Insect Biodiversity

Insect Biodiversity

Science and Society

Second Edition

Volume I

Edited by

Robert G. FoottitAgriculture and Agri-Food CanadaOttawaOntarioCanada

Peter H. AdlerClemson UniversityClemsonSouth CarolinaUSA

This edition first published 2017 © 2017 John Wiley & Sons

First edition published 2009 by John Wiley & Sons Ltd

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Library of Congress Cataloging-in-Publication Data applied for.

ISBN: 9781118945537

Cover Design: WileyCover Image: A sampling of beetle diversity in the Palearctic Region.Images by Kirill Makarov

Set in 10/12pt, WarnockPro by SPi Global, Chennai, India

10 9 8 7 6 5 4 3 2 1

v

1 Introduction 1

2 The Importance of Insects 9

Part I Insect Biodiversity: Regional Examples 45

3 Insect Biodiversity in the Nearctic Region 47

4 Amazonian Rainforests and Their Richness and Abundance of Terrestrial Arthropods on the Edge of Extinction: Abiotic–Biotic Players in the Critical Zone 65

5 Insect Biodiversity in the Afrotropical Region 93

6 Biodiversity of Australasian Insects 111

7 Insect Biodiversity in the Palearctic Region 141

Part II Insect Biodiversity: Taxon Examples 203

8 Biodiversity of Aquatic Insects 205

9 Biodiversity of Diptera 229

10 Biodiversity of Heteroptera 279

11 Biodiversity of Coleoptera 337

12 Biodiversity of Hymenoptera 419

13 Diversity and Significance of Lepidoptera: A Phylogenetic Perspective 463

Part III Insect Biodiversity: Tools and Approaches 497

14 The Science of Insect Taxonomy: Prospects and Needs 499

Brief Table of Contents

Brief Table of Contentsvi

15 Insect Species – Concepts and Practice 527

16 Molecular Dimensions of Insect Taxonomy in the Genomics Era 547

17 DNA Barcodes and Insect Biodiversity 575

18 Insect Biodiversity Informatics 593

19 Parasitoid Biodiversity and Insect Pest Management 603

20 The Taxonomy of Crop Pests: The Aphids 627

21 Adventive (Non-Native) Insects and the Consequences for Science and Society of Species that Become Invasive 641

22 Biodiversity of Blood-sucking Flies: Implications for Humanity 713

23 Reconciling Ethical and Scientific Issues for Insect Conservation 747

24 Taxonomy and Management of Insect Biodiversity 767

25 Insect Biodiversity – Millions and Millions 783

vii

List of Contributors xix Foreword, Second Edition xxiii Preface, First Edition xxvii Preface, Second Edition xxix Acknowledgements xxxi

1 Introduction 1Peter H. Adler and Robert G. Foottit

References 5

2 The Importance of Insects 9Geoffrey G. E. Scudder

2.1 Diversity 92.2 Ecological Role 102.3 Effects on Natural Resources, Agriculture, and Human Health 132.4 Insects and Advances in Science 142.4.1 Biomechanics 152.4.2 Genetics 162.4.3 Developmental Biology 162.4.4 Evolution 182.4.5 Physiology 192.4.6 Ecology 202.4.7 Paleolimnology and Climate Change 222.5 Insects and the Public 23 References 25

Part I Insect Biodiversity: Regional Examples 45

3 Insect Biodiversity in the Nearctic Region 47Hugh V. Danks and Andrew B. T. Smith

3.1 Influence of Insect Biodiversity on Society in the Nearctic Region 493.2 Insect Conservation 503.3 Species Diversity and the State of Knowledge 533.3.1 Assembling the Data 533.3.2 Synopsis of Biodiversity 54

Detailed Table of Contents

Detailed Table of Contentsviii

3.3.3 State of Knowledge 553.4 Variations in Biodiversity 563.4.1 Regional Variation 573.4.2 Habitats 583.5 Conclusions and Needs 58 Acknowledgments 60 References 60

4 Amazonian Rainforests and Their Richness and Abundance of Terrestrial Arthropods on the Edge of Extinction: Abiotic–Biotic Players in the Critical Zone 65Terry L. Erwin, Laura S. Zamorano and Christy J. Geraci

4.1 The Climatic Setting and Critical Zone Establishment 694.2 Characterization of Typical Lowland Rainforest Composition in the Western Basin 714.3 Sampling Arthropod Biodiversity in Amazonian Forests 734.4 Richness of Various Lineages and Guilds 794.5 General Patterns 794.6 Morphospecies Richness to Biodiversity 804.7 Beetles: Life Attributes Have Led to Contemporary Hyperdiversity 834.8 Summary and Guide to Future Research, or “Taking a Small Step into the Biodiversity

Vortex” 85 Acknowledgments 86 References 86

5 Insect Biodiversity in the Afrotropical Region 93Clarke H. Scholtz and Mervyn W. Mansell

5.1 What Do We Know about Afrotropical Insects? 955.2 An Information-Management Program 955.3 The Role of Insects in Ecosystem Processes and as Indicators of Environmental

Quality – Dung Beetles as a Case Study 985.3.1 Dung Beetles as Indicators of Regional Biodiversity 995.3.2 Dung Beetles as Indicators of Habitat Transformation 1005.4 Africa-Wide Pests and Training Appropriate Taxonomists – Fruit Flies as

a Case Study 1005.4.1 Invasive Species of Concern in Africa 1015.4.2 African Indigenous Fruit Flies of Economic Importance 1025.5 Sentinel Groups 1035.5.1 Neuroptera 1035.5.2 Dung Beetles (Coleoptera: Scarabaeidae: Scarabaeinae) 1045.6 Conclusions 105 References 107

