Ecology of Invertebrate Diseases

<p>List of Contributors xvii</p>
<p>Preface xxi</p>
<p>Section I Introduction 1</p>
<p>1 General Concepts in the Ecology of Invertebrate Diseases 3<br />Ann E. Hajek and David I. Shapiro–Ilan</p>
<p>1.1 Introduction 3</p>
<p>1.1.1 What Is Disease? 4</p>
<p>1.1.2 Terminology and Measurements 5</p>
<p>1.1.2.1 Prevalence/Incidence 5</p>
<p>1.1.2.2 Pathogenicity/Virulence 5</p>
<p>1.1.2.3 Infection/Infectivity 6</p>
<p>1.1.2.4 Immunity 6</p>
<p>1.1.2.5 Transmission 7</p>
<p>1.1.2.6 Epizootic and Enzootic Diseases 7</p>
<p>1.1.2.7 Cycles of Infection 8</p>
<p>1.1.2.8 R0 and the Host Density Threshold 10</p>
<p>1.1.3 Factors Influencing the Ecology of Invertebrate Diseases 11</p>
<p>1.1.3.1 Host Range 12</p>
<p>1.2 Types of Studies 13</p>
<p>1.3 Why Study the Ecology of Invertebrate Diseases? 13</p>
<p>1.4 What this Book Covers 14</p>
<p>Acknowledgments 16</p>
<p>References 16</p>
<p>2 Methods for Studying the Ecology of Invertebrate Diseases and Pathogens 19<br />Raquel Campos–Herrera and Lawrence A. Lacey</p>
<p>2.1 Introduction 19</p>
<p>2.2 Traditional Methods for Studying Diseases 19</p>
<p>2.2.1 Sampling Goals 19</p>
<p>2.2.2 Sampling Regimes 20</p>
<p>2.2.3 Methodologies 20</p>
<p>2.2.3.1 Searching for Infected Insects Using General Entomological Sampling Methods 23</p>
<p>2.2.3.2 Selective Media 23</p>
<p>2.2.3.3 Extraction Methods 24</p>
<p>2.2.3.4 Airborne Spore Sampling 24</p>
<p>2.2.3.5 Insect Baiting 24</p>
<p>2.2.3.6 Dispersal of Entomopathogens: Mark Release Recapture Method 25</p>
<p>2.3 Molecular Tools to Assist in the Detection and Quantification of Pathogens and their Impact on the Host 25</p>
<p>2.3.1 Employment of Proteins: The Beginning of the Molecular Era in Invertebrate Pathology 26</p>
<p>2.3.2 Techniques Based on the Nucleic Acids: the Pre ]Omics Era 27</p>
<p>2.3.3 Advanced Techniques: qPCR, NGS, and the Arrival of the ]Omics Era 31</p>
<p>2.4 Traditional Versus Molecular Methods: Advantages and Limitations 33</p>
<p>2.5 Advancing the Frontiers of Ecology using Pathogens and Diseases 36</p>
<p>2.6 Conclusion38</p>
<p>Acknowledgments 38</p>
<p>References 38</p>
<p>Section II The Basics of Invertebrate Pathogen Ecology 49</p>
<p>3 The Pathogen Population 51<br />Leellen F. Solter and James J. Becnel</p>
<p>3.1 Introduction 51</p>
<p>3.2 Characteristics of Pathogens 51</p>
<p>3.2.1 Invasiveness and Infectivity 52</p>
<p>3.2.1.1 Routes of Entry 53</p>
<p>3.2.1.2 Establishment of Infection and Tissue Tropism 54</p>
<p>3.2.2 Pathogenicity, Virulence, and Pathogen Replication 55</p>
<p>3.2.2.1 Virulence Factors 57</p>
<p>3.2.2.2 Attenuation or Enhancement of Virulence 58</p>
<p>3.2.3 Latency 59</p>
<p>3.2.4 Obligate, Opportunistic, and Facultative Pathogens 60</p>
