Generic relationships of selected African genera of Apiaceae by Anthony Richard Magee Thesis submitted in fulfilment of the requirements for the degree PHILOSOPHIAE DOCTOR in BOTANY in the FACULTY OF SCIENCE at the UNIVERSITY OF JOHANNESBURG SUPERVISOR: PROF. B.-E. VAN WYK CO-SUPERVISOR: PROF. P. M. TILNEY CO-SUPERVISOR: PROF. S. R. DOWNIE September 2009 AFFIDAVIT: MASTER'S AND DOCTORAL STUDENTS TO WHOM IT MAY CONCERN This serves to confirm that I Anthony Richard Magee Full Name(s) and Surname ID Number 8109065065089 Student number 920001887 enrolled for the Qualification PhD (Botany) Faculty Science Herewith declare that my academic work is in line with the Plagiarism Policy of the University of Johannesburg which I am familiar. I further declare that the work presented in the Generic relationships of selected African genera of Apiaceae (thesis) is authentic and original unless clearly indicated otherwise and in such instances full reference to the source is acknowledged and I do not pretend to receive any credit for such acknowledged quotations, and that there is no copyright infringement in my work. I declare that no unethical research practices were used or material gained through dishonesty. I understand that plagiarism is a serious offence and that should I contravene the Plagiarism Policy notwithstanding signing this affidavit, I may be found guilty of a serious criminal offence (perjury) that would amongst other consequences compel the UJ to inform all other tertiary institutions of the offence and to issue a corresponding certificate of reprehensible academic conduct to whomever request such a certificate from the institution. Signed at Johannesburg on this the 2 S' September 2009 Signature Print name A f-) Ti-V7 H RqC-C. STAMP COMMISSIONER OF OATHS Affidavit certified by a Commissioner of Oaths This affidavit conforms with the requirements of the JUSTICES OF THE PEACE AND COMMISSIONERS OF OATHS ACT 16 OF 1963 and the applicable Regulations published in the GG GNR 1258 of 21 July 1972; GN 903 of 10 July 1998; GN 109 of 2 February 2001 as amended. "There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved." — CHARLES R. DARWIN (The Origin of Species) TABLE OF CONTENTS SUMMARY i ACKNOWLEDGEMENTS VII CHAPTER 1: GENERAL INTRODUCTION AND OBJECTIVES 1 1 .1 GENERAL INTRODUCTION 1 1.2 OBJECTIVES 3 CHAPTER 2: GENERAL MATERIALS AND METHODS 6 2.1 MORPHOLOGY 6 2.2 FRUIT ANATOMY AND VITTAE STRUCTURE 7 2.3 DATA CAPTURING 8 2.4 PHYLOGENETIC ANALYSES 8 CHAPTER 3: SYSTEMATICS OF THE CAPNOPHYLLUM GROUP AND PUTATIVE RELATIVES 12 3.1 INTRODUCTION 12 3.2 MATERIAL AND METHODS 14 3.3 RESULTS 18 3.4 DISCUSSION 30 3.5 KEY TO GENERA OF THE CAPNOPHYLLUM GROUP 35 CHAPTER 4: TAXONOMIC REVISION OF THE GENUS CAPNOPHYLLUM 36 4.1 INTRODUCTION 36 4.2 VEGETATIVE AND REPRODUCTIVE MORPHOLOGY 37 4.3 TAXONOMY OF THE GENUS CAPNOPHYLLUM.. 44 CHAPTER 5: TAXONOMIC REVISION OF THE GENUS DASISPERMUM 58 5.1 INTRODUCTION 58 5.2 VEGETATIVE AND REPRODUCTIVE MORPHOLOGY 59 5.3 TAXONOMY OF THE GENUS DASISPERMUM 68 CHAPTER 6: TAXONOMIC REVISION OF THE GENUS SCARABOIDES 97 6.1 INTRODUCTION 97 6.2 VEGETATIVE AND REPRODUCTIVE MORPHOLOGY 98 6.3 TAXONOMY OF THE GENUS SCARABOIDES.. 100 CHAPTER 7: SYSTEMATICS OF AFRICAN AND MALAGASY PIMPINELLA AND RELATED GENERA 104 7.1 INTRODUCTION 104 7.2 MATERIAL AND METHODS 105 7.3 RESULTS AND DISCUSSION 106 CHAPTER 8: SYSTEMATICS OF MALAGASY PEUCEDANUM 117 8.1 INTRODUCTION 117 8.2 MATERIAL AND METHODS 118 8.3 RESULTS AND DISCUSSION 119 8.4 TAXONOMY OF THE GENUS BILLBURTTIA 125 CHAPTER 9: SYSTEMATICS OF THE GENUS EZOSCIADIUM 128 9.1 INTRODUCTION 128 9.2 MATERIAL AND METHODS 129 9.3 RESULTS AND DISCUSSION 130 9.4 TAXONOMY OF THE GENUS EZOSCIADIUM 139 CHAPTER 10: NEW TRIBAL DELIMITATIONS OF THE EARLY DIVERGING LINEAGES OF APIACEAE SUBFAMILY APIOIDEAE 143 10.1 INTRODUCTION 143 10.2 MATERIAL AND METHODS 144 10.3 RESULTS 148 10.4 DISCUSSION 155 10.5 TAXONOMY OF THE PROTOAPIOID TRIBES 165 CHAPTER 11: GENERAL CONCLUSIONS 171 LITERATURE CITED 177 APPENDIX A: VOUCHER SPECIMENS OF FRUIT MATERIAL STUDIED 198 APPENDIX B: DNA ACCESSIONS USED FOR THE PHYLOGENETIC ANALYSES 202 APPENDIX C: PUBLICATIONS RESULTING FROM THIS STUDY 216 SUMMARY Recent anatomical and molecular studies have highlighted the importance of the African and Malagasy Apiaceae, many of which have been found to occupy early diverging positions within the subfamilies Apioideae and Saniculoideae. Despite the recent interest in the African contingent however, there remain several anomalous and poorly known African and Malagasy taxa in which generic boundaries remain unclear and which have yet to be incorporated within