6 Biodiversity of Australasian Insects 111Peter S. Cranston

6.1 Australasia – The Locale 1116.2 Some Highlights of Australasian Insect Biodiversity 1126.2.1 The Lord Howe Island Stick Insect 1146.2.2 Australasian Birdwing Conservation 115

Detailed Table of Contents ix

6.3 Drowning by Numbers? How Many Insect Species are in Australasia? 1166.3.1 Australia 1166.3.2 New Zealand (Aotearoa), Chatham Islands, and Subantarctic Islands 1176.3.3 New Guinea 1186.3.4 New Caledonia and the West Pacific 1186.4 Australasian Insect Biodiversity – Overview and Special Elements 1186.4.1 Australia 1186.4.2 New Zealand 1216.4.3 New Caledonia, New Guinea, and Melanesia 1216.5 Threatening Processes to Australasian Insect Biodiversity 1236.5.1 Land Clearance and Alteration 1236.5.2 Introduced Animals 1236.5.3 Climate Change 1266.6 Australasian Biodiversity Conservation 1276.7 Conclusion 129 References 129

7 Insect Biodiversity in the Palearctic Region 141Boris A. Korotyaev, Alexander S. Konstantinov and Mark G. Volkovitsh

7.1 Preface: Societal Importance of Biodiversity in the Palearctic Region 1417.2 Introduction 1447.3 Geographic Position, Climate, and Zonality 1447.4 General Features of Palearctic Insect Biodiversity 1487.5 Biodiversity of Some Insect Groups in the Palearctic 1537.6 Biodiversity of Insect Herbivores 1587.7 Boundaries and Insect Biodiversity 1627.8 Local Biodiversity 1647.9 Insect Biodiversity and Habitats 1667.10 Insect Biodiversity and the Mountains 1697.11 Temporal Changes in Insect Biodiversity 1717.12 Insect Diversity in Major Biogeographical Divisions of the Palearctic 1727.12.1 Arctic (Circumpolar Tundra) Region 1737.12.2 Forest Regions 1747.12.3 Taiga 1767.12.4 Nemoral European and Stenopean Forests 1777.12.5 Hesperian and Orthrian Evergreen Forests 1797.12.6 Steppe (Scythian) Region 1807.12.7 Desert (Sethian) Region 183 Acknowledgments 187 References 189

Part II Insect Biodiversity: Taxon Examples 203

8 Biodiversity of Aquatic Insects 205John C. Morse

8.1 Overview of Taxa 2068.1.1 Mayflies (Ephemeroptera) 206

Detailed Table of Contentsx

8.1.2 Dragonflies and Damselflies (Odonata) 2078.1.3 Stoneflies (Plecoptera) 2078.1.4 Cockroaches (Blattodea) 2088.1.5 Grasshoppers and Crickets (Orthoptera) 2088.1.6 Earwigs (Dermaptera) 2088.1.7 Lice (Phthiraptera) 2088.1.8 Bugs (Hemiptera) 2088.1.9 Wasps (Hymenoptera) 2098.1.10 Hellgrammites and Alderflies (Megaloptera) 2098.1.11 Nerve‐winged Insects (Neuroptera) 2098.1.12 Scorpionflies (Mecoptera) 2108.1.13 Beetles (Coleoptera) 2108.1.14 Caddisflies (Trichoptera) 2108.1.15 Moths (Lepidoptera) 2118.1.16 Flies (Diptera) 2118.2 Species Numbers 2128.3 Societal Benefits and Risks 2148.3.1 Societal Benefits of Aquatic Insect Diversity in Food Webs 2148.3.2 Societal Benefits of Aquatic Insect Diversity in Biomonitoring 2158.3.3 Societal Benefits of Aquatic Insect Diversity in Fishing 2178.3.4 Societal Benefits of Aquatic Insect Diversity in Control of Noxious Weeds 2178.3.5 Societal Risks of Aquatic Insects 2178.4 Biodiversity Concerns for Aquatic Insects 2188.4.1 Threats to Freshwater Species of Insects 2188.4.2 Need for Biodiversity Discovery and Description of Aquatic Insects 2198.4.3 Need to Refine Definitions of Species of Aquatic Insects 2198.4.4 Need for New Generation of Aquatic Entomologists 219 References 220

9 Biodiversity of Diptera 229Gregory W. Courtney, Thomas Pape, Jeffrey H. Skevington and Bradley J. Sinclair

9.1 Overview of Taxa 2399.1.1 Lower Diptera 2399.1.2 Brachycera 2419.1.2.1 Lower Brachycera 2419.1.2.2 Empidoidea 2429.1.2.3 Lower Cyclorrhapha 2429.1.2.4 Non-calyptrate Schizophora 2439.1.2.5 Calyptratae 2459.2 Societal Importance 2469.2.1 Diptera as Plant Pests (Agriculture, Silviculture, and Floriculture) 2469.2.2 Medical and Veterinary Importance 2479.2.2.1 Disease transmission 2479.2.2.2 Myiasis 2489.2.3 Invasive Alien Diptera 2499.2.4 Diptera as a General Nuisance 249

Detailed Table of Contents xi

9.2.5 Diptera in Biological Control 2509.2.6 Pollination 2519.2.7 Other Ecological Services (Scavengers and Decomposers) 2529.3 Diptera of Forensic, Medicolegal, and Medical Importance 2539.4 Diptera as Model Organisms and Research Tools 2539.4.1 Physiology and Genetics 2539.4.2 Technology 2549.5 Diptera in Conservation 2549.5.1 Bioindicators 2549.5.2 Vanishing Species 2559.6 Diptera as Part of Our Cultural Legacy 256 References 257