<p>3.2.5 Transmission 61</p>
<p>3.2.5.1 Horizontal Transmission 61</p>
<p>3.2.5.2 Vertical Transmission 62</p>
<p>3.2.5.3 Indirect Transmission 63</p>
<p>3.2.6 Genetic Variability and Potential for Coevolution with Hosts 64</p>
<p>3.2.6.1 Species and Strains 64</p>
<p>3.2.6.2 Host Specificity 65</p>
<p>3.3 Pathogen Effects on Host Development and Behavior 66</p>
<p>3.4 Pathogen Populations 67</p>
<p>3.4.1 Density ]Dependent Pathogens 67</p>
<p>3.4.2 Density ]Independent Pathogens 68</p>
<p>3.4.3 Pathogen Persistence in the Host Population 68</p>
<p>3.4.3.1 Chronic Infections and Vertical Transmission 69</p>
<p>3.4.3.2 Alternative and Alternate/Intermediate Hosts 69</p>
<p>3.4.3.3 Pathogen Survival in Cadavers and in Plant Tissues 70</p>
<p>3.4.3.4 Latency in Host Populations 70</p>
<p>3.4.4 Persistence of Pathogen Stages in the Environment 71</p>
<p>3.5 Dispersal and Spatial Distribution of Pathogens 72</p>
<p>3.5.1 Physical Factors: Wind and Water Dispersal 73</p>
<p>3.5.2 Biological Factors 73</p>
<p>3.5.3 Spatial Distribution 74</p>
<p>3.6 Pathogen Interactions 75</p>
<p>3.6.1 Interactions with other Biological Agents 75</p>
<p>3.6.2 Interactions with Pesticides and other Chemicals 77</p>
<p>3.6.3 Enhancing Factors 77</p>
<p>3.7 Conclusion 78</p>
<p>References 79</p>
<p>4 The Host Population 101<br />Louela A. Castrillo</p>
<p>4.1 Introduction 101</p>
<p>4.2 General Host Factors 103</p>
<p>4.2.1 Routes of Pathogen Acquisition 103</p>
<p>4.2.2 Insect Species, Life Stage, Age, and Gender 103</p>
<p>4.2.3 Population Density 104</p>
<p>4.3 Barriers to Microbial Infection 105</p>
<p>4.3.1 Insect Integument 105</p>
<p>4.3.2 Tracheae 107</p>
<p>4.3.3 Insect Gut 108</p>
<p>4.3.3.1 Peritrophic Membrane and Basal Lamina 108</p>
<p>4.3.3.2 Conditions in the Gut Lumen 108</p>
<p>4.3.3.3 Sloughing of Infected Epidermal Cells 109</p>
<p>4.4 Defenses against Microbial Infection 110</p>
<p>4.4.1 Innate Immune System 110</p>
<p>4.4.1.1 Constitutive Innate Immunity: Cellular Immunity 110</p>
<p>4.4.1.2 Constitutive Innate Immunity: Phenoloxidase 113</p>
<p>4.4.1.3 Induced Innate Immunity: Reactive Oxygen Species 113</p>
<p>4.4.1.4 Induced Innate Immunity: Antimicrobial Peptides 114</p>
<p>4.4.2 Microbiome ]Based Defenses 117</p>
<p>4.4.2.1 Gut Microbiota 117</p>
<p>4.4.2.2 Intracellular Symbionts 118</p>
<p>4.4.3 Behavioral Defenses 119</p>
<p>4.4.3.1 Avoidance/Evasion 120</p>
<p>4.4.3.2 Grooming and Hygienic Behaviors 121</p>
<p>4.4.3.3 Diet ]Based Prophylactic and Therapeutic Defenses 122</p>
<p>4.4.3.4 Thermoregulation 123</p>
<p>4.4.3.5 Deposition of Antimicrobial Compounds 123</p>
<p>4.5 Resistance via Priming 124</p>
<p>4.6 Conclusion 125</p>
<p>Acknowledgments 126</p>
<p>References 126</p>
