ECOLOGY AND BEHAVIOUR OF THE LADYBIRD () ECOLOGY AND BEHAVIOUR OF THE LADYBIRD BEETLES (COCCINELLIDAE)

Edited by I. Hodek, H.F. van Emden and A. Honeˇk

A John Wiley & Sons, Ltd., Publication This edition fi rst published 2012 © 2012 by Blackwell Publishing Ltd

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Library of Congress Cataloging-in-Publication Data Ecology and behaviour of the ladybird beetles (Coccinellidae) / edited by I. Hodek, H.F. van Emden, and A. Honeˇk. p. cm. Includes index. ISBN 978-1-4051-8422-9 (cloth) 1. Ladybugs. I. Hodek, Ivo. II. Van Emden, Helmut Fritz. III. Honeˇk, A. (Alois) QL596.C65E26 2012 595.76'9–dc23 2011045545

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1 2012 To all those scientists who have now passed on and who laid the foundation of our present knowledge of Coccinellidae, particularly to Michael Majerus who intended to be one of authors of this volume. Contents

Detailed contents, ix 8. NATURAL ENEMIES OF LADYBIRD BEETLES, 375 Contributors, xvii Piotr Ceryngier, Helen E. Roy and Remy L. Poland Preface, xviii 9. COCCINELLIDS AND Introduction, xix SEMIOCHEMICALS, 444 Jan Pettersson Taxonomic glossary, xx 10. QUANTIFYING THE IMPACT OF 1. PHYLOGENY AND CLASSIFICATION, 1 COCCINELLIDS ON THEIR PREY, 465 Oldrich Nedveˇd and Ivo Kovárˇ J. P. Michaud and James D. Harwood

2. GENETIC STUDIES, 13 11. COCCINELLIDS IN BIOLOGICAL John J. Sloggett and Alois Honeˇk CONTROL, 488 J. P. Michaud 3. LIFE HISTORY AND DEVELOPMENT, 54 12. RECENT PROGRESS AND POSSIBLE Oldrich Nedveˇd and Alois Honeˇk FUTURE TRENDS IN THE STUDY OF COCCINELLIDAE, 520 4. DISTRIBUTION AND HABITATS, 110 Helmut F. van Emden and Ivo Hodek Alois Honeˇk Appendix: List of Genera in Tribes and Subfamilies, 526 5. FOOD RELATIONSHIPS, 141 Oldrich Nedveˇd and Ivo Kovárˇ Ivo Hodek and Edward W. Evans Subject index, 532 6. DIAPAUSE/DORMANCY, 275 Colour plate pages fall between pp. 250 and pp. 251 Ivo Hodek

7. INTRAGUILD INTERACTIONS, 343 Éric Lucas

vii DETAILED CONTENTS

Contributors, xvii 1.4.9 Coccidulinae, 8 1.4.10 Scymninae, 9 Preface, xviii 1.5 Future Perspectives, 10 References, 10 Introduction, xix

Taxonomic glossary, xx 2. GENETIC STUDIES, 13 John J. Sloggett and Alois Honeˇk 1. PHYLOGENY AND CLASSIFICATION, 1 2.1 Introduction, 14 Oldrich Nedveˇd and Ivo Kovárˇ 2.2 Genome Size, 14 1.1 Position of the Family, 2 2.3 Chromosomes and Cytology, 14 1.1.1 The Cerylonid complex, 2 2.3.1 Chromosome numbers and 1.1.2 Sister families, 2 banding, 14 1.1.3 Feeding habits, 2 2.3.2 Sex determination, 14 1.1.4 Monophyly of Coccinellidae, 2 2.3.3 Supernumerary (B) 1.2 Characteristics of the Family, 3 chromosomes, 15 1.3 Changes in the Classifi cation of Subfamilies, 3 2.3.4 Cytogenetic changes, intraspecifi c 1.3.1 Morphologically based cytogenetic variation and classifi cations, 3 speciation, 16 1.3.2 Split of Sticholotidinae, 3 2.4 Colour Pattern Variation, 18 1.3.3 Monophyly of other subfamilies, 4 2.4.1 The nature of colour patterns, 19 1.3.3.1 Contribution of immature 2.4.2 Genetic determination of colour stages, 4 patterns, 20 1.3.4 Molecular analyses, 4 2.4.3 Geographic variation, 23 1.3.4.1 Alternative molecular 2.4.4 Temporal variation, 25 methods, 5 2.4.5 Signifi cance and evolution, 25 1.3.5 Rejection of the monophyly of 2.5 The Inheritance of Other Traits, 27 subfamilies, 5 2.5.1 Morphological characters: wing 1.4 Characteristics of the Subfamilies and polymorphism, 27 Tribes, 5 2.5.2 Life history characters: heritability, 1.4.1 Proposed classifi cation, 5 selection experiments and genetic 1.4.2 Microweiseinae, 5 trade-offs, 28 1.4.3 Sticholotidinae, 5 2.6 Molecular Genetic Studies, 29 1.4.4 , 6 2.6.1 Sequence evolution, 29 1.4.5 , 7 2.6.1.1 Mitochondrial DNA and 1.4.6 Exoplectrinae, 7 the inference of 1.4.7 Chilocorinae, 7 evolutionary history, 29 1.4.8 Ortaliinae, 8 2.6.1.2 The ITS1 region, 31 ix x Detailed contents

2.6.2 Molecular studies of coccinellid 3.5.6.3 Female choice and biology, 33 melanism, 83 2.6.2.1 Species, population 3.5.6.4 Hybridization, 85 and strain 3.5.7 Oviposition, 85 identifi cation, 33 3.5.7.1 Oviposition substrate, 85 2.6.2.2 Phylogenetics, 37 3.5.7.2 Oviposition 2.6.2.3 Population genetic and rhythmicity, 86 phylogeographic 3.5.7.3 Oviposition rate, 86 studies, 38 3.5.7.4 Oviposition period, 86 2.6.2.4 Reproductive success, 3.5.8 Fecundity, 88 paternity and sperm 3.5.9 Longevity, 90 competition, 42 3.5.9.1 Voltinism, 90 2.7 Conclusions, 43 3.5.9.2 Effect of temperature, Acknowledgements and Dedication, 44 photoperiod and References, 44 humidity, 90 3.5.9.3 Effect of food, 91 3.5.9.4 Effect of sexual 3. LIFE HISTORY AND DEVELOPMENT, 54 activity, 91 Oldrich Nedveˇd and Alois Honeˇk 3.6 Temperature and Development, 91 3.1 Introduction, 55 3.6.1 Thermal constants, 92 3.2 Egg, 55 3.6.2 Relationship between LDT 3.2.1 Egg morphology, 55 and SET, 92 3.2.2 Egg size, 56 3.6.3 Thermal window and development 3.2.3 Cluster size, 56 rate isomorphy, 95 3.2.4 Hatching rate, 68 3.6.4 Other events affected by 3.2.5 Trophic eggs, 69 temperature, 95 3.3 Larva, 71 3.6.4.1 Tolerance to extreme 3.3.1 Larval morphology, 71 temperatures, 96 3.3.2 Instars, 71 Acknowledgements, 97 3.3.3 Development, 72 References, 97 3.3.4 Body size, 73 3.4 Pupa, 73 4. DISTRIBUTION AND HABITATS, 110 3.4.1 Prepupal stage, 73 3.4.2 Pupal morphology, 74 Alois Honeˇk 3.4.3 Timing of pupation, 75 4.1 Introduction, 111 3.4.4 Places of pupation, 75 4.2 Sampling, 111 3.4.5 Pupal defence, 76 4.3 Factors Determining the Composition of 3.4.6 Colouration and thermal Communities, 114 melanism, 76 4.3.1 Local faunas, 114 3.5 Adult, 76 4.3.1.1 Geographic 3.5.1 Teneral development, 76 differences, 114 3.5.2 Wings and fl ight, 77 4.3.1.2 Invasion and 3.5.3 Pre-oviposition period, 77 extinction, 115 3.5.4 Size, 78 4.3.1.3 Climatic changes, 115 3.5.5 Ovarioles, 78 4.3.2 Locality determinants, 116 3.5.6 Mating, 82 4.3.2.1 Prey, 116 3.5.6.1 Frequency and duration 4.3.2.2 Host plant, 120 of mating, 82 4.3.2.3 Microclimate, 124 3.5.6.2 Sperm competition, 83 4.3.2.4 Landscape, 125 Detailed contents xi