10 Biodiversity of Heteroptera 279Thomas J. Henry

10.1 Overview of the Heteroptera 28010.1.1 Euheteroptera 28510.1.1.1 Infraorder Enicocephalomorpha 28510.1.1.2 Infraorder Dipsocoromorpha 28510.1.2 Neoheteroptera 28710.1.2.1 Infraorder Gerromorpha 28710.1.3 Panheteroptera 28810.1.3.1 Infraorder Nepomorpha 28810.1.3.2 Infraorder Leptopodomorpha 29010.1.3.3 Infraorder Cimicomorpha 29010.1.3.4 Infraorder Pentatomomorpha 30110.2 The Importance of Heteropteran Biodiversity 311 Acknowledgments 313 References 313

11 Biodiversity of Coleoptera 337Patrice Bouchard, Andrew B. T. Smith, Hume Douglas, Matthew L. Gimmel, Adam J. Brunke and Kojun Kanda

11.1 Overview of Extant Taxa 34411.1.1 Suborders Archostemata and Myxophaga 34411.1.2 Suborder Adephaga 34611.1.3 Suborder Polyphaga 34711.1.3.1 Series Staphyliniformia 34711.1.3.2 Series Scarabaeiformia 34911.1.3.3 Series Elateriformia 35011.1.3.4 Series Derodontiformia 35111.1.3.5 Series Bostrichiformia 35211.1.3.6 Series Cucujiformia 35211.2 Overview of Fossil Taxa 35711.3 Societal Benefits and Risks 35711.3.1 Beetles of Economic Importance 357

Detailed Table of Contentsxii

11.3.1.1 Negative Effects of Beetles 35711.3.1.2 Positive Effects of Beetles 36111.3.2 Beetles of Cultural Importance 37811.3.3 Beetles of Medical and Legal Importance 37811.3.3.1 Medical Entomology 37811.3.3.2 Forensic Entomology 37911.3.4 Beetles as Research Tools 38011.3.4.1 DNA Barcoding of Beetles – A North American Case Study 38211.4 Threatened Beetles 39411.5 Conclusions 395 Acknowledgments 395 References 395

12 Biodiversity of Hymenoptera 419John T. Huber

12.1 Evolution and Higher Classification 42212.2 Numbers of Species and Individuals 42612.3 Morphological and Biological Diversity 42812.4 Importance to Humans 42912.4.1 Food and Other Products 42912.4.2 Stings and Bites 43012.5 Ecological Importance 43112.6 Conservation 43212.7 Fossils 43212.8 Collecting, Preservation, and Study Techniques 43312.9 Taxonomic Diversity 43612.9.1 Symphyta 43712.9.2 Parasitica 43712.9.2.1 Stephanoidea 43712.9.2.2 Megalyroidea 43712.9.2.3 Trigonaloidea 43712.9.2.4 Mymarommatoidea 43712.9.2.5 Evanioidea 43812.9.2.6 Ichneumonoidea 43812.9.2.7 Cynipoidea 43912.9.2.8 Proctotrupoidea 43912.9.2.9 Platygastroidea 43912.9.2.10 Diaprioidea 43912.9.2.11 Ceraphronoidea 44012.9.2.12 Chalcidoidea 44012.9.3 Aculeata 44312.9.3.1 Chrysidoidea 44312.9.3.2 Vespoidea 44412.9.3.3 Apoidea 44612.10 Summary and Conclusions 446 Acknowledgments 446 References 447

Detailed Table of Contents xiii

13 Diversity and Significance of Lepidoptera: A Phylogenetic Perspective 463Paul Z. Goldstein

13.1 Relevance of Lepidoptera: Science 46413.2 Relevance of Lepidoptera: Society 46513.3 Diversity and Diversification: A Clarification of Numbers and Challenges 46613.4 State of Lepidopteran Systematics and Phylogenetics 46713.5 General Overview 46813.5.1 Primitive Lepidoptera 47713.5.2 Ditrysia 47813.5.2.1 Tineoidea 47813.5.2.2 Gracillarioidea 47913.5.2.3 Yponomeutoidea 48013.5.3 Apoditrysia 48013.5.3.1 Gelechioidea 48013.5.3.2 Pterophoroidea 48113.5.3.3 Tortricoidea 48113.5.3.4 Cossoidea 48113.5.3.5 Zygaenoidea 48213.5.4 Obtectomera 48213.5.5 Macroheterocera 48513.6 Needs and Challenges for Advancing Lepidopteran Studies 488 Acknowledgments 489 References 489

Part III Insect Biodiversity: Tools and Approaches 497

14 The Science of Insect Taxonomy: Prospects and Needs 499Quentin D. Wheeler and Kelly B. Miller

14.1 The What and Why of Taxonomy 50014.1.1 Improving Biology’s “General Reference System” 50514.1.2 Inter‐Generational Ethics 50614.1.3 Fulfilling Our Intellectual Manifest Destiny 50614.1.4 Solving Problems 50614.1.5 Model Organisms 50614.1.6 Molecular Tools of the Trade 50714.1.7 Aesthetics 50814.1.8 Creating the Vocabulary and Syntax of a Language of Biodiversity 50814.1.9 Mapping the Biosphere 50914.2 Insect Taxonomy: Missions and “Big Questions” 50914.3 Insect Taxonomy’s Grand Challenge Questions 51014.3.1 What Is a Species? 51014.3.2 What (and How Many) Insect Species Are There? 51114.3.3 What Is the Phylogeny of Insects? 51214.3.4 What Are the Histories of Character Transformation in Insects? 51214.3.5 Where Are Insect Species Distributed? 51214.3.6 How Have Insect Distributions Changed through Time? 51314.3.7 How Can Insect Classifications and Names Be Most Predictive and Informative? 513