<p>5 Abiotic Factors 143<br />Dana Ment, Ikkei Shikano and Itamar Glazer</p>
<p>5.1 Introduction 143</p>
<p>5.2 The Surviving Unit 143</p>
<p>5.2.1 Nematodes 143</p>
<p>5.2.2 Fungi 144</p>
<p>5.2.3 Viruses 145</p>
<p>5.2.4 Bacteria 146</p>
<p>5.3 Abiotic Factors Affecting Invertebrate Pathogens 146</p>
<p>5.3.1 Temperature 146</p>
<p>5.3.1.1 Nematodes 147</p>
<p>5.3.1.2 Fungi 148</p>
<p>5.3.1.3 Viruses 150</p>
<p>5.3.1.4 Bacteria 152</p>
<p>5.3.2 Moisture and Humidity 152</p>
<p>5.3.2.1 Nematodes 153</p>
<p>5.3.2.2 Fungi 154</p>
<p>5.3.2.3 Viruses 155</p>
<p>5.3.2.4 Bacteria 156</p>
<p>5.3.3 Ultraviolet Radiation 156</p>
<p>5.3.3.1 Nematodes 157</p>
<p>5.3.3.2 Fungi 157</p>
<p>5.3.3.3 Viruses 158</p>
<p>5.3.3.4 Bacteria 158</p>
<p>5.3.4 Chemical Inputs 159</p>
<p>5.3.4.1 Nematodes 159</p>
<p>5.3.4.2 Fungi 160</p>
<p>5.3.4.3 Viruses 160</p>
<p>5.3.4.4 Bacteria 161</p>
<p>5.3.5 Other Habitat Characteristics 162</p>
<p>5.3.5.1 Nematodes 162</p>
<p>5.3.5.2 Fungi 163</p>
<p>5.3.5.3 Viruses 164</p>
<p>5.3.5.4 Bacteria 165</p>
<p>5.4 Mechanisms of Survival 165</p>
<p>5.4.1 Nematodes 165</p>
<p>5.4.2 Fungi 166</p>
<p>5.4.3 Viruses 166</p>
<p>5.4.4 Bacteria 167</p>
<p>5.5 Conclusion 167</p>
<p>References 169</p>
<p>6 The Biotic Environment 187<br />Jenny S. Cory and Pauline S. Deschodt</p>
<p>6.1 Introduction 187</p>
<p>6.2 Tritrophic Interactions 188</p>
<p>6.2.1 Further Complexity 190</p>
<p>6.3 Pathogen Natural Enemy Interactions 191</p>
<p>6.3.1 Entomopathogen Entomopathogen Interactions 191</p>
<p>6.3.2 Entomopathogen Parasitoid Interactions 195</p>
<p>6.3.2.1 Effects of Pathogens on Parasitoids 195</p>
<p>6.3.2.2 Effects of Parasitoids on Pathogens 197</p>
<p>6.3.2.3 Population Level Effects 198</p>
<p>6.3.3 Pathogen Predator Interactions 199</p>
<p>6.3.4 Conclusion 200</p>
<p>6.4 Microbe– Mediated Defense 200</p>
<p>6.4.1 Heritable Symbionts 201</p>
<p>6.4.2 Do Gut Microflora Influence Pathogen Susceptibility? 202</p>
<p>6.4.3 Future Directions 204</p>
<p>6.5 Conclusion 204</p>
<p>Acknowledgments 204</p>
<p>References 205</p>
<p>Section III Ecology of Pathogen Groups 213</p>
<p>7 Viruses 215<br />Trevor Williams</p>
<p>7.1 Introduction 215</p>
<p>7.2 Diversity of Invertebrate Pathogenic Viruses 216</p>
<p>7.3 Distribution of Invertebrate Pathogenic Viruses 219</p>
<p>7.4 Key Aspects of Pathogen Ecology 220</p>
<p>7.5 Transmission 221</p>
<p>7.5.1 Horizontal Transmission 221</p>
<p>7.5.1.1 Estimating Horizontal Transmission 223</p>
<p>7.5.2 Vertical Transmission 223</p>
<p>7.6 Persistence 225</p>
<p>7.6.1 Persistence within the Host 225</p>
<p>7.6.2 Persistence Outside of the Host 226</p>
<p>7.6.2.1 Persistence on Plants 227</p>
<p>7.6.2.2 Persistence in Soil 229</p>
<p>7.6.2.3 Persistence in Water 230</p>
