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Taxonomic revision of the Memecylon natalense species-complex in southern Africa by IMERCIA GRACIOUS MONA Submitted in partial fulfilment of the academic requirements of Master of Science in Plant Taxonomy School of Life Sciences College of Agriculture, Engineering and Science University of KwaZulu-Natal Westville South Africa Date: (September, 2018) PREFACE The research contained in this dissertation was completed by the candidate while based in the Discipline of Biology School of Life Sciences of the College of Agriculture, Engineering and Science, University of KwaZulu-Natal, Westville, South Africa. The research was financially supported by the National Research Foundation and the Botanical Educational Trust. The contents of this work have not been submitted in any form to another university and, except where the work of others is acknowledged in the text, the results reported are due to investigations by the candidate. _________________________ Signed: Dr Syd Ramdhani Date: 10/09/2018 _________________________ Signed: Dr Robert Douglas Stone Date: 10/09/2018 i DECLARATION 1: PLAGIARISM I, Imercia Gracious Mona, declare that: (i) the research reported in this dissertation, except where otherwise indicated or acknowledged, is my original work; (ii) this dissertation has not been submitted in full or in part for any degree or examination to any other university; (iii) this dissertation does not contain other persons’ data, pictures, graphs or other information, unless specifically acknowledged as being sourced from other persons; (iv) this dissertation does not contain other persons’ writing, unless specifically acknowledged as being sourced from other researchers. Where other written sources have been quoted, then: (a) their words have been re-written but the general information attributed to them has been referenced; (b) where their exact words have been used, their writing has been placed inside quotation marks, and referenced; (c) where I have used material for which publications followed, I have indicated in detail my role in the work; (v) this dissertation is primarily a collection of material, prepared by myself, published as journal articles or presented as poster or oral presentations at conferences. In some cases, additional material has been included; (vi) this dissertation does not contain text, graphics or tables copied and pasted from the Internet, unless specifically acknowledged, and the source being detailed in the dissertation and in the References sections. ____________________ Signed: Imercia Date: 10/09/2018 ii DECLARATION 2: PUBLICATIONS My role in each paper is indicated. The * indicates corresponding author. 1. Mona, I.G.,* Stone, R.D., 2016. Morphometric analysis and taxonomic revision of the Memecylon natalense species-complex (Melastomataceae) in southern Africa. South African Journal of Botany 103, 334. [Conference Abstract] https://doi.org/10.1016/j.sajb.2016.02.11 Contribution - I contributed to the writing of the abstract, mini oral and poster presentation at the South African Association of Botanists meeting held in January 2016. 2. Stone, R.D.,* Mona, I.G., Ramdhani, S., 2017. Revised treatment of Mozambican Memecylon (Melastomataceae—Olisbeoideae), with descriptions of four new species in M. section Buxifolia. Phytotaxa 331, 151–168. https://biotaxa.org/Phytotaxa/article/view/phytotaxa.331.2.1/30718 Contribution - I contributed to the morphological analyses and species descriptions, and together with the other two authors we all contributed substantially to the writing of the manuscript. ___________________ Signed: Imercia Date: 10/09/2018 iii ABSTRACT Two Memecylon species (M. natalense and M. bachmannii) were previously recognized in southern Africa, and three species of M. section Buxifolia as occurring in Mozambique (M. natalense, M. torrei and M. insulare). However, a recent nrDNA phylogeny revealed that M. natalense as previously circumscribed is not a monophyletic group and includes some geographically outlying populations warranting recognition as distinct evolutionary lineages. In this dissertation, I present the results of a comprehensive morphological study and revision of the ‘M. natalense species-complex’ including the following taxa: M. natalense sensu stricto (endemic to South Africa in KwaZulu-Natal, Eastern Cape, Mpumalanga and Limpopo), M. incisilobum (southern Mozambique), M. nubigenum (southern Malawi), M. rovumense (southeastern Tanzania), M. aenigmaticum (northern Mozambique), M. sp. nov. 1 (northeastern Kwazulu-Natal, South Africa), M. sp. nov. 2 (Eastern Cape province, South Africa), and M. sp. nov. 3 (Limpopo province, South Africa). Although M. bachmannii (Eastern Cape and KwaZulu-Natal) is morphologically distinct, it was also