Detailed Table of Contentsxiv

14.4 Transforming Insect Taxonomy 51314.5 Insect Taxonomy: Needs and Priorities 51414.5.1 Education 51414.5.2 Planetary‐Scale Projects and Virtual Species Observatories 51514.5.3 Cybertaxonomy Infrastructure 51514.5.4 Web‐Based Revisions, Taxon‐Knowledge Communities and Taxon‐Knowledge

Banks 51614.5.5 Collection Development and Growth 51714.5.6 Integrative Insect Taxonomy 51714.6 Accelerating Descriptive Taxonomy 51714.6.1 (1) Inventories to collections 51714.6.2 (2) Species descriptions 51914.6.3 (3) Species tests 51914.6.4 (4) Species tests to databases 52014.6.5 (5) Collection data 52014.6.6 (6) Cladistic analysis 52014.6.7 (7–10) Phylogenetic classifications, names, and identifications 52014.6.8 (12–16) Inputs 52114.7 Beware Sirens of Expediency 52114.8 Conclusions 522 References 522

15 Insect Species – Concepts and Practice 527Michael F. Claridge

15.1 Early Species Concepts – Linnaeus 52815.2 Biological Species Concepts 52915.2.1 Agamospecies 53115.2.2 Allopatric Forms 53115.3 Phylogenetic Species Concepts 53315.4 Species Concepts and Speciation – a Digression? 53415.5 Insect Species – Practical Problems 53515.5.1 Parthenogenetic Insects 53615.5.2 Species, Host Races, and Biotypes 53615.5.3 Specific Mate Recognition and Sibling Species 53815.6 Conclusions 540 References 540

16 Molecular Dimensions of Insect Taxonomy in the Genomics Era 547Amanda Roe, Julian Dupuis and Felix Sperling

16.1 Opportunities in Insect Taxonomy 54716.1.1 Determination 54816.1.2 Discovery 54916.1.3 Delimitation 55016.1.4 Phylogeny 55216.2 Genomic Methods 55316.2.1 Sequencing Technologies 553

Detailed Table of Contents xv

16.2.2 Genomic Sampling Strategies 55416.3 General Challenges and Considerations 55616.3.1 Data Quantity Versus Quality 55616.3.2 Phylogenetic Considerations 55716.3.2.1 Locus Selection 55716.3.2.2 Missing Data 55816.3.2.3 Gene Tree/Species Tree Incongruence 55816.3.3 Computational/Logistical/Bioinformatic Bottlenecks 55916.3.4 The Role of Morphology in a Post-genomic Era 56016.4 Conclusions 560 References 561

17 DNA Barcodes and Insect Biodiversity 575John-James Wilson, Kong-Wah Sing, Robin M. Floyd and Paul D. N. Hebert

17.1 Species Concepts and Recognition 57617.2 DNA Barcoding Methodology 57717.3 Basal Hexapod Orders 57817.4 Archaeognatha (Bristletails) and Zygentoma (Silverfish) 58017.5 Odonata (Dragonflies) 58017.6 Ephemeroptera (Mayflies) 58017.7 Orthoptera (Grasshoppers) 58017.8 Phasmatodea (Walking Sticks), Embioptera (Webspinners), Grylloblattodea

(Icecrawlers), and Mantophasmatodea (Gladiators) 58117.9 Plecoptera (Stoneflies) and Dermaptera (Earwigs) 58117.10 Mantodea (Mantids) 58117.11 Blattodea (Cockroaches) and Isoptera (Termites) 58117.12 Psocoptera (Booklice) and Phthiraptera (Lice) 58117.13 Thysanoptera (Thrips) and Hemiptera (True Bugs) 58217.14 Hymenoptera (Wasps) 58217.15 Strepsiptera (Twisted-wing Parasites) 58217.16 Coleoptera (Beetles) 58217.17 Neuroptera (Lacewings), Megaloptera (Dobsonflies), and Raphidioptera

(Snakeflies) 58317.18 Trichoptera (Caddisflies) 58317.19 Lepidoptera (Butterflies and Moths) 58317.20 Diptera (Flies) 58417.21 Siphonaptera (Fleas) and Mecoptera (Scorpionflies) 58417.22 Conclusions 584 Acknowledgments 585 References 585

18 Insect Biodiversity Informatics 593Norman F. Johnson

18.1 Biodiversity Data 59418.2 Technical Infrastructure 59518.3 Standards 597

Detailed Table of Contentsxvi

18.4 Current Status and Impediments to Progress 59918.5 Prospects 600 Acknowledgments 601 References 601

19 Parasitoid Biodiversity and Insect Pest Management 603John Heraty

19.1 What Is a Parasitoid? 60419.2 Biodiversity and Success of Insect Parasitoids 60519.2.1 Hymenoptera (Apocrita) 60619.2.2 Phoridae 60819.2.3 Tachinidae 60819.2.4 Other Groups 61019.2.5 Where Are Parasitoids Most Diverse? 61019.2.6 Leaf‐mining Parasitoids and Native Landscapes 61019.2.7 Are Parasitoids More Diverse in Tropical Versus Temperate Climates? 61219.3 Systematics, Parasitoids, and Pest Management 61219.3.1 Molecules and Parasitoid Biodiversity 61319.3.2 Cryptic Species 61419.3.3 DNA Barcoding and Biodiversity of Parasitoids 61619.3.4 Can Molecular Markers Be Applied to Understanding Biodiversity? 61719.4 Summary 617 Acknowledgments 618 References 618

20 The Taxonomy of Crop Pests: The Aphids 627Gary L. Miller and Robert G. Foottit

20.1 Historical Background 62720.2 Economic Importance and Early Taxonomy 62820.3 Early Aphid Studies – A North American Example 62820.4 Recognizing Aphid Species 63120.5 The Focus Becomes Finer 63220.6 Adventive Aphid Species 63320.7 Conclusions 634 References 634

