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Analysis of Varroa Destructor Infestation of Southern African Honeybee Populations

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Analysis of Varroa Destructor Infestation of Southern African Honeybee Populations ANALYSIS OF VARROA DESTRUCTOR INFESTATION OF SOUTHERN AFRICAN HONEYBEE POPULATIONS Dissertation submitted in fulfilment of the requirements for the degree of Master of Science in the Faculty of Natural & Agricultural Science University of Pretoria Pretoria MIKE ALLSOPP June 2006 ___________________________________________________________________________ ABSTRACT ___________________________________________________________________________ The discovery of the honeybee-specific ectoparasitic mite Varroa destructor in South Africa in October 1997 raised the spectre of massive honeybee colony losses as has occurred in most parts of the world where the varroa mite has been found. This was particularly concerning in Africa because of the importance of honeybees in the pollination of indigenous and commercial crops, and because of the numbers of small-scale beekeepers in Africa. The mite has now spread throughout South Africa and is found in almost all honeybee populations, both commercial and wild, and is also now present in most neighbouring countries. Varroa has not left a trail of destruction in South Africa as had been expected and no large scale collapse of the honeybee population occurred, despite the majority of beekeepers deciding not to protect their hives with chemical varroacides. Some colony losses did occur at the front of the varroa spread, and all colonies were found to be deleteriously affected by the mite which developed populations of 50 000 and more in some colonies. Infected colonies were also not as efficient as pollinators as uninfected colonies. Colonies exhibited all the same varroa effects witnessed in other parts of the world, with the exception that the majority of colonies did not die as a result of the infestation. The relative tolerance of African bees to the varroa mite has been confirmed by the long- term monitoring of both wild honeybee populations and commercial stock, and by population dynamic studies of the mites. In both wild and managed honeybee populations varroa appears to have been reduced to the status of an incidental pest. The development of mite tolerance took 3-5 years in the Cape honeybee (Apis mellifera capensis) and 6-7 years in the Savanna honeybee (Apis mellifera scutellata). The rapid development of mite tolerance in the Cape bee is thought to be due to the well developed removal of varroa-infested brood and the short post-capping period of worker brood. Together these resulted in a very rapid increase in infertile mites in the colony, the collapse of the mite population, and varroa tolerance. Tolerance does not develop as rapidly in Savanna honeybees as the post-capping period in these bees is similar to that of European bees and does not result in as many infertile mites. Nonetheless, varroa tolerance in Savanna bees develops more rapidly than would be the case in European bees because of more effective hygienic removal of varroa-infested brood. In both Cape and Savanna bees, the absence of varroacide applications and a “live-and-let-die” approach to the wild and commercial honeybee populations was crucial to the developed of population-wide varroa tolerance, in contrast to the selective breeding and pesticide treadmill practised in most parts of the world in an effort to get rid of the varroa mite. Varroa destructor is concluded not to be a serious threat to honeybees and beekeeping in Africa, and efforts should be made to prevent the use of pesticides and techniques that could hinder the development of natural mite tolerance in Africa. ii ___________________________________________________________________________ DECLARATION ___________________________________________________________________________ I declare that “Analysis of Varroa destructor infestation of southern African honeybee populations” is my own work unless otherwise indicated. ________________________ Mike Allsopp June 9th 2006 iii ___________________________________________________________________________ TABLE OF CONTENTS ___________________________________________________________________________ Abstract.......………………………………………………………………………………………………....……ii Declaration…………………………..………………………………………………………..……………….…iii Table of Contents ……………………………………………………………………………….…….……..…iv List of Figures ……………………………………..…………………………………………………….……..vii List of Tables …………………………………………………………………………………………….……...ix List of Appendices ……………………………………………………………...………………………..…….xii Acknowledgements ……………………………………………………….…...…………………….….…….xiii Publications & Conference Proceedings ….……………………………………….…………………..……xiv CHAPTER ONE: Varroa Mites and South African honeybees Varroa destructor Description …………………………………………………………………………………….1 Spread ………......................