September 2007 Summary
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The Ecological and Evolutionary Dynamics of Multiple Natural Enemies Edward Owen Jones Submitted for the degree of Doctor of Philosophy Department of Animal and Plant Sciences University of Sheffield September 2007 Summary Natural organisms are infected by many different parasites, and how these parasites interact with each other has important ecological and evolutionary implications. This study is to understand how organisms with different transmission strategies effect the evolution of each other. Parasites that transmit vertically benefit from the host having a high reproductive rate whilst hosts that transmit horizontally are not affected by host reproduction. Some horizontal parasites may reduce host fecundity to benefit their own replication within the host. Therefore there will be clear conflict between vertically and horizontally transmitted parasites. The evolution of traits such as vertically transmitted `protection' and `sabotage' are likely to be costly in terms of other life history traits. In the presence of such `trade-offs, ' the parasite population will evolve towards the evolutionarily stable strategy (ESS) that balances the costs and benefits of these strategies in response to a horizontally transmitted parasite. The ecology of the interaction of vertically and horizontally transmitting parasites is also examined. Further to this, field data is used to construct a model of two horizontally transmitting parasites, a pathogen and a parasitoid that occur on the Orkney Isles. We demonstrate how each of them, and both in combination, may have a large effect on resultant dynamics of winter moth, Operophtera brumata, populations. `A little nonsensenow and then is relished by the wisest men' Roald Dahl Contents Acknowledgements iv .................................................................................................. Declaration V ............................................................................................................... 1. Introduction 1 .......................................................................................................... 1.1 Ecological Outline: Host interactions 2 and multiple enemy ........................................... 1.1.1 Ecology 2 of multiple enemy systems ..................................................................... 1.1.2 Models 4 of multiple enemy systems ..................................................................... 1.1.3 The Model The Winter Moth Orkney 7 system: on ............................................... 1.1.4 Natural Enemies: Parasitoids, Pathogens Predators 8 and .................................... 1.2 Evolutionary Outline: Host Multiple Enemy Interactions 11 and .................................... 1.2.1 Evolution Parasites 11 of ......................................................................................... 1.2.2 Models Evolution Parasites 12 of the of ................................................................. 1.2.3 Models Evolution Multiple Parasites 12 of the of ............................................... 1.3 Theoretical Approach 14 .................................................................................................... 1.3.1 Population Dynamics 14 ......................................................................................... 1.3.2 Winter dynamics 15 moth ........................................................................................ 1.3.3 Evolutionarily Stable Strategies Adaptive Dynamics 15 and ............................... 1-4 Thesis 17 outline ................................................................................................................. 2. Interference Persistence Vertically Transmitted Parasites 19 and the of ..................... 2.1 Introduction 20 .................................................................................................................... 2.2 Models Analysis 22 and ...................................................................................................... 2.3 Discussion 33 ....................................................................................................................... 3. The Evolutionary implications of conflict between parasites with different transmission 36 modes .................................................................................................. 3.1 Introduction 37 .................................................................................................................... 3.2 The Model 39 ...................................................................................................................... 3.2.1 Evolution the horizontally transmitted 40 of parasite ............................................ 3.2.2 Evolution 44 of the vertically transmitted parasite ................................................ 3.2.4 Coevolution of both the horizontally and vertically transmitted parasites...... 50 3.3 Discussion 52 ....................................................................................................................... 4. The direct between 57 evolution of protection and sabotage: conflict parasites......... 4.1 Introduction 58 .................................................................................................................... 4.2 The Evolutionary Dynamics Protection 60 of ................................................................... 4.2.1 The in transmitted 61 evolution of protection vertically parasites ........................ 4.2.2 Evolution of a horizontally transmittedparasite in the presenceof a protecting 63 vertically transmitted parasite...................................................................................... 4.2.3 Coevolution of a protecting vertically transmitted parasite and horizontally 65 transmitted parasite...................................................................................................... 4.3 The dynamics `Sabotage' 68 evolutionary of .................................................................... 4.3.1 Evolution of the horizontally transmitting parasitein the presenceof a transmitting 69 sabotaging vertically parasite ................................................................. 4.3.2 Evolution of sabotage in a vertically transmitted parasite in the presence of a horizontally transmitted 71 castrating parasite ................................................................ 4.3.3 How do the horizontally transmitted parasite and the vertically transmitted 7 72 parasite coevolve ........................................................................................................ 4.4 Discussion 74 ....................................................................................................................... 5. The its Orkney 78 winter moth and natural enemies on ............................................. 5.1 Introduction 79 .................................................................................................................... 5.2 Methods 82 .......................................................................................................................... 5.2.1 Larval Collection 82 ................................................................................................ 5.2.2 Stage Dependency 82 ............................................................................................... 5.2.3 Bioassay 84 ............................................................................................................... 5.2.4 Statistical 85 analysis ............................................................................................... 5.3 Results 85 ............................................................................................................................. 5.3.1 Field 85 collection .................................................................................................... 5.3.2 Stage 93 structure..................................................................................................... 5.3.3 Bioassay 103 ............................................................................................................. 5.4 Discussion 106 ..................................................................................................................... 6. The its Orkney: A Model 113 winter moth and natural enemies on ........................... 6.1 Introduction 114 .................................................................................................................. 6.2 Host Model 117 ................................................................................................................... 6.3 Host NPV 120 and model ................................................................................................... 6.3.1 Host-pathogen dynamics 123 .................................................................................. 6.4 Parasitoid Host Model 131 and .......................................................................................... 6.4.1 Host-parasitoid dynamics 133 ................................................................................. 6.5 Host-Pathogen-Parasitoid Model 141 ................................................................................ 6.5.1 Host-pathogen-parasitoid dynamics 141 ................................................................ 6.6 Discussion 144 ..................................................................................................................... 7. Discussion 151 chapter............................................................................................