21 Adventive (Non-Native) Insects and the Consequences for Science and Society of Species that Become Invasive 641Alfred G. Wheeler, Jr and E. Richard Hoebeke

21.1 Terminology 64221.2 Distributional Status: Native or Adventive? 64321.3 Global Transport: Pathways and Vectors 64521.4 Early History of Adventive Insects in North America 64821.5 Numbers, Taxonomic Composition, and Geographic Origins of Adventive Insects 64921.6 Impact of Adventive Insects 65321.6.1 Beneficial 656

Detailed Table of Contents xvii

21.6.2 Detrimental 65721.7 Economic Considerations: Agriculture, Forestry, and Horticulture 65821.7.1 Crop Losses 65821.7.2 Plant Diseases and Transmission of Pathogens 66021.8 Implications for Animal and Human Health 66121.9 Ecological Impacts 66321.9.1 Ants 66521.9.2 Bees and Wasps 66621.9.3 Forest Pests 66721.10 Biological Control 66721.11 Biological Invasions and Global Climate Change 67021.12 Systematics, Biodiversity, and Adventive Species 67121.13 Concluding Thoughts 671 Acknowledgments 674 References 675

22 Biodiversity of Blood-sucking Flies: Implications for Humanity 713Peter H. Adler

22.1 Numbers and Estimates 71422.2 Overview of Blood-sucking Flies and Diseases 71722.2.1 Athericidae 72022.2.2 Ceratopogonidae 72122.2.3 Corethrellidae 72122.2.4 Culicidae 72122.2.5 Glossinidae 72222.2.6 Hippoboscidae 72322.2.7 Muscidae 72322.2.8 Psychodidae 72422.2.9 Rhagionidae 72422.2.10 Simuliidae 72422.2.11 Tabanidae 72522.3 Rationale for Biodiversity Studies of Blood-sucking Flies 72522.4 Biodiversity Exploration 72722.5 Societal Consequences of Disregarding Biodiversity 72922.6 Present and Future Concerns 73022.7 Conclusions 733 Acknowledgments 734 References 734

23 Reconciling Ethical and Scientific Issues for Insect Conservation 747Michael J. Samways

23.1 Valuing Nature 74923.1.1 Types of Value 74923.1.2 Sensitive Use of Ecosystem Services 74923.1.3 Common‐Good Approaches 75123.1.4 Intrinsic Value and Conservation Action 752

Detailed Table of Contentsxviii

23.1.5 Reconciling Values 75323.2 Insects and Ecosystems 75523.2.1 Interactions and Multiple Effects 75523.2.2 Insects and Food Webs 75723.2.3 Importance of Maintaining Landscape Connectance 75723.3 Two Challenges 75823.3.1 The Taxonomic Challenge 75823.3.2 The Challenge of Complementary Surrogates 75923.4 Synthesizing Deeper Values and Practical Issues 75923.5 Summary 760 Acknowledgments 760 References 760

24 Taxonomy and Management of Insect Biodiversity 767Ke Chung Kim

24.1 Insect Biodiversity 76824.2 Biodiversity Loss and Humanity 76924.3 Biodiversity and Taxonomy 77024.4 Biodiversity Inventory and Ecology 77224.5 Backyard Biodiversity and Sustainability 77424.6 Taxonomic Bottlenecks in Managing Insect Biodiversity 77524.7 Advancing the Science of Insect Biodiversity 776 References 777

25 Insect Biodiversity – Millions and Millions 783May Berenbaum

Acknowledgments 789 References 791

Index 1 (Arthropoda-Hierarchic) 793

Index 2 (Arthropoda-Alphabetic) 825

Index 3 (Non-arthropods) 849

Index 4 (General Index Terms) 855

xix

Peter H. AdlerDepartment of Plant and Environmental SciencesClemson UniversityClemsonSouth CarolinaUSA

May BerenbaumDepartment of EntomologyUniversity of IllinoisUrbanaIllinoisUSA

Patrice BouchardCanadian National Collection of Insects, Arachnids and NematodesAgriculture and Agri‐Food CanadaOttawaOntarioCanada

Adam J. BrunkeCanadian National Collection of Insects, Arachnids and NematodesAgriculture and Agri‐Food CanadaOttawaOntarioCanada

Michael F. ClaridgeSchool of BiosciencesCardiff UniversityCardiffWalesUK

Gregory W. CourtneyDepartment of EntomologyIowa State UniversityAmesIowaUSA

Peter S. CranstonEvolution, Ecology and GeneticsResearch School of BiologyThe Australian National UniversityCanberraAustralia

Hugh V. DanksBiological Survey of CanadaCanadian Museum of NatureOttawaOntarioCanada

Hume DouglasCanadian National Collection of Insects, Arachnids and NematodesAgriculture and Agri‐Food CanadaOttawaOntarioCanada

Julian DupuisDepartment of Biological SciencesUniversity of AlbertaEdmontonAlbertaCanada

List of Contributors

List of Contributorsxx

Terry L. ErwinDepartment of EntomologyNational Museum of Natural HistorySmithsonian InstitutionWashington DCUSA

Robin M. FloydWellcome Trust/MRC Stem Cell InstituteUniversity of CambridgeCambridge UK

and

Centre for Biodiversity GenomicsBiodiversity Institute of OntarioUniversity of GuelphGuelphOntarioCanada

Robert G. FoottitCanadian National Collection of Insects, Arachnids and NematodesAgriculture and Agri‐Food CanadaOttawaOntarioCanada

Christy J. GeraciDepartment of EntomologyNational Museum of Natural History, Smithsonian InstitutionWashington DCUSA