the emerging tribal classification for the family. Generic circumscriptions and affinities amongst hitherto poorly known African and Malagasy genera are here explored using anatomical, cytological, morphological and molecular sequence data. Substantial rearrangements at almost all infrafamilial levels are formalized in order to incorporate the unique African and Malagasy members for the first time. Generic circumscriptions and phylogenetic relationships of the Cape genera Capnophyllum, Dasispermum, and Sonderina are explored through parsimony and Bayesian inference analyses of nrDNA ITS and cpDNA rps16 intron sequences, morphology, and combined molecular and morphological data. The relationship of these genera with the North African genera Krubera and Stoibrax is also assessed. Analyses of both molecular data sets place Capnophyllum, Dasispermum, Sonderina, and the only southern African species of Stoibrax (S. capense) within the newly recognized Lefebvrea Glade of tribe Tordylieae. Capnophyllum is strongly supported as monophyletic and is distantly related to Krubera. The monotypic genus Dasispermum and Stoibrax capense are embedded within a paraphyletic Sonderina. This complex is distantly related to the North African species of Stoibrax in tribe Apieae, in which the type species, Stoibrax dichotomum, occurs. Consequently, i SUMMARY Dasispermum is expanded to include both Sonderina and Stoibrax capense. A taxonomic revision of Dasispermum s.l. is presented which includes keys to the species, complete nomenclature, typifications, descriptions as well as geographical distributions. New combinations are formalized for Dasispermum capense, 0 hispidum, D. humile, and D. tenue. In addition two new species, namely D. grandicarpum and D. perrenans, are described. As a result seven species of Dasispermum s.l. are recognised and can be distinguished from one another by their habit (life history and growth form), leaf morphology (leaf texture, leaf colour and breadth of the ultimate leaflet segments), inflorescence structure (length of the peduncle, presence or absence and division of involucre and involucel bracts), fruit morphology (relative length of the styles, fruit size, rib prominence and relative orientation) and fruit anatomy (shape of the cells external to the vittae). The Cape endemic genus Capnophyllum is revised. As a result of valuable recent collections and extensive fieldwork, this hitherto neglected genus was found to comprise four annual species, two of which are newly described, namely Capnophyllum lutzeyeri and C. macrocarpum. The four species are distinguished from one another by their fruit morphology (relative length of the styles, the shape and position of the stylopodium, fruit size, surface sculpturing, and the presence or absence of a sterile apical portion) and fruit anatomy (marginal wings slightly or prominently involute and secondary ribs present or absent). A comprehensive key to the species, their complete nomenclature and typification, together with complete descriptions and known geographical distributions for all the species are presented and illustrated. The genus Scaraboides is described herein to accommodate a new species, S. manningii, from the Tanqua Karoo in South Africa. This monotypic genus shares ii SUMMARY the dorsally compressed fruit and involute marginal wings with Capnophyllum, but is easily distinguished by its erect branching habit, green leaves, scabrous umbels, and fruit with indistinct median and lateral ribs, additional solitary vittae in each marginal wing, and parallel, closely spaced commissural vittae. Despite the marked fruit similarities with Capnophyllum, analyses of DNA sequence data place Scaraboides closer to Dasispermum, with which it shares the erect habit, green (non-glaucous) leaves, and scabrous umbels. The phylogenetic position of the African and Malagasy species of Pimpinella are assessed using nrITS sequence data. These species are found to ally with their Eurasian counterparts within the tribe Pimpinelleae. The genus Pimpinella is rendered paraphyletic by the inclusion of African Cryptotaenia and the small African and Malagasy endemic genera Frommia and Phellolophium. Within the paraphyletic Pimpinella three major clades are recovered, with the African species widely separated into two of the three clades. Based on the results of the molecular analyses it is clear that the current sectional classification for the genus, based largely on fruit vestiture, is largely artificial. Chromosome base number, however, was found to be consistent with the groupings recovered in the molecular trees. Optimisation