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Crassulaceae, Eurytoma Bryophylli, Fire, Invasions, Madagascar, Osphilia Tenuipes, Rhembastus Sp., Soil

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Crassulaceae, Eurytoma Bryophylli, Fire, Invasions, Madagascar, Osphilia Tenuipes, Rhembastus Sp., Soil B I O L O G I C A L C O N T R O L O F B R Y O P H Y L L U M D E L A G O E N S E (C R A S S U L A C E A E) Arne Balder Roderich Witt A thesis submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg, in fulfillment of the requirements for the degree of Doctor of Philosophy JOHANNESBURG, 2011 DECLARATION I declare that this thesis is my own, unaided work. It is being submitted for the Degree of Doctor of Philosophy in the University of the Witwatersrand, Johannesburg. It has not been submitted before for any degree or any other examination in any other University. ______________________ ______ day of ______________________ 20_____ ii ABSTRACT Introduced plants will lose interactions with natural enemies, mutualists and competitors from their native ranges, and possibly gain interactions with new species, under new abiotic conditions in their new environment. The use of biocontrol agents is based on the premise that introduced species are liberated from their natural enemies, although in some cases introduced species may not become invasive because they acquire novel natural enemies. In this study I consider the potential for the biocontrol of Bryophyllum delagoense, a Madagascan endemic, and hypothesize as to why this plant is invasive in Australia and not in South Africa. Of the 33 species of insects collected on B. delagoense in Madagascar, three species, Osphilia tenuipes, Eurytoma bryophylli, and Rhembastus sp. showed potential as biocontrol agents in Australia. Surveys in southern Africa revealed that B. delagoense had acquired 14 species of insects, compared to only two in Australia, which supported the hypothesis that an introduced plant is less likely to become invasive in regions where there are many closely related species. A beetle, Alcidodes sedi, was found to be widespread and damaging in South Africa and also sufficiently host specific for release in Australia. Studies were also undertaken to determine the biology of B. delagoense and characteristics of the recipient environments in Australia and South Africa. Analysis of soils in South Africa found that B. delagoense has the ability to grow on a range of soils and is tolerant of moderate shade. Bryophyllum delagoense exhibited phenotypic plasticity by re-allocating resources to stem production in light-limited; and to root production in nutrient- or water limited, environments. Competition trials indicated that B. delagoense is a weak competitor with overgrazing and the concomitant reduction in fires facilitating invasions. The absence of damaging natural enemies and characteristics of the Australian iii environment may make it more prone to B. delagoense invasions than in southern Africa. The study indicates that no single trait or group of traits can completely explain the invasive ability of a plant species because the success of an invader often depends on complex interactions between the species and its recipient environment. This has widespread implications for predicting the invasibility of imported plants. Keywords: Alcidodes sedi, biological control, Bryophyllum delagoense, Crassulaceae, Eurytoma bryophylli, fire, invasions, Madagascar, Osphilia tenuipes, Rhembastus sp., soil. iv ACKNOWLEDGEMENTS Thanks to Ms. Y. Rabenantoandro [Centre National de la Recherche Appliquée au Développement Rural (FOFIFA)] for organizing permits for the collection of insects in Madagascar and their subsequent exportation to South Africa and to Mr. J. Rajaonarison (FOFIFA) for assistance in the field in Madagascar. Thanks also go to a host of taxonomists who assisted with the identification of a number of insect species: Mme. H. Perrin (Muséum National d’Histoire Naturelle, Paris), and Dr. M.L. Cox and Z. Boućek, both from the Natural History Museum, London, for identifying O. tenuipes, Rhembastus sp., and Eurytoma sp., respectively; Mr. R. Stals, Mrs. O.C. Neser, Ms. E. Grobbelaar and other taxonomists from the National Collection of Insects of the Plant Protection Research Institute of the Agricultural Research Council (ARC-PPRI), Pretoria, for assisting with the identification of a host of insect species; Professor Wills Flowers (Center for Biological Control (FAMU-CESTA), Florida A&M University) for his opinions concerning the Eumolpinae; and Mr. J. Harrison, head of the Coleoptera Department, Transvaal Museum, for access to that collection. Prof. M. Byrne (University of the Witwatersrand), Prof. M.P. Hill (Rhodes University), Prof. E. Witkowski (University of the Witwatersrand), Prof. J. Hoffmann (University of Cape Town), Dr. W.A. Palmer (Queensland Department of Natural Resources & Mines, Australia), Dr. T. Olckers (University of Kwazulu-Natal), Mr. R. Stals (ARC-PPRI) and Mr. J.C. Scanlan (Department of Primary Industries, Australia) are thanked for the valuable comments/suggestions on previous drafts of some of the chapters/publications. Thanks also to some of the co-authors (Dr. A.J. McConnachie, Ms. S. Docherty, Ms. A. Nongogo) who assisted with laboratory and field trials while other co-authors (Dr. W.A. Palmer, Ms. E. Grobbelaar, Mr. R. Stals and Prof. M. Byrne) contributed valuable technical information. Mr. H.E. Steenkamp who assisted with laboratory trials is v also thanked. Thanks to Ms. L. Henderson who granted me permission to use the line drawing of B. delagoense by Ms. G. Condy and Dr B. Erasmus who kindly provided the South African frost frequency map in Chapter 10 (modified from Schultze et al., 1997) The contribution of Ms. L. Morey and others in the Biometry Division (ARC- PPRI) for assistance with the statistical analyses and to the Weeds Research Division (ARC-PPRI) for making their facilities available is also gratefully acknowledged. Climatic data was provided by the ARC-Institute for Soil Climate and Water and the study was partly funded by the Queensland government. An additional word of thanks to Prof. M. Byrne for his supervision, patience, valuable suggestions and constant encouragement over the past 4 years – his contribution has been invaluable. Last but not least I would like to thank Imelda for her help and for making me knuckle down and finish my PhD – thanks for your support. vi TABLE OF CONTENTS CONTENTS PAGE DECLARATION .......................................................................................................II ABSTRACT ..............................................................................................................III ACKNOWLEDGEMENTS ......................................................................................V TABLE OF CONTENTS .......................................................................................VII LIST OF FIGURES ...............................................................................................XIV LIST OF TABLES .............................................................................................XVIII CHAPTER 1: INTRODUCTION AND THESIS OUTLINE .................................1 CHAPTER 2: LITERATURE REVIEW ................................................................11 CHAPTER 3: INSECTS ASSOCIATED WITH BRYOPHYLLUM DELAGOENSE (CRASSULACEAE) IN MADAGASCAR AND PROSPECTS FOR BIOLOGICAL CONTROL OF THIS WEED Justification .................................................................................................................31 Abstract .......................................................................................................................33 Introduction .................................................................................................................33 Materials and Methods ................................................................................................34 Plant surveys .........................................................................................................34 Insect surveys ........................................................................................................35 Results .........................................................................................................................36 Plant surveys .........................................................................................................36 Insect surveys ........................................................................................................37 Discussion ...................................................................................................................40 Acknowledgements .....................................................................................................43 References ...................................................................................................................44 vii CHAPTER 4: INITIAL SCREENING OF THE STEM-BORING WEEVIL OSPHILIA TENUIPES, A CANDIDATE AGENT FOR THE BIOLOGICAL CONTROL OF BRYOPHYLLUM DELAGOENSE IN AUSTRALIA. Justification .................................................................................................................47 Abstract .......................................................................................................................49 Introduction .................................................................................................................50 Materials and methods ................................................................................................50 Field observations .................................................................................................50 Laboratory cultures and biological studies ...........................................................52
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