4.3.3 Dominance, diversity and niche 5.2.12 Tritrophic studies, 196 differentiation, 127 5.2.13 Food of phytophagous 4.4 Coccinellid Communities of Particular Coccinellidae, 198 Habitats, 128 5.2.14 Food of mycophagous 4.5 Conclusion, 132 Coccinellidae, 200 Acknowledgements, 133 5.3 Quantitative Aspects of Food Relations, 201 References, 133 5.3.1 Effect of physical factors on consumption, 202 5.3.1.1 Daily consumption 5. FOOD RELATIONSHIPS, 141 rate, 202 Ivo Hodek and Edward W. Evans 5.3.1.2 Total food 5.1 Introduction, 142 consumption, 202 5.2 Food Specifi city, 142 5.3.2 Effect of prey density on 5.2.1 Food range, 142 consumption: functional 5.2.1.1 Methods for detection of response, 204 food range, 144 5.3.3 Effects of consumption on growth 5.2.2 Nutritional suitability of food, 145 and reproduction, 207 5.2.3 Prey size–density hypothesis, 147 5.3.3.1 Larval development, 207 5.2.4 Euryphagous and stenophagous 5.3.3.2 Adult performance, 208 species / Generalist and specialist 5.3.4 Conversion and utilization of species, 150 consumed food, 209 5.2.4.1 Generalists, 152 5.3.5 Aggregative numerical 5.2.4.2 Specialists, 154 response, 211 5.2.5 Mixed and combined diet, 155 5.3.5.1 Temporal and spatial 5.2.5.1 Complementation across patterns, 211 stages, 156 5.3.5.2 Modeling of aggregative 5.2.5.2 Prey switching, 157 responses, 211 5.2.5.3 Prey specialization 5.3.5.3 Factors other than focal through selection, 157 prey density, 211 5.2.6 Lower quality prey 5.4 Food-Related Behaviour, 213 (mostly aphids), 157 5.4.1 Foraging behaviour, 213 5.2.6.1 Toxic prey, 157 5.4.1.1 Indirect factors in 5.2.6.2 Rejected prey, 165 foraging, 215 5.2.6.3 ‘Problematic’ prey, 167 5.4.1.2 The role of senses in 5.2.7 Prey other than aphids/coccids, 169 foraging, 218 5.2.7.1 Developmental stages of 5.4.1.3 Finding an oviposition Holometabola, 169 site, 226 5.2.7.2 Non-aphid 5.4.1.4 Foraging of fi rst hemipterans, 174 instars, 233 5.2.8 Cannibalism, 175 5.4.1.5 Movement among 5.2.8.1 Sibling egg habitats/patches in the cannibalism, 175 landscape, 234 5.2.8.2 Non-sibling egg 5.4.1.6 Ladybird foraging and cannibalism, 177 ants, 236 5.2.9 Non- food (pollen, nectar, 5.4.2 Prey capture, 238 spores of fungi), 180 5.4.3 Food intake, 241 5.2.10 Substitute diets and food supplements 5.5 Conclusions, 242 (sprays), 185 Acknowledgements, 242 5.2.11 Essential foods, 187 References, 243 xii Detailed contents

6. DIAPAUSE/DORMANCY, 275 6.2.16 Ecophysiological regulation of Ivo Hodek diapause in coccinellids, 300 6.3 Behaviour Patterns Related to Diapause, 301 6.1 Introduction: Mechanisms and 6.3.1 Phases of dormancy behaviour, 301 Defi nitions, 276 6.3.1.1 Pre-diapause, 301 6.1.1 Hibernation and aestivation, 276 6.3.1.2 Migration, 302 6.1.2 Termination/completion of 6.3.1.3 Flight and methods for its diapause, 277 study, 303 6.1.3 Phases of dormancy, 277 6.3.1.4 Aggregations, 305 6.1.4 Endocrinological aspects of adult 6.3.1.5 Emergence from diapause, 277 dormancy sites, 309 6.2 Ecophysiological Regulation of Diapause in 6.3.2 Behaviour of individual Coccinellids, 278 species, 309 6.2.1 Coccinella septempunctata, 278 6.3.2.1 Ceratomegilla 6.2.1.1 Central Europe, 278 (=Semiadalia) 6.2.1.2 Western Europe: undecimnotata, 309 , 282 6.3.2.2 Hypsotactic species 6.2.1.3 Northern Europe, 283 Harmonia axyridis, 6.2.1.4 Mediterranean Har. conformis and region, 284 Aiolocaria hexaspilota 6.2.1.5 Nearctic region, 285 (=mirabilis), 310 6.2.1.6 Potential multivoltines 6.3.2.3 Coccinella septempunctata in univoltine and other species populations, 286 dormant in the litter, 311 6.2.2 Coccinella septempunctata 6.3.2.4 Coleomegilla maculata, 313 brucki, 287 6.3.2.5 Adalia bipunctata, 313 6.2.2.1 Central Japan (central 6.3.2.6 Myrrha octodecimguttata Honshu), 287 and other forest species 6.2.2.2 Sapporo, Hokkaido which hibernate in bark (Japan), 288 crevices, 314 6.2.2.3 Northern Honshu 6.3.2.7 Hippodamia (Japan), 289 convergens, 314 6.2.3 Coccinella novemnotata, 289 6.3.2.8 Hippodamia 6.2.4 Adalia bipunctata, 290 quinquesignata, 316 6.2.5 Propylea quatuordecimpunctata and 6.3.2.9 Hibernation of P. dissecta, 291 mycophagous and 6.2.6 Hippodamia tredecimpunctata, 292 phytophagous 6.2.7 Hippodamia convergens, 292 species, 316 6.2.8 Ceratomegilla (=Semiadalia) 6.4 Anatomical and Physiological Changes undecimnotata, 292 Related to Dormancy, 316 6.2.9 Harmonia axyridis, 295 6.4.1 Anatomical state, 316 6.2.10 Coccinella leonina (=repanda), 297 6.4.1.1 Fat body and digestive 6.2.11 Apolinus (=Scymnodes) lividigaster and tract, 316 (=Leptothea) galbula, 297 6.4.1.2 Ovary, spermatheca, 317 6.2.12 Harmonia sedecimnotata, 297 6.4.1.3 Male gonads, 321 6.2.13 Chilocorus spp., 298 6.4.1.4 Flight muscles, 323 6.2.14 Stethorus punctum picipes and S. 6.4.2 Metabolic changes related to japonicus, 299 diapause, 323 6.2.15 Scymnus (Neopullus) 6.4.2.1 Lipids, 323 sinuanodulus, 300 6.4.2.2 Glycogen, 325 Detailed contents xiii

6.4.2.3 Water, 326 7.9.1 Defensive mechanisms of coccinellids 6.4.2.4 Metabolic rate, 326 against intraguild predation, 355 6.4.3 Corpora allata and regulation of 7.9.1.1 Defence of all stages, 355 vitellogenesis, 327 7.9.1.2 Defence of eggs, 356 6.4.4 Cold-hardiness, 328 7.9.1.3 Defence of larvae, 356 6.5 Conclusions and Lacunae in Knowledge, 332 7.9.1.4 Defence of moulting Acknowledgements, 333 individuals and References, 333 pupae, 357 7.9.1.5 Defence of adults, 357 7.9.2 Intraguild predation on 7. INTRAGUILD INTERACTIONS, 343 coccinellids, 358 Éric Lucas 7.9.2.1 Intraguild predation by 7.1 Scope, 344 intraguild predators, 358 7.2 Coccinellids as Guild Members, 344 7.9.2.2 Intraguild predation 7.3 Coccinellids as Neutralists, 344 by intraguild 7.3.1 Temporal guild partition, 344 parasitoids, 358 7.3.2 Spatial guild partition, 344 7.9.2.3 Intraguild predation by 7.3.3 Thermal guild partition, 344 intraguild pathogens, 359 7.3.4 Body size guild partition, 345 7.10 Coccinellids Interacting with Intraguild 7.4 Coccinellids as Interacting Organisms, 345 Ants, 359 7.5 Coccinellids as Intraguild Commensalists and 7.10.1 Non-myrmecophilous coccinellid Mutualists, 345 species, 359 7.6 Coccinellids as Competitors, 347 7.10.2 Myrmecophilous coccinellid 7.6.1 Exploitative competition, 347 species, 360 7.6.2 Apparent competition, 349 7.11 Applied Aspects of Intraguild Interactions, 360 7.7 Coccinellids as Vectors of Male-Killing 7.11.1 Conservation, 360 Bacteria, 349 7.11.2 Biological control, 361 7.8 Coccinellids as Intraguild Predators, 349 7.11.2.1 Intraguild predation and 7.8.1 General rules of intraguild biocontrol, 361 predation involving coccinellid 7.11.2.2 Facilitation and predators, 349 biocontrol, 361 7.8.2 Occurrence of intraguild predation 7.11.2.3 Ants and biocontrol, 362 by coccinellids, 350 7.11.2.4 Intraguild interactions 7.8.3 Intraguild predation on intraguild and biocontrol coccinellids (by coccinellids), 351 approaches, 362 7.8.4 Intraguild predation on intraguild 7.12 Interguild Effects, 362 non-coccinellid predators, 352 7.13 Conclusion, 363 7.8.4.1 Intraguild References, 364 neuropterans, 352 7.8.4.2 Intraguild dipterans, 352 8. NATURAL ENEMIES OF LADYBIRD 7.8.4.3 Intraguild BEETLES, 375 hemipterans, 353 7.8.4.4 Other intraguild Piotr Ceryngier, Helen E. Roy and Remy L. Poland predators, 353 8.1 Introduction, 376 7.8.5 Intraguild predation on intraguild 8.2 Predation and Related Phenomena, 376 parasitoids, 353 8.2.1 Anti-predator defences, 376 7.8.6 Intraguild predation on intraguild 8.2.1.1 Aposematic colouration pathogens, 354 and other visual 7.8.7 Coccinellids as top predators, 354 signals, 376 7.9 Coccinellids as Intraguild Prey, 355 8.2.1.2 Refl ex bleeding, 376 xiv Detailed contents