<p>7.7 Dispersal 231</p>
<p>7.7.1 Host ]Mediated Dispersal 231</p>
<p>7.7.2 Environmental Factors Involved in Dispersal 232</p>
<p>7.7.3 Biotic Factors that Assist the Dispersal of Viruses 233</p>
<p>7.7.3.1 Predators 233</p>
<p>7.7.3.2 Parasitoids 234</p>
<p>7.7.3.3 Other Organisms 234</p>
<p>7.7.4 Agricultural Practices that Affect Dispersal 234</p>
<p>7.7.5 Spatial Patterns of Dispersal 235</p>
<p>7.8 Genetic Diversity in Viruses 235</p>
<p>7.8.1 Genetic Diversity is Pervasive in Virus Populations 235</p>
<p>7.8.2 Genetic Diversity Favors Virus Survival 237</p>
<p>7.8.3 What Generates So Much Genetic Diversity? 238</p>
<p>7.8.4 How Is Genetic Diversity Transmitted? 239</p>
<p>7.9 Role of Host Behavior in Virus Ecology 240</p>
<p>7.9.1 Foraging Decisions: What and Where to Eat 240</p>
<p>7.9.2 The Risks of Cannibalism 241</p>
<p>7.9.3 Sexually Transmitted Viral Diseases 241</p>
<p>7.9.4 Ecological Consequences of Host Manipulation by Viruses 242</p>
<p>7.9.4.1 Molecular Basis for Host Manipulation 243</p>
<p>7.10 Dynamics of Viruses in Host Populations 244</p>
<p>7.10.1 Pathogenic Viruses Can Regulate Populations 244</p>
<p>7.10.2 Ecosystem Characteristics that Favor Virus Transmission 246</p>
<p>7.10.3 Climate Change and Insect Virus Population Dynamics 247</p>
<p>7.11 Influence of Abiotic Factors on Viruses 248</p>
<p>7.11.1 Effect of Ultraviolet Light on Viruses 248</p>
<p>7.11.2 Seasonal Effects on Viruses 250</p>
<p>7.11.3 Effect of Temperature on Viruses 250</p>
<p>7.11.4 Humidity, Moisture and Precipitation 251</p>
<p>7.11.5 Effect of pH on Viruses 251</p>
<p>7.12 Biotic Factors that Interact with Virus Populations 253</p>
<p>7.12.1 Plant Phenology, Structure, and Nutritional Value 253</p>
<p>7.12.2 Phytochemical Virus Interactions 253</p>
<p>7.12.3 Virus Interactions with Alternative Hosts 254</p>
<p>7.12.4 Competition and Facilitation in Virus Interactions with Other Organisms 255</p>
<p>7.12.4.1 Virus Interactions with Parasitoids 255</p>
<p>7.12.4.2 Virus Interactions with Other Pathogens 257</p>
<p>7.12.4.3 Virus Interactions with Microbiota 258</p>
<p>7.13 Conclusion 258</p>
<p>Acknowledgments 259</p>
<p>References 259</p>
<p>8 Bacteria 287<br />Trevor A. Jackson, Colin Berry and Maureen O Callaghan</p>
<p>8.1 Introduction 287</p>
<p>8.2 Bacterial Pathogens and Associations with Insects 288</p>
<p>8.3 Pathogenicity and Virulence 294</p>
<p>8.3.1 Pathogenicity 295</p>
<p>8.3.2 Virulence 299</p>
<p>8.4 Disease Transmission 300</p>
<p>8.5 Survival in the Environment 301</p>
<p>8.5.1 Soil 302</p>
<p>8.5.2 Aqueous Environments 304</p>
<p>8.5.3 On the Phylloplane and In Planta 304</p>
<p>8.6 Population Dynamics: Epizootics and Enzootics 305</p>
<p>8.7 Evolution 308</p>
