Resource Management and Sustainability S c h o w a l t e INSECTS and Sustainability of r Ecosystem Services “As the most diverse taxon on earth, insects have complex roles in ecosystems and hence affect provisioning services, regulating services, cultural services and supporting services provided by ecosystems. This new synthesis enhances our understanding of the important functional roles of insects in ecosystem services, which will help us manage our natural resources in a better way to reach the goal of sustainability.” —Jung-Tai Chao, Taiwan Forestry Research Institute Timothy D. Schowalter “… the chapters of Schowalter's interesting new book range from insect ecology to the valuation and management of their benefits. … an impor- tant source for everybody interested in sustainable management of insects driving ecosystem functioning.” —Teja Tscharntke, Georg-August-University a “Dr. Schowalter brings forth many important topics within this realm as n d he ultimately leads the reader to a strong understanding of the important S interplay between humans, insects, and the shaping of environments u throughout the world. … well written and designed to allow the reader to s quickly grasp and understand even the most difficult concepts. I have little t a doubt that this will become a ‘must have’ reference for anyone interested i n insects and their role in the sustainability of ecosystem services.” a —Mathew A. Camper, Entomologist, Colorado State University b i l Features it y • Presents a complete theory written for a general audience o interested in insects and the sustainability of ecosystem services f E • Addresses the complexity of insect effects on our lives, our c resources, and ecosystem services on which we depend o • Provides recommendations for more rational approaches to s y “managing” insect populations s t • Challenges many traditional perceptions of the value of insects e m in our lives S e r v i c K15294 e s Half Title Page Series Page Title Page LOC Page Contents Series Preface ..........................................................................................................ix Preface ......................................................................................................................xi Author .....................................................................................................................xv 1 Introduction .....................................................................................................1 1.1 Ecosystem Services ...............................................................................3 1.1.1 Provisioning Services ..............................................................4 1.1.2 Cultural Services ......................................................................6 1.1.3 Supporting Services .................................................................6 1.1.4 Regulating Services .................................................................7 1.2 Ecosystem Characteristics ...................................................................8 1.3 Insects and Insect Control .................................................................12 1.4 Scope of This Book ..............................................................................15 References .......................................................................................................16 2 Humans versus Insects: The Good, the Bad, and the Ugly ..................23 2.1 Honey Bees (the “Good”) ...................................................................24 2.2 Other Insects Used as Food or Cosmetics .......................................26 2.3 Medical Uses ........................................................................................28 2.4 Industrial Uses ....................................................................................29 2.5 Cultural Icons ......................................................................................31 2.6 Indicators of Environmental Change ...............................................34 2.7 Nuisances (the “Bad”) .........................................................................35 2.8 Venomous Insects and Other Arthropods ......................................37 2.9 Vectors of Human and Livestock Diseases .....................................39 2.10 Crop Pests .............................................................................................42 2.11 Instruments of Warfare (the “Ugly”) ...............................................44 2.12 Summary ..............................................................................................45 References .......................................................................................................45 3 Insect Responses to Environmental Changes ........................................53 3.1 Adaptive Attributes ............................................................................53 3.2 Dispersal ...............................................................................................55 3.2.1 Life History Strategy .............................................................55 3.2.2 Crowding.................................................................................55 3.2.3 Nutritional Status ..................................................................56 3.2.4 Mechanism of Dispersal .......................................................57 3.2.5 Habitat and Resource Conditions ........................................59 3.3 Environmental Variation ....................................................................60 3.4 Disturbance ..........................................................................................63 © 2008 Taylor & Francis Group, LLC v vi Contents 3.4.1 Type ..........................................................................................64 3.4.2 Magnitude ...............................................................................64 3.4.3 Frequency ................................................................................64 3.4.4 Extent .......................................................................................66 3.5 Insect Responses .................................................................................67 3.5.1 Direct Effects of Abiotic Changes ........................................67 3.5.1.1 Temperature Extremes ..........................................67 3.5.1.2 Precipitation Extremes ...........................................68 3.5.1.3 Wind Speed and Water Flow ................................69 3.5.1.4 Air and Water Quality ...........................................69 3.5.2 Indirect Effects of Post-Disturbance Changes ...................70 3.5.2.1 Exposure ..................................................................71 3.5.2.2 Resource Abundance and Quality ......................71 3.5.2.3 Predator/Parasite Abundance and Foraging Activity ....................................................................80 3.6 Rate of Adaptation by Insects ...........................................................80 3.7 Summary ..............................................................................................83 References ......................................................................................................84 4 Changes in Insect Abundance and Distribution ...................................97 4.1 Population Fluctuation through Time .............................................98 4.2 Factors Affecting Population Size .....................................................106 4.2.1 Abiotic Factors ......................................................................107 4.2.2 Biotic Factors .........................................................................109 4.2.2.1 Resource Suitability .............................................109 4.2.2.2 Resource Availability ...........................................111 4.2.2.3 Predation ...............................................................113 4.3 Regulation of Population Size .........................................................114 4.4 Population Fluctuation in Space .....................................................117 4.4.1 Population Expansion .........................................................117 4.4.2 Metapopulation Dynamics .................................................122 4.4.3 Habitat Connectivity ...........................................................125 4.5 Summary ............................................................................................126 References .....................................................................................................127 5 How Do Ecosystems Provide Services? .................................................141 5.1 Ecosystem Structure and Function ................................................143 5.2 Interactions among Species .............................................................150 5.2.1 Direct Interactions ...............................................................150 5.2.2 Indirect Interactions ............................................................152 5.3 Climate Control .................................................................................154 5.4 Regulation of Ecosystem Conditions .............................................157 5.4.1 Evidence for Self-Regulation ..............................................158 5.4.2 Biodiversity ...........................................................................161 © 2008 Taylor & Francis Group, LLC Contents vii 5.4.3 Herbivory ..............................................................................163 5.4.4 Predation ...............................................................................166 5.5 Ecosystem Development ..................................................................169 5.6 Differences among Ecosystems ......................................................171 5.7 Predicting Change ............................................................................172 5.8 Summary ............................................................................................174 References .....................................................................................................175 6 Effects of Anthropogenic Changes and Management ........................189 6.1 How Humans Affect Ecosystems ...................................................190 6.1.1 Disturbances .........................................................................190 6.1.2 Global Change ......................................................................193 6.1.3 Pollution and Invasive Species ...........................................196 6.1.4 Ecosystem Fragmentation and Conversion ......................201 6.1.5 Induction of Insect Problems .............................................205 6.2 Insecticide Effects .............................................................................208 6.3 Alternative Control Options ............................................................214 6.4 Summary ............................................................................................216 References .....................................................................................................217 7 Effects of Insects on Ecosystem Services ..............................................233 7.1 Effects on Provisioning Services .....................................................237 7.2 Effects on Cultural Services .............................................................239 7.3 Effects on Supporting Services .......................................................241 7.3.1 Primary Production .............................................................241 7.3.2 Pollination .............................................................................244 7.3.3 Decomposition and Soil Formation...................................246 7.4 Effects on Regulating Services ........................................................253 7.4.1 Primary Production .............................................................253 7.4.2 Biological Control.................................................................254 7.4.3 Disease Cycles ......................................................................254 7.4.4 Nutrient Cycling ..................................................................256 7.4.5 Climate and Disturbance ....................................................258 7.5 Summary ............................................................................................259 References .....................................................................................................260 8 Valuation of Insect and Management Effects .......................................275 8.1 Valuation of Insect Effects on Provisioning Services ..................276 8.2 Valuation of Insect Effects on Cultural Services ..........................280 8.3 Valuation of Insect Effects on Supporting Services .....................281 8.4 Valuation of Insect Effects on Regulating Services ......................285 8.5 Valuation of Insect Management Strategies ..................................286 8.6 Summary ............................................................................................288 References .....................................................................................................289 © 2008 Taylor & Francis Group, LLC viii Contents 9 Conclusions and Recommendations .......................................................297 9.1 Why New Pest Management Approaches Are Needed ..............298 9.2 Management Goals ...........................................................................300 9.3 Deciding When to Control ...............................................................301 9.4 Greater Emphasis on Ecological Principles ...................................305 9.5 Use of a Broader Range of Tactics ...................................................307 9.5.1 Preventive Tactics .................................................................309 9.5.1.1 Protecting or Augmenting Natural Enemies ...309 9.5.1.2 Manipulating Habitat or Resource Suitability and Availability .................................309 9.5.2 Control Measures .................................................................312 9.5.3 Marketing Insects as Food..................................................315 9.6 Implementation of Integrated Pest Management Strategy .........315 9.7 Integrated Pest Management: A Case Study .................................317 9.8 Urban Responsibilities .....................................................................318 9.9 Concluding Comments ....................................................................322 References .....................................................................................................324 © 2008 Taylor & Francis Group, LLC