8.2.1.3 Morphological anti- 8.4 Parasites and Pathogens, 411 predator adaptations, 377 8.4.1 Acarina, 411 8.2.2 Vertebrate predators, 377 8.4.1.1 Phoretic mites, 411 8.2.3 Invertebrate predators, 377 8.4.1.2 Coccipolipus Husband 8.2.4 Hemiptera-tending ants, 379 (Prostigmata: 8.2.5 Social aphids with a soldier caste, 382 Podapolipidae), 411 8.3 Parasitoids, 383 8.4.2 Nematodes, 414 8.3.1 General characteristics of parasitoids 8.4.2.1 Allantonematidae of ladybirds, 390 (Tylenchida), 414 8.3.1.1 Host specifi city of 8.4.2.2 Mermithidae parasitoids recorded from (Mermithida), 415 Coccinellidae, 390 8.4.3 Fungal pathogens, 415 8.3.1.2 Parasitoids of different 8.4.3.1 Hypocreales subfamilies of (Ascomycota), 415 Coccinellidae, 390 8.4.3.2 Hesperomyces spp. 8.3.1.3 Parasitism in different (Ascomycota: developmental stages of Laboulbeniales, Coccinellidae, 390 Laboulbeniaceae), 416 8.3.2 Review of the more important 8.4.3.3 Nosematidae taxa, 391 (Microsporidia), 418 8.3.2.1 Dinocampus Foerster 8.4.4 Protozoan pathogens, 419 (Hymenoptera: 8.4.4.1 Septate eugregarines Braconidae, (Apicomplexa: Euphorinae), 391 Eugregarinida: 8.3.2.2 Uga Girault Septatorina), 419 (Hymenoptera: 8.4.5 Bacteria, 421 Chalcididae), 399 8.4.5.1 General pathogenic 8.3.2.3 Cowperia Girault bacteria, 421 (Hymenoptera: 8.4.5.2 Male-killing bacteria, 421 Encyrtidae), 399 8.5 Impact of Natural Enemies on Ladybird 8.3.2.4 Homalotylus Mayr Populations, 425 (Hymenoptera: 8.5.1 Impact on phytophagous Encyrtidae), 400 Coccinellidae, 426 8.3.2.5 Nothoserphus Brues 8.5.2 Impact on predatory (Hymenoptera: Coccinellidae, 426 Proctotrupidae), 404 8.5.3 Concluding note, 428 8.3.2.6 Metastenus Walker References, 428 (Hymenoptera: Pteromalidae), 405 9. COCCINELLIDS AND 8.3.2.7 Phalacrotophora Enderlein SEMIOCHEMICALS, 444 (Diptera: Phoridae), 406 8.3.2.8 Oomyzus scaposus Jan Pettersson (Thomson) 9.1 Introduction, 445 (Hymenoptera: 9.2 Aposematism and Refl ex Bleeding Eulophidae, Chemistry, 445 Tetrastichinae), 408 9.2.1 Refl ex bleeding, 445 8.3.2.9 Pediobius foveolatus 9.2.2 Refl ex bleeding substances, 445 (Crawford) (Hymenoptera: 9.2.2.1 Experiments, 445 Eulophidae, 9.2.2.2 Identifi cation, 445 Entedoninae), 409 9.2.2.3 Sources, 445 Detailed contents xv

9.2.2.4 Production cost, 446 9.7 Hibernation and Aggregation, 457 9.2.2.5 Age and stage 9.8 Habitat Preferences: Responses to Plants and modifi cations, 447 Plant Volatiles, 457 9.2.2.6 Species specifi c 9.8.1 Habitat selection, 457 chemistry, 448 9.8.2 Plant stand traits, 457 9.2.3 Relation to enemies and 9.8.3 Avoided plants, 458 competitors, 449 9.8.4 Plant–plant interactions/attractive 9.2.3.1 Parasitoids, 449 plants, 458 9.2.3.2 Ants, 449 9.9 Conclusions and Future Challenges, 459 9.2.3.3 Spiders, 449 References, 460 9.3 Semiochemicals Related to Food, 449 9.3.1 Plant volatiles, 449 10. QUANTIFYING THE IMPACT OF 9.3.1.1 Herbivore-induced plant COCCINELLIDS ON THEIR PREY, 465 volatiles, 449 9.3.1.2 Adult receptors for food J. P. Michaud and James D. Harwood semiochemicals, 450 10.1 Introduction, 466 9.3.1.3 Responses of larvae, 450 10.2 Assays of Consumption, 467 9.3.1.4 Prey sex pheromones: 10.3 Indirect Impacts, 468 predator kairomones, 451 10.4 Traditional Approaches to the Study of 9.3.2 Prey alarm pheromones, 451 Predation, 468 9.3.2.1 Aphid alarm 10.4.1 Selective exclusion, 469 pheromone, 451 10.4.2 Field cages, 469 9.3.2.2 Mechanism for modifi ed 10.4.3 Cage inclusion, 470 responses, 451 10.4.4 Manual removal, 471 9.3.2.3 EBF release: aphid 10.5 Statistical Approaches, 471 individual risks, 451 10.6 Resolving Coccinellid Impact within Complex 9.3.3 Toxic substances in prey, 451 Communities, 472 9.3.4 Feeding stimulants for phytophagous 10.6.1 Multi-species combinations, 472 coccinellids, 452 10.6.2 Demographic-based 9.3.5 Learning, 452 estimates, 473 9.4 Mating and Sex Pheromones, 453 10.7 Post-Mortem Analysis of Predation, 474 9.4.1 Chemoreceptors on the 10.7.1 Antibody-based analysis of antennae, 453 predation, 474 9.4.2 Hydrocarbons on elytra, 454 10.7.2 Protein marking for predation 9.4.3 Ultrastructure of the integumentary analysis, 477 glands, 454 10.7.3 Detection of prey-specifi c 9.5 Oviposition, 454 DNA, 478 9.5.1 Oviposition deterrence: oviposition 10.8 Conclusions, 481 deterrence pheromones, 454 References, 482 9.5.2 Species-specifi city of oviposition deterrence pheromone 11. COCCINELLIDS IN BIOLOGICAL substances, 454 CONTROL, 488 9.5.3 Active substances in oviposition deterrence pheromone J. P. Michaud tracks, 455 11.1 Introduction, 489 9.5.4 Relation to other aphid enemies, 455 11.2 The Roles of Coccinellidae in Biological 9.5.5 Aphid abundance, 456 Control, 489 9.6 Egg and Pupa Protection, 456 11.2.1 Prey specifi city, 489 9.6.1 Protection of eggs, 456 11.2.2 Generalist coccinellids, 490 9.6.2 Protection of pupae, 456 11.2.3 Intraguild predation, 490 xvi Detailed contents