<p>8.8 Ecology Guiding Use of Bacterial Entomopathogens in Microbial Control 309</p>
<p>8.9 Conclusion 311</p>
<p>References 312</p>
<p>9 Fungi 327<br />Ann E. Hajek and Nicolai V. Meyling</p>
<p>9.1 Introduction 327</p>
<p>9.1.1 Fungal Systematics and Taxonomy 328</p>
<p>9.1.2 Relevance of Fungal Systematics and Taxonomy in Ecology 330</p>
<p>9.2 Fungal Biology and Pathology 331</p>
<p>9.2.1 Biology and Pathology of Major Groups of Fungal Pathogens 331</p>
<p>9.2.1.1 Entomophthoromycotina, Entomophthorales 331</p>
<p>9.2.1.2 Ascomycota, Hypocreales 333</p>
<p>9.2.2 Distribution Patterns and Habitat Associations of Invertebrate Pathogenic Fungi 334</p>
<p>9.2.2.1 Patterns of Fungal Abundance and Distribution: Insights from the Use of Molecular Markers 335</p>
<p>9.2.3 Factors Governing Diversity Patterns of Fungal Pathogens 337</p>
<p>9.3 Dynamics of Fungal Pathogens 338</p>
<p>9.3.1 Disease Transmission 338</p>
<p>9.3.2 Fungal Dispersal 341</p>
<p>9.3.3 Fungal Environmental Survival and Persistence 341</p>
<p>9.3.4 Impacts on Host Population Densities over Space and Time 342</p>
<p>9.4 Interactions between Fungal Pathogens and Host Individuals 344</p>
<p>9.4.1 Host Responses to Fungal Pathogens to Prevent or Cure Infections 344</p>
<p>9.5 Impact of Abiotic Factors on Infected Hosts and Pathogen Inocula 347</p>
<p>9.6 Impact of Biotic Factors on Pathogenic Fungi 349</p>
<p>9.6.1 Endophytic and Rhizosphere Associations of Invertebrate Fungal Pathogens 349</p>
<p>9.6.1.1 Natural Occurrence and Distribution of Invertebrate Pathogenic Fungi as Plant Associates 350</p>
<p>9.6.1.2 Experimental Inoculations of Plants with Entomopathogenic Fungi 351</p>
<p>9.6.1.3 Direct and Indirect Fungal Interactions with Insects and Plants 352</p>
<p>9.6.2 Interactions between Host Symbionts and Fungal Pathogens 353</p>
<p>9.6.3 Interactions between Fungal Pathogens and Other Natural Enemies 354</p>
<p>9.6.3.1 Interactions among Co ]infecting Pathogens 355</p>
<p>9.6.3.2 Interactions of Fungal Pathogens with Parasitoids and Predators 356</p>
<p>9.6.4 Mycoparasitism of Fungal Pathogens 357</p>
<p>9.7 Use of Pathogenic Fungi for Biological Control of Invertebrates 358</p>
<p>9.8 Conclusion 361</p>
<p>Acknowledgments 361</p>
<p>References 362</p>
<p>10 Microsporidia 379<br />Gernot Hoch and Leellen F. Solter</p>
<p>10.1 Introduction 379</p>
<p>10.1.1 Mechanisms of Infection 380</p>
<p>10.1.2 Microsporidian Life Cycles 381</p>
<p>10.1.3 Pathology 381</p>
<p>10.2 Host Population 383</p>
<p>10.2.1 Susceptibility to Microsporidiosis 383</p>
<p>10.2.2 Immune Response 383</p>
<p>10.2.3 Behavioral Response 384</p>
<p>10.3 Pathogen Population 385</p>
<p>10.3.1 Virulence of Microsporidian Pathogens 385</p>
<p>10.3.2 Host Specificity 386</p>
