DIVERSITY OF HONEY BEE Apis mellifera SUBSPECIES (HYMENOPTERA: APIDAE) AND THEIR ASSOCIATED ARTHROPOD PESTS IN CAMEROON BY DAVID TEMBONG CHAM (I80/92221/2013) (B.Sc. UNIVERSITY OF BUEA-CAMEROON, M.PHIL. UNIVERSITY OF GHANA-LEGON) A THESIS SUBMITTED IN FULFILLMENT OF REQUIREMENTS FOR THE AWARD OF THE DEGREE OF DOCTOR OF PHILOSOPHY IN ENTOMOLOGY SCHOOL OF BIOLOGICAL SCIENCES UNIVERSITY OF NAIROBI 2017 i DECLARATION Candidate I, DAVID TEMBONG CHAM, Registration Number I80/92221/2013, declare that this thesis is my original work and has not been submitted for award of a degree in any other University. Signature__________________________ Date ____________________________ Supervisors This thesis has been submitted with our approval Prof. Paul N. Ndegwa School of Biological Sciences, University of Nairobi, P.O. Box 30197-00100, Nairobi, Kenya Signature __________________________ Date ____________________________ Prof. Lucy W. Irungu School of Biological Sciences, University of Nairobi, P.O. Box 30197-00100, Nairobi, Kenya Signature __________________________ Date ____________________________ Dr. Ayuka T. Fombong International Centre of Insect Physiology and Ecology (ICIPE), P. O. Box 30772-00100, Nairobi, Kenya Signature __________________________ Date ____________________________ Prof. Suresh Raina International Centre of Insect Physiology and Ecology (ICIPE), PO. BOX 30772-00100, Nairobi, Kenya Signature __________________________ Date ____________________________ ii DEDICATION This thesis is dedicated to the Cham’s family iii ACKNOWLEDGEMENTS My sincere gratitude to my supervisors Prof. Paul N. Ndegwa, and Prof. Lucy W. Irungu of the University of Nairobi, and Dr. Ayuka T. Fombong and Prof. Suresh K. Raina of ICIPE for their guidance, invaluable suggestions, support, and reviews that led to the successful completion of this thesis and associated manuscripts. My gratitude also goes to Prof. Baldwyn Torto, Dr(s) Everlyn Nguku, Kiatoko Nkoba, Muli Elliud and David P. Tchouassi for their useful suggestions during research and contributions to the manuscripts that emanated from this thesis. I am also thankful to Dr. Daisy Salifu and Mr. Benedict Orindi for their guidance in data analysis. I also acknowledge the invaluable contributions from lecturers of the School of Biological Sciences, University of Nairobi during Annual Postgraduate Seminars that contributed in improving the quality of this thesis. Special thanks to the anonymous reviewers of this thesis. I am very grateful to all the staff of Environmental Health, especially in the Bee Health Unit, ICIPE for their assistance and useful contributions during scientific presentations. My sincere gratitude go to Capacity Building and Institutional Development staff, especially Dr. Robert Skilton, Mrs. Lilian Igweta-Tonnang, Ms Atieno Vivian, Mrs Lisa Omondi, Mama Margaret Ochanda for administrative assistance and organization of the numerous trainings that boosted my research and writing skills. I acknowledge with appreciation the assistance provided by the Human Resources, Procurement, IT and other administrative Units of ICIPE. Special thanks go to the Ministry of Livestock, Fisheries and Animal Industries (MENIPIA) Cameroon for their support and guidance on the selection of apiary sites and beekeepers. My appreciation also goes to all the Regional Delegates, MINEPIA and the Delegates of beekeeping groups that provided assistance to me in any form during field surveys. I am very grateful to all the beekeepers that provided their apiaries for this study and for their assistance during sampling. Sincere gratitude to Dr (s) David Kupesa, Foba Caroline, Mr. David Makori, Mrs. Nelly Ndungu and to all my colleagues of ICIPE and University of Nairobi for their encouragement, scientific discussions and support. I am also very grateful to all my family members and friends especially my mum Angelina Cham, my aunt Prisca Cham, Dr. P. Ching, Dr. Ndong Cheng, Mr. and Mrs. iv Njana, Mr. and Mrs. Kah Hycinth, Mr. and Mrs. Fuh-Cham, Mr. Awah, Mr. and Mrs Cham Peter for their support, encouragement and prayers. Most importantly, I gratefully acknowledge the financial support for this research by the European Union (EU) under grant number EU/ICIPE 2013/313-659 and the German Academic Exchange Service (DAAD) In-Region Postgraduate Scholarship for my Ph.D. studies. I remain ever grateful to God for good health and guidance throughout this study, and especially for life. v TABLE OF CONTENTS DECLARATION ..................................................................................................ii DEDICATION .................................................................................................... iii ACKNOWLEDGEMENTS.................................................................................iv TABLE OF CONTENTS.....................................................................................vi LIST OF TABLES...............................................................................................xi LIST OF FIGURES ............................................................................................xii APPENDICES ...................................................................................................xiv ABBREVIATIONS AND ACRONYMS...........................................................xv DEFINITION OF TERMINOLOGIES .............................................................xvi ABSTRACT..................................................................................................... xvii CHAPTER ONE .................................................................................................1 INTRODUCTION ................................................................................................1 1.1 General introduction ............................................................................................................1 1.2 Problem statement................................................................................................................3 1.3 Justification and significance of the study...........................................................................4 1.4 Objectives ............................................................................................................................5 1.4.1 Main objective ..................................................................................................................5 1.4.2 Specific objectives ............................................................................................................5 CHAPTER TWO ................................................................................................................6 LITERATURE REVIEW ..........................................................................................................6 2.1 Apis mellifera and related species of the genus Apis ...........................................................6 2.1.1 Diversity and geographic distribution of Apis species......................................................6 2.1.2 Origin and classification of Apis mellifera .......................................................................7 2.1.3 Diversity of Apis mellifera subspecies..............................................................................8 2.1.4 Apis mellifera lineages and their distribution .................................................................10 2.1.5 Methods for characterising Apis mellifera subspecies....................................................10 2.1.5.1 Morphological methods ...............................................................................................11 vi 2.1.5.2 Molecular techniques ...................................................................................................12 2.1.6 The Apis mellifera colony and life history......................................................................13 2.1.7 Ecological and economic importance of honey bees......................................................14 2.2 Honey bee parasites and pests............................................................................................16 2.2.1 Honey bee parasites ........................................................................................................16 2.2.1.1 Varroa mite..................................................................................................................17 2.2.1.2 Tracheal mite Acarapis woodi .....................................................................................18 2.2.1.3 Tropilaelaps clareae ....................................................................................................19 2.2.1.4 Endoparasitic flies........................................................................................................21 2.2.2 Honey bee pests ..............................................................................................................23 2.2.2.1 Aethina tumida and other Nitidulids............................................................................23 2.2.2.2 Large hive beetles ........................................................................................................24 2.2.2.3 Moths ...........................................................................................................................25 2.2.2.3 Bee louse......................................................................................................................26 2.3 Predatory flies of honey bees.............................................................................................26