11.3 Scale Versus Aphids as Targets of 11.6.3 Interaction of biological control by Exotic Introductions, 491 coccinellids with plant structure and 11.3.1 Coccidophagous coccinellids, 491 chemistry, 504 11.3.2 Aphidophagous coccinellids, 492 11.6.4 Selective use of pesticides, 506 11.3.3 Invasive coccinellids, 493 11.7 Conclusions, 509 11.3.4 Competitive displacement, 494 References, 509 11.4 Augmentation of Coccinellids, 495 11.4.1 The mealybug destroyer, 496 12. RECENT PROGRESS AND POSSIBLE 11.4.2 Redistribution of FUTURE TRENDS IN THE STUDY OF coccinellids, 496 COCCINELLIDAE, 520 11.4.3 Selection of source material for Helmut F. van Emden and Ivo Hodek augmentation, 497 Acknowledgements, 525 11.4.4 Voltinism and diapause, 497 References, 525 11.4.5 Dietary requirements, 497 11.4.6 Life stages for release, 498 APPENDIX: LIST OF GENERA IN TRIBES 11.5 Conservation, 498 AND SUBFAMILIES, 526 11.5.1 Alternative or supplementary Oldrich Nedveˇd and Ivo Kovárˇ food, 499 Coccinellidae Latreille, 1807, 527 11.5.2 Hibernation refuges, 500 Chilocorinae Mulsant, 1846 [2:25], 527 11.5.3 Habitat management, 500 Coccidulinae Mulsant, 1846 [6:46], 527 11.5.3.1 Strip-harvesting, 501 Coccinellinae Latreille, 1807 [5:94], 528 11.5.3.2 Floral diversity (non-crop Epilachninae Mulsant, 1846 [4:24], 529 plants), 501 Exoplectrinae Crotch, 1874 [2:24], 529 11.5.3.3 Intercropping, 501 Microweiseinae Leng, 1920 [4:23], 529 11.5.3.4 Reduced tillage, 502 Ortaliinae Mulsant, 1850 [2:14], 530 11.6 Ancillary Factors Infl uencing Biological Scymninae Mulsant, 1846 [11:51], 530 Control by Coccinellids, 502 Sticholotidinae Weise, 1901 [6:58], 531 11.6.1 Ant-attendance of aphid colonies, 502 Subject index, 532 11.6.2 Timing of arrival in annual crops, 503 Colour plate pages fall between pp. 250 and pp. 251 Contributors

PIOTR CERYNGIER Centre for Ecological Research, J.P. MICHAUD Department of Entomology, Kansas Polish Academy of Sciences, Dziekanow Lesny, 05 - 092 State University, 1232 240th Ave., Hays, KS 67601 USA Lomianki, POLAND OLDRICH NEDVE ˇ D Faculty of Science, University HELMUT F. VAN EMDEN School of Biological Sci- of South Bohemia and Institute of Entomology, Academy ences, University of Reading, Whiteknights, Reading RG6 of Sciences, CZ 37005 C ˇ esk é Bud eˇ jovice, CZECH 6AS UK REPUBLIC

EDWARD W. EVANS Department of Biology, Utah JAN PETTERSSON Department of Ecology, Swedish State University, Logan, UT 84322 USA University of Agricultural Sciences, Box 7044, SE - 750 07 Uppsala, SWEDEN JAMES D. HARWOOD Department of Entomology, University of Kentucky, Lexington, KY 40546 - 0091 REMY L. POLAND Department of Genetics, Univer- USA sity of Cambridge, Cambridge CB2 3EH UK , recent address: Clifton College, 32 College Road, Clifton, Bristol, IVO HODEK Institute of Entomology, Academy of Sci- BS8 3JH, UK ences, CZ 37005 Cˇ esk é Bud eˇjovice, HELEN E. ROY NERC Centre for Ecology and Hydrol- ALOIS HON Eˇ K Department of Entomology, Crop ogy, Crowmarsh Gifford, Oxfordshire OX10 8BB UK Research Institute, CZ 16106 Prague 6, CZECH REPUBLIC JOHN J. SLOGGETT Maastricht Science Programme, Maastricht University, P.O. Box 616, 6200 MD Maas- IVO KOVÁR ˇ Emer. Scientist of the National Museum, tricht, THE Prague; Current address: Zichovec, CZECH REPUBLIC

ERIC LUCAS D é partement des Sciences Biologiques, Universit é du Qu é bec à Montr é al, C.P. 8888 Succ. Centre - ville, Montr é al, Qu é bec H3C 3P8 CANADA

xvii Preface

For more than a decade, no volume on the general (for parasites and pathogens in Chapter 8 ), who fulfi lled aspects of ladybirds has been published, although the their task with great success. Many thanks ! predaceous major part of this coleopteran family We should like to thank the other authors of chap- represents an important component of the natural ters for contributing their expertise so readily and for enemies of Sternorrhyncha (aphids, coccids, aleyro- reacting so positively and constructively to our dids and psyllids). These sucking insects are among suggestions for revision. We are also grateful for the most dangerous pests of crops, as under suitable the expert help we have received with taxonomic conditions their populations increase exponentially, nomenclature: Dr O. Nedve ˇd of the University of South especially when parthenogenesis and viviparity Bohemia, Czech Republic, and from the UK Drs V.F. occur. Eastop, C.H. Lyal and D.J. Williams (Natural History Although classical biological control with coccinel- Museum), Drs R.T.V. Fox and S.L.Jury (University of lids has recently been mostly abandoned, the increasing Reading) and Professor D.L.J. Quicke (Imperial College). concern about chemical pest control has increased the Dr Lyal deserves additional thanks for his advice in need for modern types of biological control, mostly relation to Chapter 1 . involving conservation and augmentation, within We should also like to thank Dr Ward Cooper of the framework of integrated pest management. For Wiley - Blackwell for enthusiastically agreeing to the success of these sophisticated methods of control, publish our book and Mr Kelvin Matthews for his help precise knowledge of behaviour and ecological rela- with the publication process. We should also like to tions is indispensable. thank the other Wiley- Blackwell staff who have worked Such knowledge, particularly in the areas of ethol- so helpfully and effi ciently on the production of this ogy and molecular genetics, has accumulated in the book. last decade and reviewing these areas is an important novel contribution of this book. We hope that, at this Ivo Hodek stage, it will help to improve conservation and aug- C ˇesk é Bude ˇjovice, Czech Republic mentation control, but also that it will stimulate further research. Helmut F. van Emden The book was very unfortunately interrupted by Reading, UK the premature death of Professor Michael Majerus, Department of Genetics, Cambridge University, who Alois Hon eˇk was our proposed author for Chapters 2 and 8 . We Prague, Czech Republic were extremely lucky to fi nd very able successors, John Sloggett (Chapter 2 ), and Helen Roy and Remy Poland March 2012

xviii Introduction

The reader may fi nd the following information helpful have usually not changed, and the older names are also in order to use the book more easily. listed in the glossary, any confusion will be avoided. To keep the fl ow of the text unbroken and to save The very common generic names are always abbre- repetition and space, taxonomic affi liations of organ- viated as follows throughout the text of all chapters: A . isms and species authorities are given only in the Taxo- for Adalia ; C. , Coccinella ; Cer. , Ceratomegilla ; Chil. , Chilo- nomic Glossary (following section, while the Subject corus ; Col. , Coleomegilla ; Har. , Harmonia ; Hip. , Hippoda- Index is at the end of book, as usual). Readers should mia ; P. , Propylea . also note that the Latin names as given in this glossary Some phenomena are discussed in more than one are used in the text and tables, and therefore may not chapter in the book, but from different angles; the be the same as the older Latin names given in the origi- reader’ s attention is directed to this by cross- references nal papers cited. However, because the species names to the section number.

