ben-Erik van Wyk Co-supervisor: Dr M. Marianne le Roux May 2018 Thesium goetzeanum complex Content Contents Summary .................................................................................................................................... 1 Chapter 1 Introduction ............................................................................................................... 1 Chapter 2 Material and Methods................................................................................................ 5 2.1 Herbarium material ....................................................................................................................... 5 2.2 Field studies .................................................................................................................................. 5 2.3 Gross morphology ......................................................................................................................... 5 2.4 Distribution maps .......................................................................................................................... 6 2.5 Anatomical sections ...................................................................................................................... 7 2.5.1 Flowers and leaves ................................................................................................................. 7 2.5.1 Stems ...................................................................................................................................... 7 2.7 Flower clearing ............................................................................................................................. 7 2.6 Conservation status assessments ................................................................................................... 8 2.7 DNA extraction, amplification, sequencing and alignment .......................................................... 8 2.8 Phylogenetic analyses ................................................................................................................. 10 2.9 Morphological characters ............................................................................................................ 11 2.9.1 Cladistic analysis of the Thesium goetzeanum complex ..................................................... 12 2.9.2 Ancestral character reconstruction ....................................................................................... 12 Chapter 3 Ethnobotany ............................................................................................................ 20 3.1 General uses of Thesium ............................................................................................................. 20 3.2 Southern Africa ........................................................................................................................... 21 3.3 Africa .......................................................................................................................................... 24 3.4 Asia ............................................................................................................................................. 25 3.5 Australia ...................................................................................................................................... 26 3.6 Conclusions ................................................................................................................................. 26 Chapter 4 Morphology ............................................................................................................. 31 4.1 Vegetative morphology ............................................................................................................... 31 4.1.1 Habit ..................................................................................................................................... 31 4.1.2 Rootstock ............................................................................................................................. 34 4.1.3 Stems .................................................................................................................................... 37 4.1.4 Leaves .................................................................................................................................. 41 4.2 Reproductive morphology .......................................................................................................... 44 4.2.1 Inflorescences ...................................................................................................................... 44 4.2.2 Bracts and bracteoles ........................................................................................................... 47 4.2.3 Number of tepals .................................................................................................................. 49 4.2.4 External perianth ‘glands’ .................................................................................................... 49 i Thesium goetzeanum complex Content 4.2.5 Style length and stigma position .......................................................................................... 53 4.2.6 Ovaries and placenta ............................................................................................................ 55 4.2.7 Fruit ...................................................................................................................................... 56 4.3 Conclusions ................................................................................................................................. 58 Chapter 5 Phylogenetic relationships....................................................................................... 59 5.1 Introduction ................................................................................................................................. 59 5.2 Results ......................................................................................................................................... 61 5.2.1 Cladistic analysis of the Thesium goetzeanum complex ...................................................... 61 5.2.2 ITS dataset ........................................................................................................................... 62 5.2.3 TrnL-trnF dataset ................................................................................................................. 62 5.2.4 Combined ITS and trnL-trnF dataset ................................................................................... 63 5.2.5 Ancestral character reconstruction ....................................................................................... 64 5.3 Discussion ................................................................................................................................... 65 5.4 Conclusions ................................................................................................................................. 68 Chapter 6 Taxonomy of the Thesium goetzeanum species complex ....................................... 78 6.1 Key to the species in the Thesium goetzeanum complex ............................................................ 78 6.2 Taxonomic treatment .................................................................................................................. 81 6.2.1 T. goetzeanum ...................................................................................................................... 81 6.2.2 T. gracilarioides ................................................................................................................... 92 6.2.3 T. gracile .............................................................................................................................. 97 6.2.4 T. gypsophiloides ............................................................................................................... 101 6.2.5 T. infundibulare .................................................................................................................. 106 6.2.6 T. lobelioides ...................................................................................................................... 109 6.2.7 T. magalismontanum .......................................................................................................... 113 6.2.8 T. procerum .......................................................................................................................