<p>10.3.2.1 Physiological vs. Ecological Host Specificity 386</p>
<p>10.3.2.2 Host Range 386</p>
<p>10.3.2.3 Alternate Hosts 387</p>
<p>10.3.2.4 Microsporidia Crossing the Invertebrate Vertebrate Barrier 387</p>
<p>10.3.3 Persistence in the Environment 388</p>
<p>10.4 Transmission 390</p>
<p>10.4.1 Horizontal Transmission 390</p>
<p>10.4.1.1 Transmission from Living Hosts 390</p>
<p>10.4.1.2 Transmission after Host Death 392</p>
<p>10.4.1.3 Transmission by Parasitoid Vectors 393</p>
<p>10.4.1.4 Effects of Host Development and Host Microsporidia Interactions on Transmission 393</p>
<p>10.4.2 Vertical Transmission 394</p>
<p>10.5 Epizootiology 397</p>
<p>10.5.1 Microsporidian Prevalence in Invertebrate Populations and Impact on Host Populations 397</p>
<p>10.5.2 Microsporidia in Cultured Insects 398</p>
<p>10.5.3 Microsporidia as Potential Biological Control Agents 399</p>
<p>References 400</p>
<p>11 Nematodes 415<br />David I. Shapiro–Ilan, Ivan Hiltpold and Edwin E. Lewis</p>
<p>11.1 Introduction 415</p>
<p>11.1.1 Diversity and Life Histories 415</p>
<p>11.1.2 EPN Distribution 420</p>
<p>11.2 Transmission 421</p>
<p>11.3 Host Population 421</p>
<p>11.4 Pathogen Population 422</p>
<p>11.4.1 Pathogenicity and Virulence 422</p>
<p>11.4.2 Persistence and Recycling 423</p>
<p>11.4.3 Dispersal and Foraging Behavior 423</p>
<p>11.5 Abiotic Environmental Factors 424</p>
<p>11.5.1 Soil Moisture 424</p>
<p>11.5.2 Soil Temperature 425</p>
<p>11.5.3 Soil Characteristics and Chemistry 425</p>
<p>11.5.4 Ultraviolet Light 426</p>
<p>11.6 Biotic Interactions 426</p>
<p>11.6.1 Interactions with Predators and Pathogens, Including Intraguild Competition 426</p>
<p>11.6.2 Cues Used in Host ]Finding and Navigation 427</p>
<p>11.6.3 Tri ]trophic Interactions (Plant, Insect, Nematode) 427</p>
<p>11.7 Applied Ecology and Aspects in Microbial Control 427</p>
<p>11.7.1 Production, Formulation, and Application 427</p>
<p>11.7.2 Approaches to Microbial Control 428</p>
<p>11.8 Conclusion 430</p>
<p>References 431</p>
<p>Section IV Applied Ecology of Invertebrate Pathogens 441</p>
<p>12 Modeling Insect Epizootics and their Population–Level Consequences 443<br />Bret D. Elderd</p>
<p>12.1 Introduction 443</p>
<p>12.2 The Pathogen and its Hosts 445</p>
<p>12.3 Modeling Disease Transmission: A Single Epizootic 447</p>
<p>12.3.1 Phenomenological and Mechanistic Models 448</p>
<p>12.4 Fitting Models to Data 450</p>
<p>12.4.1 Akaike Information Criterion 451</p>
<p>12.4.2 An Example of the AIC in Action 452</p>
<p>12.5 A Bayesian Approach 453</p>
<p>12.5.1 Fitting a Bayesian Model 454</p>
<p>12.5.2 An Example of the WAIC in Action 456</p>
<p>12.6 Long–Term Dynamics 457</p>
<p>12.6.1 Long ]Term Dynamics: Confronting Models with Data 458</p>
<p>12.6.2 Time ]Series Diagnostics 459</p>