included in the present study because of recent nrDNA work indicating a close relationship with M. natalense. Species boundaries were investigated using phenetic and cladistic methods based on morphology. Phenetic analysis using the Unweighted Pair Group Method with Arithmetic Mean (UPGMA) revealed that M. bachmannii clustered independently, M. sp. nov. 1 clustered close to M. incisilobum, M. aenigmaticum clustered close to M. rovumense, and M. nubigenum clustered close to a subcluster consisting of M. natalense, M. sp. nov. 2 and M. sp. nov. 3. The morphological cladistic results were congruent with a previous molecular analysis in that M. sect. Buxifolia was monophyletic. Memecylon rovumense + M. aenigmaticum and M. incisilobum + M. sp. nov. 1 appeared in separate clades. Memecylon bachmannii and M. nubigenum were each monophyletic, while M. natalense, M. sp. nov. 2 and M. sp. nov. 3 appeared in an unresolved polytomy. Overall, the cladistic results were in agreement with the UPGMA analysis but were less resolved. Vegetative and fruit morphology have proven to be of great diagnostic value in delimiting the members of the M. natalense species-complex. Four new species have been named and described, while a further three new species are proposed. Dichotomous keys are provided, and the conservation status of each species is assessed according to the criteria of the iv IUCN. A lectotype should be designated for M. natalense. With these additions, the flora of Mozambique now has seven species of Memecylon (including six species in M. sect. Buxifolia). In South Africa, the number of recognized Memecylon species has increased from two to five. v ACKNOWLEDGMENTS A number of people deserve specific mention: Mrs Jane T. Mabilane, my Mother, for her support; My supervisor Dr Syd Ramdhani and my co-supervisor Dr Robert Douglas Stone of the University of KwaZulu-Natal (UKZN); The curators of the following herbaria for providing loans and or access to specimens and specimen-images: Buffelskloof Nature Reserve (BNRH), Botanic Garden Meise (BR), California Academy of Sciences (CAS), University of Coimbra (COI), Royal Botanical Gardens, Kew (K), Instituto de Investigação Científica Tropical (LISC), Institute for Agricultural Research of Mozambique (LMA), Eduardo Mondlane University (LMU), Missouri Botanical Garden (MO), Natal Herbarium (NH), Bews Herbarium (NU), Muséum National d’Histoire Naturelle, Paris (P), National Herbarium, Pretoria (PRE), H.G.W.J. Schweickerdt Herbarium (PRU), National Herbarium, Harare (SRGH), University of Bayreuth (UBT), Ward Herbarium (UDW), Naturalis (WAG); The line drawings and photographs were provided by: Sandie Burrows, David Styles, Dr Robert Douglas Stone and Prabha Amarashinghe, Dr Syd Ramdhani, Hermenegildo Matimele, John Burrows and Richard Boon; Dr Christina Potgieter and Dr Benny Bytebier of Bews Herbarium (UKZN) for technical support; The following people for assistance with field work: the Pondoland CREW team and Teddy Govender (at Umtamvuna Nature Reserve); Sharon Louw (at Ongoye Forest Reserve); Bobby Stone and Daniel Joel Engotto (at Cwebe and Hluleka Nature Reserves, Eastern Cape) and David Styles (at Sihadla Forest, Kosi Bay); Dr Jacques van Rooyen of SANBI (PRE) for the conference poster and oral presentation facilitation. The anonymous reviewers of our manuscript submitted for publication in the journal Phytotaxa (Stone, Mona & Ramdhani, 2017); The various agencies for funding support: the South African Department of Science and Technology and NRF (DST-NRF); the Botanical Education Trust (BET) and the South African National Biodiversity Institute (SANBI). vi TABLE OF CONTENTS PREFACE ....................................................................................................................... i DECLARATION 1: PLAGIARISM ............................................................................. ii DECLARATION 2: PUBLICATIONS ........................................................................ iii ABSTRACT ................................................................................................................. iv ACKNOWLEDGMENTS ............................................................................................ vi TABLE OF CONTENTS ............................................................................................ vii LIST OF TABLES ...................................................................................................... viii LIST OF FIGURES ...................................................................................................... ix CHAPTER 1: INTRODUCTION ................................................................................... 1 CHAPTER 2: MORPHOLOGY ....................................................................................