Matthew L. GimmelInvertebrate ZoologySanta Barbara Museum of Natural HistorySanta BarbaraCaliforniaUSA

Paul Z. GoldsteinSystematic Entomology LaboratoryPlant Science InstituteAgriculture Research ServiceUS Department of Agriculturec/o Smithsonian InstitutionWashington DCUSA

Paul D. N. HebertCentre for Biodiversity GenomicsBiodiversity Institute of OntarioUniversity of GuelphGuelphOntarioCanada

Thomas J. HenrySystematic Entomology LaboratoryPlant Science InstituteAgriculture Research ServiceUS Department of Agriculturec/o Smithsonian InstitutionWashington DCUSA

John HeratyDepartment of EntomologyUniversity of CaliforniaRiversideCaliforniaUSA

E. Richard HoebekeGeorgia Museum of Natural History and Department of EntomologyUniversity of GeorgiaAthensGeorgiaUSA

John T. HuberNatural Resources CanadaCanadian Forestry Servicec/o Canadian National Collection of Insects, Arachnids and NematodesOttawaOntarioCanada

List of Contributors xxi

Norman F. JohnsonDepartment of Evolution, Ecology and Organismal Biology and Department of EntomologyOhio State UniversityColumbusOhioUSA

Kojun KandaDepartment of Biological SciencesNorthern Arizona UniversityFlagstaffArizonaUSA

Ke Chung KimFrost Entomological MuseumDepartment of EntomologyPennsylvania State UniversityUniversity ParkPennsylvaniaUSA

Alexander S. KonstantinovSystematic Entomology LaboratoryPlant Science InstituteAgriculture Research ServiceUS Department of Agriculturec/o Smithsonian InstitutionWashington DCUSA

Boris A. KorotyaevZoological InstituteRussian Academy of SciencesSt PetersburgRussia

Mervyn W. MansellDepartment of Zoology and EntomologyUniversity of PretoriaPretoriaSouth Africa

Gary L. MillerSystematic Entomology LaboratoryPlant Science InstituteAgricultural Research ServiceUS Department of AgricultureBeltsvilleMarylandUSA

Kelly B. MillerDepartment of BiologyUniversity of New MexicoAlbuquerqueNew MexicoUSA

John C. MorseDepartment of Plant and Environmental SciencesClemson UniversityClemsonSouth CarolinaUSA

Thomas PapeNatural History Museum of DenmarkUniversity of CopenhagenCopenhagenDenmark

Amanda RoeNatural Resources CanadaCanadian Forest ServiceGreat Lakes Forestry CentreSault Ste. MarieOntarioCanada

Michael J. SamwaysDepartment of Conservation Ecology and EntomologyStellenbosch UniversityMatielandSouth Africa

List of Contributorsxxii

Clarke H. ScholtzDepartment of Zoology and EntomologyUniversity of PretoriaPretoriaSouth Africa

Geoffrey G. E. ScudderDepartment of ZoologyUniversity of British ColumbiaVancouverBritish ColumbiaCanada

Bradley J. SinclairCanadian National Collection of Insects and Canadian Food Inspection AgencyOttawa Plant Laboratory – EntomologyOttawaOntarioCanada

Kong-Wah SingState Key Laboratory of Genetic Resources and EvolutionKunming Institute of ZoologyChinese Academy of Sciences KunmingP. R. China

and

Institute of Biological SciencesUniversity of MalayaKuala LumpurMalaysia

Jeffrey H. SkevingtonCanadian National Collection of Insects, Arachnids and NematodesAgriculture and Agri‐Food CanadaOttawaOntarioCanada

Andrew B. T. SmithResearch DivisionCanadian Museum of NatureOttawaOntarioCanada

Felix SperlingDepartment of Biological SciencesBiological Sciences CentreUniversity of AlbertaEdmontonAlbertaCanada

Mark G. VolkovitshZoological InstituteRussian Academy of SciencesSt PetersburgRussia

Alfred G. Wheeler, JrDepartment of Plant and Environmental SciencesClemson UniversityClemsonSouth CarolinaUSA

Quentin D. WheelerCollege of Environmental Science and ForestryState University of New YorkSyracuseNew YorkUSA

John-James WilsonInternational College BeijingChina Agricultural UniversityBeijingP. R. China

and

Institute of Biological SciencesUniversity of MalayaKuala LumpurMalaysia

Laura S. ZamoranoDepartment of EntomologyNational Museum of Natural HistorySmithsonian InstitutionWashington DCUSA

xxiii

Insects are the most exuberant manifestation of Earth’s many and varied life forms. Their rather simple unifying body plan has become modified and adapted to produce an enormous variety of species, and insects exploit virtually all terres­trial and freshwater environments on the planet, as well as many brackish waters. However, as a paradox debated extensively a few decades ago, they are largely absent from the seas and oceans. Features such as wings and the complete meta­morphosis of many species have been cited fre­quently as fostering this massive diversity. The “success” of the insects can be measured by many parameters: their long‐term persistence and stability of their basic patterns, the variety of higher groups (with almost 30 orders com­monly recognized) and, as emphasized in this book, the wealth of species and similar entities. Each of these species has its individual biologi­cal peculiarities, ecological role, distribution, and interactions within the local community. And each may differ in habit and appearance, both from its close relatives and across its range, to reflect local influences and conditions. Every species is thus a mosaic of physical variety and genetic constitution that can lead to taxonomic and ecological ambiguity in interpreting its integrity and the ways in which it may evolve and persist.