xix Taxonomic Glossary

With common names where appropriate Anatis quindecimpunctata (DeGeer) Anegleis cardoni (Weise) [Anisolemnia dilatata (F.)] = Megalocaria dilatata (F.) Coleoptera: Family Coccinellidae Anisolemnia tetrasticta Fairmaire Anisosticta bitriangularis (Say) Synonyms [in square brackets] as well as currently Anisosticta novemdecimpunctata (L.) – water ladybird valid names are listed in alphabetical order. Anisosticta sibirica Bielawski Aphidecta obliterata (L.) – larch ladybird Adalia bipunctata (L.) – two spot ladybird [Adalia Apolinus lividigaster (Mulsant) [ Scymnodes lividigaster fasciatopunctata (Faldermann)] (Mulsant)] Adalia conglomerata (L.) Axinoscymnus cardilobus Ren & Pang Adalia decempunctata (L.) – ten spot ladybird [ Azya trinitatis (Marshall)] = Pseudoazya trinitatis Adalia defi ciens Mulsant (Marshall) [Adalia fasciatopunctata (Faldermann)] = Adalia Azya orbigera Mulsant bipunctata (L.) Brachiacantha quadripunctata Melsheimer [Adalia fl avomaculata (De Geer)] = Lioadalia Brachiacantha ursina (F.) fl avomaculata (De Geer) Brumoides septentrionis (Weise) Adalia tetraspilota (Hope) Brumoides suturalis (F.) – three - striped lady [Adonia ] = Hippodamia [Brumus suturalis (Mani)] [Adonia arctica (Schneider)] = Hippodamia arctica [Brumus quadripustulatus (L.)] = Exochomus (Schneider) quadripustulatus (L.) [Adonia variegata (Goeze)] = Hippodamia variegata [Brumus suturalis (Mani)] = Brumoides suturalis (F.) (Goeze) Bulaea lichatschovi (Hummel) Afi denta misera (Weise) [ Afi denta mimetica Dieke] Callicaria superba (Mulsant) Aphidentula bisquadripunctata (Gyllenhal) [ Epilachna Calvia albida Bielawski bisquadripunctata (Gyllenhal)] Calvia decemguttata (L.) Afi ssula rana Kapur Calvia duodecimmaculata Gebler Afi ssula sanscrita (Crotch) Calvia muiri Timberlake [Eocaria muiri Timberlake] Aiolocaria hexaspilota (Hope) [ Aiolocaria mirabilis Calvia quatuordecimguttata (L.) – cream - spot ladybird (Motschulsky)] Calvia quindecimguttata (F.) [Aiolocaria mirabilis (Motschulsky)] = Aiolocaria Calvia shiva Kapur hexaspilota (Hope) Calvia shiva pasupati Kapur Alloneda dodecaspilota (Hope) Calvia shiva pinaki Kapur Anatis halonis Lewis Calvia shiva trilochana Kapur Anatis labiculata (Say) – fi fteen spotted lady beetle Ceratomegilla barovskii kiritschenkoi (Semenov - Tian - Anatis mali (Say) – eyespotted lady beetle Shanski) [Spiladelpha barovskii kiritschenkoi Anatis ocellata (L.) – eyed ladybird Semenov - Tian - Shanski] xx Taxonomic glossary xxi

Ceratomegilla notata (Laicharting) Coccinella hieroglyphica L. – hieroglyphic ladybird Ceratomegilla undecimnotata (Schneider) [ Semiadalia Coccinella leonina transversalis F. [Coccinella repanda undecimnotata (Schneider)] Thunberg, Coccinella transversalis F.] Cheilomenes lunata (F.) Coccinella luteopicta (Mulsant) Cheilomenes propinqua vicina (Mulsant) [ Cheilomenes Coccinella magnifi ca Redtenbacher – scarce seven spot vicina (Mulsant), Cydonia vicina nilotica Mulsant] ladybird [Coccinella divaricata Olivier)] [Cheilomenes sexmaculata (F.)] = Menochilus Coccinella monticola Mulsant sexmaculatus (F.) Coccinella nigrovittata Kapur [ Tytthaspis trilineata Cheilomenes sulphurea (Olivier) (Weise)] [Cheilomenes vicina ] = Cheilomenes propinqua vicina Coccinella novemnotata Herbst – nine - spotted lady (Mulsant) beetle [Chilocorus baylei (Blackburn)] = Chilocorus malasiae Coccinella quinquepunctata L. – fi ve spot ladybird Crotch Coccinella reitteri Weise Chilocorus bijugus Mulsant [Coccinella repanda Thunberg] = Coccinella leonina F. Chilocorus bipustulatus (L.) – heather ladybird Coccinella septempunctata L. – seven spot ladybird Chilocorus braeti Weise Coccinella septempunctata algerica Kov á rˇ [Coccinella Chilocorus cacti (L.) algerica Kov á rˇ ] Chilocorus circumdatus (Gyllenhal) Coccinella septempunctata brucki Mulsant Chilocorus discoideus Crotch [Coccinella sinuatomarginata Faldermann] = Coccinula Chilocorus distigma Klug sinuatomarginata (Faldermann) Chilocorus geminus Zaslavsky [Coccinella transversalis F.] = Coccinella leonina Chilocorus hauseri Weise transversalis F. Chilocorus hexacyclus Smith Coccinella transversoguttata Faldermann – transverse Chilocorus infernalis Mulsant lady beetle Chilocorus inornatus Weise Coccinella transversoguttata richardsoni Brown Chilocorus kuwanae Silvestri Coccinella trifasciata L. – three - banded lady beetle Chilocorus malasiae Crotch [ Chilocorus baylei Coccinella undecimpunctata L. – eleven spot ladybird (Blackburn)] [Coccinella undecimpunctata aegyptiaca Chilocorus nigripes Mader Reiche] = Coccinella undecimpunctata menetriesi Chilocorus nigritus (F.) Mulsant Chilocorus orbus Casey Coccinella undecimpunctata menetriesi Mulsant Chilocorus quadrimaculatus (Weise) [Coccinella undecimpunctata aegyptiaca Reiche] Chilocorus renipustulatus (Scriba) – kidney - spot Coccinula crotchi (Lewis) ladybird Coccinula quatuordecimpustulata (L.) Chilocorus rubidus Hope Coccinula redimita (Weise) Chilocorus similis (Rossi) Coccinula sinensis (Weise) Chilocorus stigma (Say) – twice - stabbed lady beetle Coccinula sinuatomarginata (Faldermann) [ Coccinella Chilocorus tricyclus Smith sinuatomarginata Faldermann] Chnootriba similis (Thunberg) Coelophora biplagiata Swartz [ Lemnia biplagiata Cleobora mellyi (Mulsant) – Tasmanian ladybird (Swartz)] Clitostethus arcuatus (Rossi) Coelophora bissellata Mulsant Clitostethus oculatus (Blatchley) [ Nephaspis oculatus Coelophora duvaucelii (Mulsant) (Blatchley)] Coelophora inaequalis (F.) – common Australian lady Coccidophilus citricola Br è thes beetle Coccidula rufa (Herbst) Coelophora mulsanti (Montrouzier) Coccidula scutellata (Herbst) Coelophora quadrivittata Fauvel [Coccinella algerica Kov á rˇ ] = Coccinella septempunctata Coelophora saucia Mulsant algerica Kov á rˇ Coleomegilla maculata (DeGeer) – spotted lady Coccinella californica Mannerheim beetle Coccinella explanata Miyatake Coleomegilla maculata fuscilabris (Mulsant) xxii Taxonomic glossary