<p>12.7 Modifying and Applying the Model 462</p>
<p>12.8 Conclusion 463</p>
<p>Acknowledgments 463</p>
<p>References 463</p>
<p>13 Leveraging the Ecology of Invertebrate Pathogens in Microbial Control 469<br />Surendra K. Dara, Tarryn A. Goble and David I. Shapiro–Ilan</p>
<p>13.1 Basics of Microbial Control and Approaches 469</p>
<p>13.1.1 Classical Microbial Control 469</p>
<p>13.1.2 Inoculative Release 471</p>
<p>13.1.3 Inundative Release 472</p>
<p>13.1.4 Conservation/Environmental Manipulation 472</p>
<p>13.2 Ecological Considerations 472</p>
<p>13.2.1 Host Specificity 472</p>
<p>13.2.2 Dispersal Ability 473</p>
<p>13.2.3 Virulence 473</p>
<p>13.2.4 Pathogen Density 473</p>
<p>13.2.5 Host ]Related Factors 474</p>
<p>13.2.6 Transmission 474</p>
<p>13.2.7 Environmental Persistence 475</p>
<p>13.3 Methods to Improve Microbial Control 476</p>
<p>13.3.1 Improving the Organism as a Microbial Control Agent 476</p>
<p>13.3.2 Improving Production Methods 477</p>
<p>13.3.3 Improving Formulation and Application Technologies 478</p>
<p>13.3.4 Improving the Environment 479</p>
<p>13.4 Incorporating Microbial Control into Integrated Pest–Management Systems 480</p>
<p>13.4.1 Regulatory Issues 480</p>
<p>13.4.2 Standalone vs. Integrated Approaches 481</p>
<p>13.4.3 Case Studies 481</p>
<p>13.4.3.1 Orchard Crops 482</p>
<p>13.4.3.2 Row Crops 482</p>
<p>13.4.3.3 Forests 483</p>
<p>13.4.3.4 Greenhouses 483</p>
<p>13.5 Conclusion 484</p>
<p>References 484</p>
<p>14 Prevention and Management of Diseases in Terrestrial Invertebrates 495<br />J&oslash;rgen Eilenberg and Annette Bruun Jensen</p>
<p>14.1 Introduction 495</p>
<p>14.1.1 Types of Production Facilities 496</p>
<p>14.1.2 Transmission of Insect Diseases in Production Facilities 499</p>
<p>14.2 Major uses of Insects and Mites in the Production and Transmission of Insect Pathogens within Production Systems 500</p>
<p>14.2.1 Pollination and Honey Production 500</p>
<p>14.2.2 Silk Production 507</p>
<p>14.2.3 Biological Control 509</p>
<p>14.2.4 Production of Insects for Food and Feed 512</p>
<p>14.3 Status of Diagnostic Services 516</p>
<p>14.4 Ensuring Production of Healthy Insects 516</p>
<p>14.5 Conclusion 519</p>
<p>Acknowledgments 519</p>
<p>References 519</p>
<p>15 Prevention and Management of Infectious Diseases in Aquatic Invertebrates 527<br />Jeffrey D. Shields</p>
<p>15.1 Scope 527</p>
<p>15.1.1 Myriad Pathogens Infect Aquatic Invertebrates 527</p>
<p>15.1.2 Overview of Disease Issues in Assessing Epidemics in Aquatic Invertebrates 531</p>
<p>15.2 Oyster Diseases 539</p>
<p>15.3 Crustacean Diseases 543</p>
<p>15.3.1 Outbreaks in Shrimp Aquaculture 544</p>
<p>15.3.2 Disease Management in Shrimp Aquaculture 548</p>