…………………...……………………………………………2 Life Cycle …………………………………………………….………………………………..3 Impact …..………………...……………………………………….....................................5 Treatment ……………………………………………………………………………....……..6 South African honeybees Biology ………………………………………………………….……………………………..9 Value of honeybees in South Africa...……………………………………………………..10 Potential Impact of varroa mites on beekeeping & honeybees in South Africa .......................12 Scope & Aims …………...……………………………………………………………………………14 CHAPTER TWO: The introduction and spread of varroa mites in South Africa Introduction ……………………………………………………….…………………………….……..15 Materials & Methods August/October 1997 Survey ………………………………..……………………….……18 Subsequent surveys and ad hoc sampling……………………………………………….18 Survey of the wild honeybee population …………………………………………….……19 Robben Island ………………………………………………………………………….……20 Results August/October 1997 Survey ………………………………..……………………….……20 Subsequent surveys and ad hoc sampling……………………………………………….23 Survey of the wild honeybee population …………………………………………….……26 Robben Island ………………………………………………………………………….……27 iv Discussion …………………………………………………………………………………………….28 CHAPTER THREE: The impact of varroa mites in South Africa Introduction …………………………………………………………………………………………...33 Materials & Methods Size of varroa population in colonies of Apis mellifera scutellata and Apis mellifera Capensis ………………………………………………………………………….………….37 Direct Mortality Caused by Varroa ………………………………………………………..37 Varroa Monitoring in Commercial Honeybees …………………………………………..38 Effect on Pollination Efficiency …………………………………………………………....39 Secondary Diseases and Pests Tracheal Mites ………………………………………………………………….….42 Capensis Problem …………………………………………………………….…..42 Statistical Analysis ……………………………………………………………………….…43 Results Size of varroa population in colonies of Apis mellifera scutellata and Apis mellifera Capensis ………………………………………………………………………………….….43 Direct Mortality Caused by Varroa ………………………………………………………..44 Varroa Monitoring in Commercial Honeybees …………………………………………..47 Effect on Pollination Efficiency …………………………………………………………....54 Secondary Diseases and Pests Tracheal Mites ………………………………………………………………….….56 Capensis Problem ………………………………………………………………...56 Discussion ……………………………………………………………………….…………………....59 CHAPTER FOUR: Population dynamics of varroa mites in African honeybees. Introduction ……………………………………………………………………………………………65 Materials & Methods Varroa population growth in Apis mellifera scutellata and Apis mellifera capensis ….68 Brood cell infestation rates and mite reproduction rates in Apis mellifera capensis ...69 Statistical analysis …………………………………………………………………………..70 Results Varroa population growth in Apis mellifera scutellata and Apis mellifera capensis ….71 Brood cell infestation rates and mite reproduction rates in Apis mellifera capensis ...75 Discussion …………………………………………………………………………………………….79 CHAPTER FIVE: Tolerance to varroa mite in Cape honeybees Introduction ……………………………………………………………………………………………84 Materials & Methods v Tolerance in Wild honeybee populations ………………………………………………...93 Tolerance in commercial honeybee populations ………………………………………..94 Potential factors affecting tolerance to Varroa destructor in Cape honeybees Hygienic behaviour & Varroa Removal …………………………………………95 Aggression towards mites ………………………………………………………..96 Attractiveness of brood …………………………………………………………...96 Mite Fertility and Reproductive Fate …………………………………………….97 Post-capping period in Apis mellifera capensis ………………………………..98 Biocontrol by pseudoscorpions ………………………………………………….98 Results Tolerance in Wild honeybee populations ………………………………………………...99 Tolerance in commercial honeybee populations ………………………………………..99 Potential factors affecting tolerance to Varroa destructor in Cape honeybees Hygienic behaviour & Varroa Removal ………………………………………..103 Aggression towards mites ………………………………………………………105 Attractiveness of brood ………………………………………………………….105 Mite Fertility and Reproductive Fate …………………………………………..107 Post-capping period in Apis mellifera capensis ………………………………108 Biocontrol by pseudoscorpions ………………………………………………...108 Discussion …………………………………………………………………………………………...110 CHAPTER SIX: General Discussion ……………………………………………………………………….118 REFERENCES: ……………………………………………………………………………………………….125 APPENDIX I: ………………………………………………………………………………………………….147 APPENDIX II: …………………………………………………………………………………………………201 APPENDIX III: ………………………………………………………………………………………………...208 APPENDIX IV: ………………………………………………………………………………………………...239 vi ___________________________________________________________________________ LIST OF FIGURES ___________________________________________________________________________ CHAPTER ONE: Figure 1.1: The varroa mite Varroa destructor Anderson and Trueman (2000). Figure 1.2: Varroa mites are typically found between honeybee abdominal segments, but also on the thorax at high densities. Figure
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