Coleomegilla maculata lengi Timberlake – twelve Epilachna karisimbica Weise spotted ladybeetle Epilachna marginella (F.) Coleomegilla quadrifasciata (Schoenherr) Epilachna marginicollis (Hope) Cryptognatha simillima Sicard Epilachna mexicana (Gu é rin - M é neville) Cryptognatha nodiceps Marshall Epilachna mystica Mulsant Cryptognatha signata Korschefsky Epilachna nigrolimbata Thomson Cryptogonus ariasi (Mulsant) [Epilachna niponica Lewis] = niponica Cryptogonus kapuri Ghorpade (Lewis) Cryptogonus orbiculus (Gyllenhal) Epilachna paenulata (Germar) Cryptogonus postmedialis Kapur [Epilachna philippinensis Dieke] = Henosepilachna Cryptogonus quadriguttatus (Weise) vigintisexpunctata (Boisduval) Cryptolaemus montrouzieri Mulsant – mealybug [Epilachna pusillanima Mulsant] = Henosepilachna destroyer pusillanima (Mulsant) Curinus coeruleus Mulsant – metallic blue lady beetle [Epilachna pustulosa K ô no] = Henosepilachna pustulosa Cycloneda ancoralis (Germar) (K ô no) Cycloneda limbifer Casey Epilachna quadricollis (Dieke) Cycloneda munda (Say) – polished lady beetle [Epilachna septima Dieke] = Henosepilachna septima Cycloneda polita Casey (Dieke) Cycloneda sanguinea (L.) – blood - red lady beetle [Epilachna sparsa orientalis Dieke] = Henosepilachna [Cydonia vicina nilotica Mulsant] = Cheilomenes vigintioctopunctata (F.) propinqua vicina (Mulsant) Epilachna undecimvariolata (Boisduval) Declivitata spp. Epilachna varivestis Mulsant – mexican bean beetle Delphastus catalinae (Horn) [Epilachna vigintioctomaculata Motschulsky] = Delphastus pusillus (LeConte) Henosepilachna vigintioctomaculata (Motschulsky) Diomus austrinus Gordon [Epilachna vigintioctomaculata Motschulsky] = Diomus hennesseyi F ü rsch Henosepilachna vigintioctomaculata (Motschulsky) Diomus pumilio Weise Epilachna vigintisexpunctata (Boisduval) Diomus seminulus (Mulsant) [Epilachna yasutomii (Katakura)] = Henosepilachna Diomus thoracicus F. yasutomii Katakura [Eocaria muiri Timberlake] = Calvia muiri (Timberlake) Epiverta chelonia (Mader) Epilachna admirabilis Crotch Eriopis connexa (Germar) [Epilachna bisquadripunctata (Gyllenhal)] = Aphidentula Exochomus childreni Mulsant bisquadripunctata (Gyllenhal) [Exochomus concavus F ü rsch] = Parexochomus troberti [Epilachna boisduvali (Mulsant)] = Henosepilachna concavus (F ü rsch) boisduvali Mulsant Exochomus fl avipes (Thunberg) Epilachna borealis (F.) – squash beetle Exochomus fl aviventris Mader Epilachna canina (F.) Exochomus fulvimanus Weise [Epilachna ‘ chrysomelina ’ (F.)]; used variably either for [Exochomus lituratus (Gorham)] = Priscibrumus Henosepilachna argus (Geoffroy) or Henosepilachna lituratus (Gorham) elaterii (Rossi) or Henosepilachna vigintioctopunctata [Exochomus melanocephalus (Zoubkoff)] = Parexochomus (F.) melanocephalus (Zoubkoff) [Epilachna cucurbitae ] = Henosepilachna sumbana [Exochomus nigromaculatus (Goeze) = Parexochomus Bielawski nigromaculatus (Goeze) Epilachna defecta Mulsant Exochomus quadripustulatus (L.) – pine ladybird Epilachna dregei Mulsant – potato ladybird [Brumus quadripustulatus (L.)] Epilachna dumerili Mulsant [Exochomus troberti Mulsant] = Parexochomus troberti Epilachna eckloni Mulsant (Mulsant) [Epilachna enneasticta Mulsant] = Henosepilachna (Boisduval) – steelblue lady beetle enneasticta (Mulsant) [Orcus chalybeus (Boisduval)] Epilachna eusema (Weise) [Halyzia hauseri (Mader)] = Macroilleis hauseri (Mader) Taxonomic glossary xxiii

Halyzia sanscrita Mulsant Hippodamia arctica (Schneider) [ Adonia arctica Halyzia sedecimguttata (L.) – orange ladybird (Schneider)] Halyzia straminea (Hope) Hippodamia caseyi Johnson Halyzia tschitscherini Semenov Hippodamia convergens Guerin – convergent ladybeetle Harmonia antipoda (Mulsant in White) – antipodean Hippodamia glacialis (F.) – glacial lady beetle ladybird Hippodamia parenthesis (Say) – parenthesis lady beetle Harmonia axyridis (Pallas) – harlequin ladybird or Hippodamia quinquesignata (Kirby) Asian multi - colored ladybeetle Hippodamia quinquesignata punctulata Le Conte [Harmonia breiti Mader] = Harmonia expalliata Sicard [Hippodamia quinquesignata ambigua LeConte] Harmonia conformis (Boisduval) Hippodamia septemmaculata (DeGeer) Harmonia dimidiata (F.) [ Leis dimidiata (F.)] Hippodamia sinuata Mulsant Harmonia eucharis (Mulsant) Hippodamia tredecimpunctata (L.) – thirteen spot Harmonia expalliata Sicard [ Harmonia breiti Mader] ladybird Harmonia quadripunctata (Pontoppidan) – cream - Hippodamia variegata (Goeze) – Adonis ladybird, streaked ladybird variegated lady beetle [ Adonia variegata (Goeze)] Harmonia octomaculata (F.) Hyperaspis aestimabilis Mader Harmonia sedecimnotata (F.) Hyperaspis bigeminata (Randall) Harmonia yedoensis (Takizawa) Hyperaspis binotata (Say) Henosepilachna argus (Geoffroy) – bryony ladybird Hyperaspis campestris (Herbst) Henosepilachna bifasciata (L.) [Hyperaspis congressis Watson] = Hyperaspis conviva Henosepilachna boisduvali (Mulsant) [ Epilachna Casey boisduvali Mulsant] Hyperaspis conviva Casey [ Hyperaspis congressis Henosepilachna dodecastigma (Wiedemann) Watson] Henosepilachna elaterii (Rossi) [ Epilachna chrysomelina Hyperaspis desertorum Weise (F.)] Hyperaspis lateralis Mulsant Henosepilachna enneasticta (Mulsant) [ Epilachna Hyperaspis notata Mulsant enneasticta Mulsant] Hyperaspis pantherina F ü rsch Henosepilachna guttatopustulata (F.) Hyperaspis raynevali Mulsant Henosepilachna indica (Mulsant) Hyperaspis reppensis (Herbst) Henosepilachna niponica (Lewis) [ Epilachna niponica Hyperaspis senegalensis Mulsant Lewis] Hyperaspis senegalensis hottentotta Mulsant Henosepilachna ocellata (Redtenbacher) Hyperaspis sphaeridioides Mulsant Henosepilachna processa Li Hyperaspis undulata (Say) Henosepilachna pusillanima (Mulsant) [ Epilachna Illeis bielawskii Ghorpade pusillanima Mulsant] Illeis cincta (F.) Henosepilachna pustulosa (K ô no) [Epilachna pustulosa Illeis galbula (Mulsant) [ Leptothea galbula (Mulsant)] K ô no] Illeis koebelei Timberlake Henosepilachna septima (Dieke) [Epilachna septima Dieke] Jauravia quadrinotata Kapur Henosepilachna sumbana Bielawski [ Epilachna [Leis dimidiata (F.)] = Harmonia dimidiata (F.) cucurbitae Richards, Henosepilachna cucurbitae [Leis ] = Harmonia Richards] [Lemnia biplagiata (Swartz)] = Coelophora biplagiata Henosepilachna vigintioctomaculata (Motschulsky) Swartz [Epilachna vigintioctomaculata Motschulsky] [Leptothea galbula (Mulsant)] = Illeis galbula Henosepilachna vigintioctopunctata (F.) [ Epilachna (Mulsant) vigintioctopunctata (F.), Epilachna sparsa orientalis Lindorus lophanthae (Blaisdell) [ Rhyzobius lophantae Dieke] (Blaisdell), Rhyzobius lorophantae (Blaisdell)] Henosepilachna vigintisexpunctata (Boisduval) Lioadalia fl avomaculata (De Geer) [Adalia fl avomaculata [Epilachna philippinensis Dieke] (De Geer)] Henosepilachna yasutomii Katakura [ Epilachna Macroilleis hauseri (Mader) [ Halyzia hauseri (Mader)] yasutomii (Katakura)] Macronaemia hauseri (Weise) xxiv Taxonomic glossary

