Disease Mitigation and Pathogenic Control in German and Common Wasps, Vespula Germanica and V Vulgaris , (Hymenoptera: V Espidae )
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Disease mitigation and pathogenic control in German and common wasps, Vespula germanica and V vulgaris , (Hymenoptera: V espidae ) By Stephen John Harcourt A thesis completed in partial fulfilment of the requirements for the degree of DOCTOR OF PHILOSOPHY at Lincoln University Canterbury, New Zealand 2002 Vespula vulgaris nest being examined during the colony hygiene experiment (see Chapter Six). 11 ABSTRACT Vespula germanica (F.) (German wasp) and V. vulgaris (L.) (common wasp) are major pests in New Zealand. They damage crops, have a significant impact on the biodiversity of native ecosystems and present a significant health hazard by stinging people and animals. Efforts to control them, both chemically and biologically, have not been as effective as initially hoped. This research contributes to a larger programme studying the potential of entomopathogenic microbes for the control of Gennan and common wasps in New Zealand. This study has demonstrated the potential and limitations which exist for control of wasps using microbial pathogens. Effective pathogens were identified and disease transmission quantified in bioassays. Behavioural and physiological adaptations of wasps were investigated as resistance mechanisms to disease., Three species of fungi (Beauveria bassiana, Aspergillus flavus and Metarhiziul1l anisop/iae) were identified as having potential to kill workers and larvae and severely disrupt the colony. In laboratory trials, fungi and bacteria were readily transmitted between workers and larvae and among workers. Workers and larvae, individually, possess few resistance mechanisms capable of suppressing a mass infection of pathogenic fungi. No tested bacteria were pathogenic to adult wasps. Although B. bassiana was extremely pathogenic in bioassays, behavioural adaptations of wasp workers and larvae restricted proliferation of fungi in a healthy nest. Anomalies in behaviour when removing sporulating or infectious cadavers indicate a 'window-of-opportunity' for the dissemination of disease if the fungus could achieve maturity. However, getting sufficient inoculum into the nest and achieving widespread sporulation to establish an epizootic will be a significant challenge. All of the bacteria tested were susceptible to antibiotics in larval saliva, and therefore not significantly pathogenic to wasps. Significant inhibitory effects of venom against A. flavus were identified, but no evidence of its use in the nest could be found. A more cryptic and infectious disease, such as a bacterium or microsporidian, holds the greatest potential for future control efforts. It is apparent that, in order to establish an infection capable of developing into an epizootic in wasp nests, a microbe needs to be gradually invasive, readily transmitted, a prolific replicator, cryptic and have little impact on host behaviour leading up to maturation. Limitations to the current use of pathogens for wasp control may be overcome by strain selection and development of delivery systems. Pathogens have the potential to complement and enhance existing biological control programmes, but much research is needed to realise this potential. Key words: Vespula vulgaris, Vespula germanica, Beauveria bassiana, Aspergillus flavus, Serratia marcescens, fungus, bacteria, bioassay, entomopathogen, mortality, venom, larval saliva, hygienic behaviour, necrophoria, necrophagia, trophallaxis. III ACKNOWLEDGEMENTS Firstly, I would like to acknowledge the sacrifice my family have made to allow me the lUxury of working on this thesis. It goes without saying that I could not have taken on such a project without their backing, but it has been difficult for you all. Mac, Ben and Cameron, in time, will not remember the hardships they have endured, but Nikki will probably never forget. For your understanding, I am grateful. I would like to thank my supervisors, Bruce Chapman and Dr. Eric Scott from Lincoln University and Dr. Richard Harris from Landcare Research for their guidance and encouragement throughout this research project. I would also like to thank the many science and support staff at Landcare Research (Lincoln and Nelson) and AgResearch who were a constant source of encouragement and enthusiasm concerning my research, especially Peter Read, Anne Rose, Dr. Travis Glare, Tracy Nelson, Dr. Trevor Jackson. Special mention must go to Dr. Ban·y Donovan who was an enduring source of inspiration about wasp research and provided me with innovative discussion to keep me on my scientific toes. Many thanks to the Applied Entomology and Pathology team (weeds and pests), my hosts at Landcare Research. Lindsay, Hugh, Lynley, Pauline, Helen, Jeremy, Tom (and Vanessa) you have been excellent colleagues and friends. I would also like to recognise the contribution made by staff from Crop and Food Research for the casual employment which made the difference financially, but particularly Dr. David Teulon and Marlon Stufkens and Damien Coup for being supportive of my research and for the great working environment and lasting friendships. For financial support and facilities I must thank the executive of Landcare Research, the trustees of the Frank Sydenham Trust and Lincoln University who made this research possible. Lastly J would like to thank Christophe Placet for his assistance with the contents of Appendix Three. A small, but infonnative, section of work that I was able to draw upon for much of my discussion. IV TABLE OF CONTENTS ABSTRACT .......................................................................................................................................... III ACKNOWLEDGEMENTS ...................................................................................................................... IV TABLE OF CONTENTS .......................................................................................................................... V LIST OF FIGURES .............................................................................................................................. VIII LIST OF TABLES .......................................................................................................................... , ..... XII LIST OF SYMBOLS AND ABBREVIATIONS .......................................................................................... XIV Chapter One General Introduction Background ......................................................................................................................................... I Vespid \vasps in New Zealand ............................................................................................................ 2 Micro-organisms in biological control ................................................................................................ 7 Study goals and objectives ................................................................................................................ 15 Chapter Two Adult Bioassays: Susceptibility of wasps to generalist entomopathogens Introduction ...................................................................................................................................... 16 Materials and methods ...................................................................................................................... 18 Results .............................................................................................................................................. 22 Discussion ......................................................................................................................................... 25 Chapter Three Transfer of pathogens between nestmates Introduction ...................................................................................................................................... 29 Materials and Methods ...................................................................................................................... 30 Results .............................................................................................................................................. 32 Discussion ......................................................................................................................................... 34 Chapter FOllr Anti-microbial properties of wasp exocrine secretions. venom and larval saliva Introduction ...................................................................................................................................... 39 Preliminary experiment~: Growth plate assays Introduction ...................................................................................................................................... 42 Materials and methods ...................................................................................................................... 43 Results .............................................................................................................................................. 44 Discussion ......................................................................................................................................... 47 V Fungicidal properties of venom: Germination assays Introduction ...................................................................................................................................... 51 Method and Materials ....................................................................................................................... 51 Results .............................................................................................................................................