<p>15.3.2.1 Switching Species and Specific Pathogen ]Free (SPF) Stocks 549</p>
<p>15.3.2.2 Surveillance 550</p>
<p>15.3.2.3 Development of Vaccines 551</p>
<p>15.3.2.4 Ecological and Biological Control 551</p>
<p>15.3.3 Crayfish and Krebspest 552</p>
<p>15.3.4 Disease Emergence in Culture of the Chinese Mitten Crab 553</p>
<p>15.4 Crustacean Fisheries 554</p>
<p>15.4.1 Snow Crabs and Bitter Crab Disease 555</p>
<p>15.4.2 American Lobster and Epizootic Shell Disease 556</p>
<p>15.4.3 Spiny Lobsters and PaV1 559</p>
<p>15.5 Agencies for Disease Management 560</p>
<p>15.6 Conclusion 563</p>
<p>Acknowledgments 563</p>
<p>References 563</p>
<p>16 Ecology of Emerging Infectious Diseases of Invertebrates 587<br />Colleen A. Burge, Amanda Shore–Maggio and Natalie D. Rivlin</p>
<p>16.1 Introduction 587</p>
<p>16.2 Host Pathogen Relationships and Anthropogenic Change 593</p>
<p>16.2.1 Ecological Context of Invertebrate Host Pathogen Relationships 593</p>
<p>16.2.2 Anthropogenic Change and Disease Emergence 594</p>
<p>16.2.2.1 Host Factors 595</p>
<p>16.2.2.2 Pathogen Factors 595</p>
<p>16.2.2.3 Environment Factors 595</p>
<p>16.3 Case Studies of Invertebrate Disease Emergence 596</p>
<p>16.3.1 Molluscan Herpesvirus Infections of Bivalves 597</p>
<p>16.3.1.1 OsHV ]1 Infections of Pacific Oysters and Other Bivalves 598</p>
<p>16.3.2 Acute Hepatopancreatic Necrosis Disease of Shrimp 601</p>
<p>16.3.3 Emerging Densoviruses of Arthropods and Echinoderms 603</p>
<p>16.3.3.1 Acheta domesticus Densovirus 603</p>
<p>16.3.3.2 Sea Star ]Associated Densovirus 604</p>
<p>16.3.4 Emerging Pathogens of Pollinators 605</p>
<p>16.3.4.1 Varroa destructor and Deformed Wing Virus 606</p>
<p>16.3.4.2 Spillover and spread of Nosema ceranae 607</p>
<p>16.3.4.3 Multi ]stressors, Bee Mortalities and Control Measures 608</p>
<p>16.3.5 Emergent Coral Diseases 608</p>
<p>16.3.5.1 Black Band Disease 609</p>
<p>16.3.5.2 Acroporid Serratosis 610</p>
<p>16.3.5.3 Problems Facing Coral Disease Investigations 610</p>
<p>16.4 Conclusion 611</p>
<p>Acknowledgments 612</p>
<p>References 612</p>
<p>17 Conclusions and Future Directions 627<br />David Shapiro–Ilan and Ann E. Hajek</p>
<p>17.1 The Increasing Urgency of the Study of Invertebrate Pathogen Ecology 627</p>
<p>17.1.1 Food Security and the Role of Microbial Control 627</p>
<p>17.1.2 Conservation of Beneficial Organisms 628</p>
<p>17.2 The Future for Invasive and Native Invertebrate Pathogens 629</p>
<p>17.3 New Directions and Novel Tools for Studying Invertebrate Ecology 630</p>
<p>17.3.1 Molecular Tools 630</p>
<p>17.3.2 Chemical Ecology and Signaling 631</p>
<p>17.3.3 Exploring Other Novel Biotic Associations 632</p>
<p>17.3.4 Interdisciplinary Studies 633</p>
<p>References 634</p>
<p>Index 637</p>
<p>&nbsp;</p>