Megalocaria dilatata (F.) [ Anisolemnia dilatata (F.)] Parastethorus nigripes (Kapur) [ Stethorus loxtoni [Menochilus quadriplagiatus (Swartz)] = Menochilus Britton & Lee] sexmaculatus (F.) Parexochomus melanocephalus (Zoubkoff) [ Exochomus Menochilus sexmaculatus (F.) [ Cheilomenes sexmaculata melanocephalus (Zoubkoff)) (F.), Menochilus quadriplagiatus (Swartz)] Parexochomus nigromaculatus (Goeze) [ Exochomus Micraspis allardi (Mulsant) nigromaculatus (Goeze)] Micraspis discolor (F.) Parexochomus troberti (Mulsant) [ Exochomus troberti Microweisea sp. (Mulsant)] Mulsantina hudsonica (Casey) Parexochomus troberti concavus (Fuersch) [ Exochomus Mulsantina picta (Randall) concavus Fuersch] Myrrha octodecimguttata (L.) – eighteen spot ladybird Pentilia insidiosa Mulsant Myzia oblongoguttata (L.) – striped ladybird Pharoscymnus anchorago (Fairmaire) Myzia subvittata (Mulsant) Pharoscymnus numidicus (Pic) Neda marginalis Mulsant Pharoscymnus ovoideus Sicard Neocalvia anastomozans Crotch Phymatosternus lewisii (Crotch) [Nephaspis oculatus (Blatchley)] = Clitostethus oculatus Platynaspis luteorubra (Goeze) (Blatchley) Priscibrumus lituratus (Gorham) [Exochomus lituratus Nephus bilucernarius (Mulsant) (Gorham)] Nephus bisignatus (Boheman) Priscibrumus uropygialis (Mulsant) Nephus fl avifrons (Melsheimer) [Scymnus fl avifrons Propylea dissecta (Mulsant) Melsheimer, North America, not Scymnus fl avifrons Propylea japonica (Thunberg) Blackburn, Australia] Propylea quatuordecimpunctata (L.) – fourteen spot Nephus guttulatus (LeConte) [Scymnus guttulatus ladybird [Propylaea quatuordecimpunctata (L.)] LeConte] Pseudoazya trinitatis (Marshall) [ Azya trinitatis Nephus includens (Kirsch) (Marshall)] Nephus kiesenwetteri (Mulsant) [Scymnus kiesenwetteri [Pseudoscymnus tsugae ] Sasaji & McClure] = Mulsant] Sasajiscymnus tsugae (Sasaji & McClure) Nephus ornatus (LeConte) [Scymnus ornatus LeConte] [Pseudoscymnus kurohime (Mityake)] = Sasajiscymnus Nephus quadrimaculatus (Herbst) [Scymnus kurohime (Miyatake) quadrimaculatus (Herbst)] Psyllobora confl uens (F.) Nephus redtenbacheri (Mulsant) Psyllobora vigintiduopunctata (L.) – twenty two spot Nephus soudanensis (Sicard) [Scymnus soudanensis ladybird [Thea vigintiduopunctata (L.)] Sicard] Psyllobora vigintimaculata (Say) – twenty - spotted lady Oenopia billieti (Mulsant) beetle Oenopia conglobata (L.) [ Synharmonia conglobata (L.)] [Pullus auritus (Thunberg)] = Scymnus auritus Oenopia kirbyi Mulsant Thunberg Oenopia lyncea (Olivier) [Pullus mediterraneus (F.)] = Scymnus marinus Oenopia sexareata (Mulsant) (Mulsant) [Olla abdominalis (Say)] = Olla v - nigrum (Mulsant) [Pullus subvillosus (Goeze)] = Scymnus subvillosus Olla v - nigrum (Mulsant) – ash- gray lady beetle [Olla (Goeze) abdominalis (Say)] Rhyzobius litura (F.) Orcus australasiae (Boisduval) [Rhyzobius lophanthae (Blaisdell)] = Lindorus [Orcus chalybeus (Boisduval)] = Halmus chalybeus lophanthae (Blaisdell) (Boisduval) [Rhyzobius lorophanthae (Blaisdell)] = Lindorus Palaeoneda auriculata (Mulsant) [Paleoneda miniata lophanthae (Blaisdell) (Hope)] Rhyzobius ventralis (Erichson) – black lady beetle [Paleoneda miniata (Hope)] = Palaeoneda auriculata Rodatus major (Blackburn) (Mulsant) Rodolia cardinalis (Mulsant) – vedalia beetle Pania luteopustulata Mulsant Rodolia fumida Mulsant Paranaemia vittigera (Mannerheim) Rodolia guerini (Crotch) Taxonomic glossary xxv

Rodolia iceryae Janson Scymnus pyrocheilus Mulsant Rodolia occidentalis Weise Scymnus quadrillum Motschulsky Sasajiscymnus kurohime (Miyatake) [ Pseudoscymnus [Scymnus (Nephus) quadrimaculatus (Herbst)] = Nephus kurohime (Miyatake)] quadrimaculatus (Herbst) [Sasajiscymnus ningshanensis (Sasaji & McClure)] = Scymnus rubromaculatus (Goeze) Scymnus ningshanensis Yu & Yao Scymnus sinuanodulus Yu & Yao Sasajiscymnus tsugae (Sasaji & McClure) Scymnus smithianus Silvestri [Pseudoscymnus tsugae Sasaji & McClure] Scymnus soudanensis Sicard [Scymnodes lividigaster (Mulsant)] = Apolinus Scymnus subvillosus (Goeze) [ Pullus subvillosus lividigaster (Mulsant) (Goeze)] Scymnus abietis (Paykull) Scymnus suturalis Thunberg [Scymnus aeneipennis Sicard] = Zagloba aeneipennis Scymnus syriacus (Marsuel) (Sicard) Scymnus tardus Mulsant Scymnus apetzi Mulsant [Semiadalia undecimnotata (Schneider)] = Ceratomegilla Scymnus ater Kugelann undecimnotata (Schneider) Scymnus auritus Thunberg [Pullus auritus (Thunberg)] Serangium parcesetosum Sicard Scymnus coccivora Ayyar [Sidis ] = Nephus Scymnus creperus Mulsant [Spiladelpha barovskii kiritschenkoi Semenov - Tian - Scymnus dorcatomoides Weise Shanski] = Ceratomegilla barovskii kiritschenkoi Scymnus fl avifrons Blackburn (Australia) (Semenov - Tian - Shanski) [Scymnus fl avifrons Melsheimer] (North Sospita vigintiguttata (L.) America) = Nephus fl avifrons (Melsheimer) Stethorus bifi dus Kapur Scymnus frontalis (F.) Stethorus gilvifrons (Mulsant) [Scymnus guttulatus LeConte] = Nephus guttulatus Stethorus japonicus Kamiya (LeConte) [ Stethorus loxtoni Britton & Lee] = Parastethorus Scymnus haemorrhoidalis Herbst nigripes (Kapur) Scymnus hilaris Motschulsky Stethorus madecassus Chazeau Scymnus hoffmanni Weise [Stethorus picipes Casey] = Stethorus punctum picipes Scymnus impexus Mulsant Casey Scymnus interruptus (Goeze) [Stethorus punctillum Weise] = Stethorus pusillus [Scymnus kiesenwetteri Mulsant] = Nephus (Herbst) kiesenwetteri (Mulsant) Stethorus punctum (LeConte) – spider - mite destroyer Scymnus lacustris LeConte Stethorus punctum picipes Casey [ Stethorus picipes Scymnus levaillanti Mulsant Casey] Scymnus loewii Mulsant – dusky lady beetle Stethorus pusillus (Herbst) [ Stethorus punctillum Scymnus louisianae Chapin (Weise)] Scymnus marginicollis Mannerheim Stethorus tridens Gordon Scymnus marinus (Mulsant) [ Scymnus mediterraneus Stethorus vegans (Blackburn) Iablokoff - Khnzorian, Pullus mediterraneus (Iablokoff- Subcoccinella vigintiquatuorpunctata (L.) – twenty four Khnzorian)] spot ladybird [Scymnus mediterraneus ] = Scymnus marinus (Mulsant) [Synharmonia conglobata (L.)] = Oenopia conglobata (L.) Scymnus morelleti Mulsant Synona obscura Poorani, S ´ lipi n´ ski & Booth Scymnus nigrinus Kugelann Synonycha grandis (Thunberg) Scymnus ningshanensis Yu & Yao [Sasajiscymnus Thalassa saginata Mulsant ningshanensis (Sasaji & McClure)] [Thea vigintiduopunctata (L.)] = Psyllobora [Scymnus ornatus LeConte] = Nephus ornatus vigintiduopunctata (L.) (LeConte) Tytthaspis sedecimpunctata (L.) – sixteen spot ladybird Scymnus otohime Kamiya [Tytthaspis trilineata (Weise)] = Coccinella nigrovittata Scymnus posticalis Sicard Kapur Scymnus postpictus Casey [Verania ] = Micraspis xxvi Taxonomic glossary