Entomologists will continue to debate the number of insect “species” that exist and the levels of past and likely future extinctions that edit any such estimate. The difficulties in gain­ing consensus have two main axes – first, lack of understanding of how these entities may be

recognized and categorized and, second, that many insect groups remain substantially under­collected and are poorly known. The first of these themes dominates much of this book  – gaining agreement over “what is a species” is difficult and sometimes contentious. Many tax­onomists hold strong and individualistic views, molded by years of study, of the limits of species and the validity of infraspecific categories such as subspecies and races that in practice can function as “evolutionarily significant units” in their insect group. One widespread trend, often not appreciated fully, is that widespread gener­alist insect species may not persist as such as their environment changes  –  loss of resources and fragmentation of previously extensive bio­topes may cause populations to become iso­lated, and restricted to a limited subset of resources, such as particular host plants, on which they must then depend and specialize. Such situations may beget speciation, perhaps especially among phytophagous insects that display many examples of such localized but obligatory isolation. Populations involved com­monly show haplotype differences and bio­logical idiosyncrasies related to their local conditions, but otherwise are not easily separa­ble. Generalist “species” may commonly com­prise complexes of cryptic species masquerading as a single entity. Conventional “typological” taxonomists may tend to mirror the more con­servative “generalist” approach, whereas other constituents (such as many butterfly collectors) may opt to recognize numerous isolated popu­lations displaying small phenotypic variations as

Foreword, Second Edition

Foreword, Second Editionxxiv

distinct (specific or subspecific) taxa. Individual specialists in any large insect group are likely to occupy different positions along the gradient of “lumpers” to “splitters” in how they treat such  variety, and may defend their stance energetically.

Biologists and philosophers alike continue to debate the merits of the numerous species con­cepts, drawing on the reality quoted by one recent commentator that “there are n+1 defini­tions of ‘species’ in a room of n biologists,” with the most common inference that “a species is whatever a taxonomist says it is.” All recognized categories, however, are dynamic. Any given fig­ure for insect diversity (as numbers of species) is a working hypothesis, as is each of the contrib­uting species – so that complete and enduring enumeration is perhaps impossible to achieve.

Documenting and cataloging insect biodiver­sity as a major component of Earth’s life is a natural quest of human inquiry, but is not an end in itself and, importantly, is not synony­mous with conserving insects or a necessary prerequisite to assuring their well‐being. Despite many ambiguities in projecting the actual num­bers of insect species, no one would query that there are a lot, and that the various ecological processes that sustain ecosystems depend heav­ily on insect activity. Indeed, “ecological ser­vices” such as pollination, recycling of materials, and the economically important activities of predators and parasitoids are signaled increas­ingly as part of the rationale for insect conserva­tion because these values can be appreciated easily through direct economic impacts. All these themes are dealt with in this book, cen­tered on questions related to our ignorance of fundamental matters of “how many are there?” and “how important are they?”, to which the broad answers of “millions” and “massive” may incorporate considerable uncertainty; this uncertainty, however, is reduced by many of the chapters here.

In any investigations of insect biodiversity, the role of inventory tends to be emphasized, despite the impracticability of achieving complete enu­meration. Documenting numbers of species

(however they are delimited or defined) gives us foci for conservation advocacy and is pivotal in helping to elucidate patterns of evolution and distribution. Recognizing and naming species allow us to transfer information, but high proportions of undescribed or unrecognizable species necessitate the use of terms such as “morphospecies” in much ecological interpreta­tion of diversity. Accompanying archival deposi­tion of voucher specimens is then needed as the only reliable means through which the consist­ency of separations can be affirmed and cross‐survey comparisons validated. Nevertheless, other than in some temperate regions, particu­larly in the northern hemisphere, many esti­mates of insect species richness and the naming of the species present are highly incomplete. Much of the tropics, for example, harbors few resident entomologists other than those involved with pressing problems of human welfare, and more basic and sustained documentation almost inevitably depends on assistance from elsewhere. Some insects, of course, have been explored more comprehensively than others, so that selected taxonomic groups (such as butterflies, larger beetles, and dragonflies) and ecological groups (“pests”) have received more attention than many less charismatic or less economically important groups. Indeed, when collecting Psocoptera in parts of the tropics, I have occa­sionally been asked by local people why I am not collecting birdwing butterflies, stag beetles, or other “popular” (or commercially desirable!) insects, and my responses have done little to change their opinions of my insanity!

In short, many gaps in knowledge of insect diversity persist, and seem unlikely to be redressed effectively in the near future, other than by “guesstimates” extrapolating from sometimes rather dubious foundations. How­ever, sufficient knowledge does exist to endorse the practical need to protect natural habitats from continued despoliation and, as far as practicable, from the effects of climate change. Citations of impressively large numbers of insect species can become valuable advocacy in  helping to conserve areas with largely

Foreword, Second Edition xxv

unheralded wealth of biodiversity. The pres­ence of unusual lineages of insects, of narrow‐range endemics, and highly localized radiations and distributional idiosyncrasies (such as iso­lated populations beyond the main range of the taxon) are all commonplace scenarios, and may in various ways help us to designate priori­ties  for allocating the limited conservation resources available. Many such examples from selected insect groups are revealed in this book – but evaluating the richness and ecologi­cal importance of the so‐called meek inheri­tors, that vast majority of insects that do not intrude notably on human intelligence and wel­fare, remains a major challenge. Many such taxa receive attention from only a handful of entomologists at any time, and some are essen­tially “orphaned” for considerable periods. Progress with their documentation is inevitably slow and sporadic. Some hyperdiverse orders and families of insects exhibit daunting com­plexity of form and biology, as “black hole groups” whose elucidation is among the major challenges that face us.

