Nymphaea Mexicana Zuccarini (Nymphaeaceae) in South Africa
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INITIATING BIOLOGICAL CONTROL FOR NYMPHAEA MEXICANA ZUCCARINI (NYMPHAEACEAE) IN SOUTH AFRICA A thesis submitted in fulfilment of the requirements for the degree of Masters in Entomology at Rhodes University By MEGAN REID December 2019 RU-AREC ERAS ethics approval number: 2019-0371-304 Table of contents Abstract .......................................................................................................................................... 1 Acknowledgements ..................................................................................................................... 3 List of figures ............................................................................................................................... 5 List of tables ................................................................................................................................. 9 Chapter 1: General introduction ............................................................................................. 11 1.1 INVASIVE ALIEN PLANTS ......................................................................................... 11 1.2 MANAGEMENT OF INVASIVE ALIEN PLANTS ..................................................... 13 Mechanical control ................................................................................................................ 14 Chemical control.................................................................................................................... 15 Biological and integrated control .......................................................................................... 15 1.3 HISTORY OF AQUATIC INVASIONS IN SOUTH AFRICA ..................................... 19 1.4 NYMPHAEA MEXICANA................................................................................................ 21 Distribution in South Africa .................................................................................................. 24 Hybrids .................................................................................................................................. 25 Impact .................................................................................................................................... 26 Current control ....................................................................................................................... 26 Potential impact ..................................................................................................................... 28 1.5 THESIS OUTLINES AND AIMS .................................................................................. 28 Chapter 2: Matching invasive Nymphaea mexicana to native range populations ....... 30 2.1 INTRODUCTION ........................................................................................................... 30 2.2 MATERIALS AND METHODS .................................................................................... 33 Sampling ................................................................................................................................ 33 DNA extraction...................................................................................................................... 34 ISSR PCR protocol and analysis ........................................................................................... 34 i Electropherograms ................................................................................................................. 39 ISSR scoring parameters ....................................................................................................... 39 Analysis of full dataset .......................................................................................................... 39 STRUCTURE analysis .......................................................................................................... 41 2.3 RESULTS ........................................................................................................................ 42 Preliminary tests .................................................................................................................... 42 ISSR data ............................................................................................................................... 43 STRUCTURE analysis .......................................................................................................... 48 2.4 DISCUSSION .................................................................................................................. 49 Chapter 3: Surveys for potential biological control agents for Nymphaea mexicana, with notes on field host specificity* .................................................................................................... 56 3.1 INTRODUCTION ........................................................................................................... 56 3.2 MATERIALS AND METHODS .................................................................................... 61 MaxEnt modelling and surveys ............................................................................................. 61 Host specificity pilot study .................................................................................................... 65 3.3 RESULTS ........................................................................................................................ 65 MaxEnt modelling ................................................................................................................. 65 Surveys .................................................................................................................................. 66 Host specificity pilot studies.................................................................................................. 78 Prioritisation .......................................................................................................................... 79 3.4 DISCUSSION .................................................................................................................. 80 Chapter 4: General discussion ................................................................................................ 86 4.1 INTRODUCTION ........................................................................................................... 86 4.2 MATCHING INVASIVE NYMPHAEA MEXICANA TO ITS NATIVE RANGE ......... 87 4.3 SURVEYS FOR POTENTIAL BIOLOGICAL CONTROL AGENTS ......................... 88 4.4 PREDICTING EFFICACY OF BIOLOGICAL CONTROL AGENTS ......................... 91 4.5 BIOLOGICAL CONTROL OF HYBRIDS .................................................................... 95 ii 4.6 IMPORTANCE OF ANTHROPOGENIC INFLUENCE ON INVASIONS.................. 99 4.7 LIMITATIONS AND RECOMMENDATIONS .......................................................... 100 4.8 CONCLUSION ............................................................................................................. 102 References ................................................................................................................................. 104 iii Abstract Nymphaea mexicana Zuccarini (Nymphaeaceae) is an aquatic plant originating from south-eastern USA that is becoming increasingly invasive in South Africa as other invasive aquatic plants are being managed successfully through biological control. Mechanical and chemical control of aquatic weeds is expensive, damaging to the environment, and only effective in the short term, so biological control is more desirable as a management strategy for N. mexicana. The biological control of invasive alien plants requires that agents are host specific so that non-target risks are mitigated. For success to be achieved, it is important to ensure that the genetic structure of invasive populations is clarified so that agents can be collected from populations in the native range that match genetically to populations in the invasive range. This is especially important in cases where the morphology of invasive alien plants does not reflect genetic differences between populations. A previous study of the genetic structure of the invasive populations of N. mexicana in South Africa suggests the presence of hybrid forms of the plant in South Africa, with only one of these populations matching with samples from the native range. However, the study only used samples from two sites in the native range using amplified fragment length polymorphisms (AFLPs), so it was necessary to conduct further genetic analyses using samples from more sites in the native range. Hence, the first aim of this study was to develop a better understanding of the genetic structure of N. mexicana populations in the native and invaded range. Genetic samples were collected from sites in the native range during field surveys for potential biological control agents, and inter-simple sequence repeats (ISSRs) were used to compare the genetic structure of invasive and native populations of N. mexicana in South Africa. The results from these analyses suggest that seven of the 14 investigated invasive populations of N. mexicana in South Africa are genetically similar to populations in the native range, while the remaining seven populations are likely to be hybrid forms of the plant. This knowledge will be useful to target populations for biological control and highlights the need for further genetic analyses to determine the parentage of these hybrids so that biological control efforts are more likely to be successful. The initiation