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Molecular Characterization of Horse Flies (Diptera: Tabanidae) and Determination of Their Role in Transmission of Haemoparasites in Southern Africa

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Molecular Characterization of Horse Flies (Diptera: Tabanidae) and Determination of Their Role in Transmission of Haemoparasites in Southern Africa Molecular characterization of horse flies (Diptera: Tabanidae) and determination of their role in transmission of haemoparasites in southern Africa M.O. Taioe 26849976 Thesis is submitted for the degree Philosophiae Doctor in Zoology at the Potchefstroom Campus of the North-West University Promoter: Prof O.M.M. Thekisoe Co-promoter: Dr M.Y. Motloang September 2017 DEDICATION This thesis is dedicated to my mother Matshidiso Sanna Taioe. For her encouragement, endless love and support that has sustained me throughout my life. I LOVE YOU MA!!! "In the middle of difficulty lies opportunity." -Albert Einstein i ACKNOWLEDGEMENTS This thesis was made possible by assistance and guidance of several people from different countries and I offer my sincere gratitude to all of them. South Africa I express my sincere gratitude to my promoter Prof. Oriel Thekisoe for his continuous support during my studies, for his patience, motivation, and immense knowledge. His guidance helped me at all times. I thank my co-promoter Dr. Makhosazana Motloang for her support, insightful critical comments and corrections of this thesis. I thank Mr Jerome Ntshangase from ARC-Onderstepoort Veterinary Institute (OVI) for his technical assistance during field work for collection of South African tabanid flies. I am grateful to Prof. Huib van Hamburg and Prof. Pieter Theron from the Unit of Environmental Sciences and Management at NWU for their advice and assistance during the identification of the fly specimens. Dr. Charlotte Mienie, I thank you for your assistance in preparing next generation sequencing (NGS) libraries, and sequencing on Illumina MiSeq platform for producing metagenomic data. Ms Joanita Viviers and Ms Jani Reeder from the African Amphibian Conservation Research Group NWU, your assistance in taking sequential photographs of tabanid specimens is greately acknowledged. I thank the following institutions: The Ezemvelo KZN Wildlife for permission to collect fly specimens in their nature reserves and ARC-OVI for allowing me to utilize their facilities at the Kuleni Tsetse station. Lesotho I thank Mr Mabusetsa Makalo from the Department of Livestock Services, Pathology Division for his assistance in sampling for tabanid flies in the Maseru District. Zambia I thank Prof. Boniface Namangala from the Department of Paraclinical Studies, School of Veterinary Sciences, University of Zambia, for his advice and support during sampling in Zambia. ii I thank Mr Amos Chota from School of Veterinary Sciences, University of Zambia for his assistance in the identification of tabanid flies. Dr. Stephan James Phiri, Mr Albert Mwiinga, Mr Samuel Wilson Sakala and Mr Banda Totti I thank you for your assistance during sampling of tabanid flies in different districts of Zambia. Japan I thank Prof. Noboru Inoue from the National Research Centre for Protozoan Diseases, Obihiro University of Agriculture and Veterinary Medicine for allowing me to use his research facilities during molecular detection of protozoan parasites. Prof. Keisuke Suganuma, Dr. Peter Musinguzi and Ms Nthatisi Molefe I thank you all for your assistance and support during my stay in Japan. Malaysia I thank Prof. Asif Khan from Centre for Bioinformatics, Perdana University for allowing me to use his facilities for metagenomic analysis. Dr. Lloyd Low from the Centre for Bioinformatics Perdana University, I thank you for advice, assistance and support for my metagenomic analysis. Mr Muhammad Farhan and Ms Tan Swan from the Centre for Bioinformatics, your assistance and support during my stay in Malaysia is greately acknowledged. Family and friends I thank my family, friends and fellow students for the words of encouragement, jokes during hard times and support throughout the duration of this study. Financial Support DST-NRF Scarce Skills Scholarship Grant UID: 95090 awarded to Mr Moeti Oriel Taioe. National Research Foundation (NRF) Development Grant for Y-rated Researchers made available to Prof. Oriel Thekisoe. Above all, I praise the almighty God, for providing me with this opportunity and granting me the strength and capability to complete this journey successfully. As stated in iii Proverbs 3:5-6, “ Trust in the Lord with all your heart. Don't put your confidence in your own understanding. In all your ways acknowledge him, and he will direct your path." God Bless You All Thank you iv TABLE OF CONTENTS DEDICATION ....................................................................................................................... i ACKNOWLEDGMENTS .................................................................................................... ii TABLE OF CONTENTS ..................................................................................................... v RESEARCH OUTPUTS ................................................................................................... xv ABSTRACT ..................................................................................................................... xvi CHAPTER 1: GENERAL INTRODUCTION ...................................................................... 1 1.1 Background ................................................................................................................... 1 1.2 Biology of Tabanidae .................................................................................................... 2 1.2.1 Taxonomy .................................................................................................................. 2 1.2.2 Morphology of adult flies ........................................................................................... 4 1.2.3 Reproduction ............................................................................................................. 6 1.2.4 Behaviour ................................................................................................................... 8 1.3 Economic, medical and veterinary importance of Tabanidae ..................................... 9 1.3.1 Tabanids as plant pollinators .................................................................................... 9 1.3.2 Tabanids as vectors of pathogens ............................................................................ 9 1.4 Justification of the study ............................................................................................. 10 1.4.2 Diseases of interest in the current study ................................................................ 11 1.4.2.1 Babesiosis ............................................................................................................ 11 1.4.2.2 Besnoitiosis........................................................................................................... 11 1.4.2.3 Theileriosis............................................................................................................ 12 1.4.2.4 Trypanosomiasis .................................................................................................. 13 1.4.2.5 Symbiotic associations in haematophagous insects ........................................... 14 1.7 Aims and objectives.................................................................................................... 16 1.7.1 Aims ......................................................................................................................... 16 v 1.7.2 Objectives ................................................................................................................ 16 CHAPTER 2: MORPHOLOGICAL AND MOLECULAR CHARACTERIZATION OF TABANID FLIES FROM LESOTHO, SOUTH AFRICA AND ZAMBIA .................. 17 2.1 Introduction ................................................................................................................. 17 2.2 Materials and methods ............................................................................................... 20 2.2.1 Study area ............................................................................................................... 20 2.2.1.1 Kingdom of Lesotho ............................................................................................. 20 2.2.1.2 The Republic of South Africa ............................................................................... 20 2.2.1.3 Zambia .................................................................................................................. 21 2.2.2 Collection of samples .............................................................................................. 22 2.2.3 Identification and characterization of tabanid flies ................................................. 27 2.2.4 DNA extraction from tabanid flies ........................................................................... 27 2.2.5 Molecular amplification of cytochrome oxidase 1 (CO1) gene .............................. 28 2.2.6 Purification of PCR amplicons ................................................................................ 28 2.2.7 Sequencing and BLASTn ........................................................................................ 29 2.2.8 Statistical analysis ................................................................................................... 29 2.2.9 Phylogenetic analysis .............................................................................................. 30 2.3 Results...... .................................................................................................................. 31 2.3.1 Tabanids from Lesotho...........................................................................................
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