Insect conservation has drawn heavily on issues relevant to biodiversity and appreciation of the vast richness of insects, not only of easily recognizable “species”, but also of the occur­rence of subspecies and other infraspecific var­iants, such as significant populations. This more complex dimension of insect biodiversity is receiving considerable attention as new molecular tools (such as DNA analysis) enable us to probe characters in ways undreamed of only a decade or so ago to augment the per­spective provided by morphological interpreta­tion, and assess relationships within lineages and their rates of differentiation. Applications of these tools proliferate, sometimes to the extent where small molecular differences treated in isolation may confuse, rather than clarify, relationships implied from more tradi­tional approaches. The vast arrays of cryptic species gradually being revealed suggest that even our most up‐to‐date estimates of species numbers based on morphological data may be

woefully inadequate. Insect diversity equates to “variety,” but the subtleties of interpopulation variations in genetic constitution and ecologi­cal performance are difficult to appraise and to categorize formally  –  and perhaps even more difficult to communicate to non‐entomologists whose powers may determine the future of the systems in which those insects participate. Education and communication, based on the soundest available information, are essential components of insect conservation. This book is a significant contribution to this endeavor, through indicating how we may come to inter­pret and understand insect biodiversity more effectively. In addition to providing a range of opinions and facts on insect richness in a vari­ety of taxonomic, geographical, and methodo­logical contexts, it helps to emphasize the scientific and political importance of accurate species recognition. Failure to recognize adven­tive alien species may have dire economic or ecological consequences, or confusion between biotypes or cryptic species may invalidate expensive management programs for their sup­pression or conservation.

A new generation of skilled insect systema­tists – whose visions encompass the wider rami­fications of insect biodiversity, its importance in understanding the natural world, and the accel­erating impacts of humans upon it – is an urgent need. They enter an exciting and challenging field of endeavor, and the perspectives included in this volume are essential background to their future contributions. The first edition of this book was a foundation and a stimulating work­ing tool toward that end, and I expect many of the renewed chapters to become key references as we progressively refine and enlarge the bases of our understanding of insects and their activi­ties in the modern world.

Tim NewDepartment of Ecology,

Environment and EvolutionLa Trobe University

xxvii

Insects are the world’s most diverse group of animals, making up more than 58 percent of the known global biodiversity. They inhabit all habi-tat types and play major roles in the function and stability of terrestrial and aquatic ecosys-tems. Insects are closely associated with our lives and affect the welfare of humanity in diverse ways. At the same time, large numbers of insect species, including those not known to sci-ence, continue to become extinct or extirpated from local habitats worldwide. Our knowledge of insect biodiversity is far from complete; for  example, barely 65 percent of the  North American insect fauna has been described. Only a relatively few species of insects have been studied in depth. We urgently need to explore and describe insect biodiversity and to better understand the biology and ecology of insects if ecosystems are to be managed sustainably and if the effect of global environment change is to be mitigated.

The scientific study of insect biodiversity is at a precarious point. Resources for the support of taxonomy are tenuous worldwide. The number of taxonomists is declining and the output of taxonomic research has slowed. Many taxono-mists are reaching retirement age and will not be replaced with trained scientists, which will result in a lack of taxonomic expertise for many groups of insects. These trends contrast with an increas-ing need for taxonomic information and ser-vices in our society, particularly for biodiversity assessment, ecosystem management, conserva-tion, sustainable development, management of climate‐change effects, and pest management.

In light of these contrasting trends, the scientific community and its leadership must increase their understanding of the science of insect bio-diversity and taxonomy and ensure that policy makers are informed of the importance of biodi-versity for a sustainable future for humanity.

We have attended and contributed to many scientific meetings and management and policy gatherings where the future, the resource needs, and importance of insect taxonomy and biodi-versity have been debated. In fact, discussion of the future of taxonomy is a favorite pastime of taxonomists; there is no shortage of “taxonomic opinion.” Considerable discussion has focused on the daunting task of describing the diversity of insect life and how many undescribed spe-cies are out there. However, we felt that there was a need for an up‐to‐date, quantitative assessment of what insect biodiversity entails, and to connect what we know and don’t know about insect biodiversity with its impact on human society.

Our approach was to ask authors to develop accounts of biodiversity in certain orders of insects and geographic regions and along selected subject lines. In all categories, we were limited by the availability of willing contribu-tors and their time and resources. Many insect groups, geographic regions, and scientific and societal issues could not be treated in a single volume. It also was apparent to us, sometimes painfully so, that many taxonomists are wildly overcommitted. This situation can be seen as part of the so‐called “taxonomic impediment” – the lack of available taxonomic expertise is

Preface, First Edition

Preface, First Editionxxviii

compounded by an overburdened community of present‐day taxonomists with too much work and perhaps too much unrealistic enthusiasm.

In Chapter 1, we introduce the ongoing chal-lenge to document insect biodiversity and develop its services. Chapter 2 provides a com-prehensive overview of the importance and value of insects to humans. The next two sec-tions deal with regional treatments and ordinal‐level accounts of insect biodiversity. These approaches were a serious challenge to the con-tributors who had to compile information from a wide array of sources or, alternatively, deal with situations in which accurate information simply is insufficient. In Section III, we document some of the tools and approaches to the science of tax-onomy and its applications. Perspective is pro-vided on the past, present, and future of the science of insect taxonomy and the all‐impor-tant influence of species concepts and their

operational treatment on taxonomic science and insect biodiversity. Contributions on the increas-ing role of informatics and molecular approaches are provided, areas with ongoing controversy and differences of opinion. These chapters are followed by contributions on the applications of taxonomic science for which biodiversity infor-mation is fundamental, including the increasing impact of adventive insects, pest detection and management, human medical concerns, and the management and conservation of biodiversity. The book ends with an historical view of the continuing attempts to document the extent of world insect biodiversity.

Robert G. FoottitOttawa, Ontario

Peter H. AdlerClemson, South Carolina