Vibidia duodecimguttata (Poda) Nilaparvata lugens (St å l) (Delphacidae) – brown Zagloba aeneipennis (Sicard) [ Scymnus aeneipennis planthopper Sicard] Philaenus spumarius (L.) (Cercopidae) – common froghopper or meadow spittlebug OTHER ORGANISMS Hemiptera: Sternorrhyncha: Aphidoidea In alphabetical order within each taxon, and with Family given in italics within brackets after the (Synonyms are in square brackets) authority Acyrthosiphon caraganae (Cholodkovsky) (Aphididae) OTHER INSECTS Acyrthosiphon ignotum Mordvilko (Aphididae) Acyrthosiphon kondoi Shinji (Aphididae) – blue alfalfa Dermaptera aphid [Acyrthosiphon nipponicum (Essig & Kuwana)] = Anechura harmandi (Burr) (Forfi culidae) – hump Neoaulacorthum nipponicum (Essig & Kuwana) earwig (Aphididae) Forfi cula auricularia L. (Forfi culidae) – European Acyrthosiphon pisum (Harris) (Aphididae) – pea aphid earwig Adelges cooleyi (Gillette) (Adelgidae) – Cooley spruce gall adelgid Adelges laricis Vallot (Adelgidae) – larch adelgid Hemiptera: Heteroptera Adelges nordmannianae (Eckstein) (Adelgidae) [Adelges nusslini (Boerner)] = Adelges nordmannianae Calocoris norvegicus (Gmelin) (Miridae) – strawberry (Eckstein) (Adelgidae) bug Adelges piceae (Ratzeburg) (Adelgidae) – balsam woolly Campylomma verbasci (Meyer) (Miridae) – mullein bug adelgid Caternaultiella rugosa (Schoutenden) (Plataspidae) Adelges tsugae Annand (Adelgidae) – hemlock woolly Eurygaster integriceps Puton (Pentatomidae) – sunn adelgid pest or corn bug Aphis carduella Walsh (Aphididae) Geocoris punctipes (Say) (Lygaeidae) – big - eyed bug [Aphis cirsiiacanthoidis Boerner] = Aphis fabae Hyaliodes vitripennis (Say) (Miridae) cirsiiacanthoidis Scopoli (Aphididae) Lygus hesperus (Knight) (Miridae) – western plant Aphis craccivora Koch (Aphididae) – cowpea aphid or bug groundnut aphid Lygus lineolaris (Palidot de Beauvois) (Miridae) Aphis cytisorum cytisorum Hartig (Aphididae) – tarnished plant bug Aphis fabae Scopoli (Aphididae) – black bean aphid Lygus (Miridae) Aphis fabae cirsiiacanthoidis Scopoli (Aphididae) Nabis (Reduviolus) americoferus Carayon (Nabidae) Aphis farinosa J.F.Gmelin (Aphididae) – common damsel bug Aphis glycines Matsumura (Aphididae) – soybean aphid Nysius huttoni White (Lygaeidae) – wheat bug Aphis gossypii Glover (Aphididae) – cotton aphid or Orius insidiosus (Say) (Anthocoridae) – insidious fl ower melon aphid bug Aphis hederae Kaltenbach (Aphididae) Podisus maculiventris (Say) (Pentatomidae) – spined [Aphis helianthi Monell] = Aphis carduella Walsh soldier bug (Aphididae) Pyrrhocoris apterus (L.) (Pyrrhocoridae) – fi rebug Aphis jacobaeae Schrank (Aphididae) Sidnia kinbergi (St å l) (Miridae) [Aphis laburni Kaltenbach] = Aphis cytisorum cytisorum Hartig (Aphididae) Hemiptera: Auchenorrhyncha Aphis nerii Boyer de Fonscolombe (Aphididae) – oleander aphid Homalodisca vitripennis (Germar) (Cicadellidae) Aphis pomi De Geer (Aphididae) – green apple aphid – glassy - winged sharpshooter Aphis punicae Passerini (Aphididae) Taxonomic glossary xxvii

Aphis sambuci L. (Aphididae) – elder aphid Eucallipterus tiliae (L.) (Aphididae) – lime aphid Aphis spiraecola Patch (Aphididae) – spiraea aphid or Euceraphis betulae (Koch) (Aphididae) green citrus aphid Euceraphis punctipennis (Zetterstedt) (Aphididae) Aphis spiraephaga F.P. M ü ller (Aphididae) Hyalopterus pruni (Geoffroy) (Aphididae) – mealy plum Aphis spiraephila Patch (Aphididae) aphid Aphis urticata J.F. Gmelin (Aphididae) Hyperomyzus carduellinus (Theobald) (Aphididae) [Aulacorthum magnoliae (Essig & Hyperomyzus lactucae (L.) (Aphididae) – blackcurrant – Kuwana)] = Neoaulacorthum magnoliae (Essig & sowthistle aphid Kuwana) (Aphididae) Laingia psammae Theobald (Aphididae) Aulacorthum solani (Kaltenbach) (Aphididae) – Liosomaphis berberidis (Kaltenbach) (Aphididae) glasshouse potato aphid or foxglove aphid Lipaphis pseudobrassicae (Davis) (Aphididae) – turnip Betulaphis brevipilosa Boerner (Aphididae) aphid or mustard aphid Betulaphis quadrituberculata (Kaltenbach) (Aphididae) [Longiunguis donacis (Passerini)] = Melanaphis donacis Brachycaudus helichrysi (Kaltenbach) (Aphididae) (Passerini) (Aphididae) – leaf - curling plum aphid Macrosiphoniella artemisiae (Boyer de Fonscolombe) Brachycaudus persicae (Passerini) (Aphididae) – black (Aphididae) peach aphid Macrosiphoniella sanborni (Gillette) (Aphididae) Brachycaudus prunicola (Kaltenbach) (Aphididae) – chrysanthemum aphid Brachycaudus tragopogonis (Kaltenbach) (Aphididae) Macrosiphum albifrons Essig (Aphididae) – lupin aphid Brevicoryne brassicae (L.) (Aphididae) – cabbage aphid Macrosiphum euphorbiae (Thomas) (Aphididae) Callipterinella calliptera (Hartig) (Aphididae) – potato aphid Capitophorus elaeagni (Del Guercio) (Aphididae) [Macrosiphum ibarae (Matsumura)] = Sitobion ibarae Cavariella konoi Takahashi (Aphididae) (Matsumura) (Aphididae) Ceratovacuna lanigera Zehntner(Aphididae) – sugar Macrosiphum rosae (L.) (Aphididae) – rose aphid cane woolly aphid Megoura viciae Buckton (Aphididae) – vetch aphid Cervaphis quercus Takahashi (Aphididae) Melanaphis donacis (Passerini) (Aphididae) Chaetosiphon fragaefolii (Cockerell) (Aphididae) Metopolophium dirhodum (Walker) (Aphididae) – strawberry aphid – rose – grain aphid. Chaitophorus capreae (Mosley) (Aphididae) Metopolophium festucae (Theobald) (Aphididae) Chaitophorus leucomelas Koch (Aphididae) – fescue aphid [Chaitophorus versicolor Koch] = Chaitophorus Microlophium carnosum (Buckton) (Aphididae) leucomelas Koch (Aphididae) Microsiphoniella artemisiae (Gillette) (Aphididae) Chromaphis juglandicola (Kaltenbach) (Aphididae) Mindarus abietinus Koch (Aphididae) – walnut aphid Myzocallis boerneri Stroyan (Aphididae) Cinara palaestinensis Hille Ris Lambers (Aphididae) Myzocallis carpini (Koch) (Aphididae) Delphiniobium junackianum (Karsch) (Aphididae) Myzocallis castanicola Baker (Aphididae) Diuraphis noxia (Kurdjumov) (Aphididae) – Russian Myzocallis coryli (Goetze) (Aphididae) – hazel aphid or wheat aphid fi lbert aphid Drepanosiphum platanoidis (Schrank) (Aphididae) Myzus cerasi (F.) (Aphididae) – cherry blackfl y – sycamore aphid Myzus persicae (Sulzer) (Aphididae) – peach – potato Dysaphis crataegi (Kaltenbach) (Aphididae) – aphid hawthorn – parsnip aphid Myzus persicae nicotianae Blackman (Aphididae) Dysaphis devecta (Walker) (Aphididae) – rosy leaf - Neoaulacorthum magnoliae (Essig & Kuwana) curling aphid (Aphididae) Dysaphis plantaginea (Passerini) (Aphididae) – rosy Neoaulacorthum nipponicum (Essig & Kuwana) apple aphid (Aphididae) Elatobium abietinum (Walker) (Aphididae) – spruce Neomyzus circumfl exus (Buckton) (Aphididae) aphid Neophyllaphis podocarpi Takahashi (Aphididae) Eriosoma lanigerum (Hausmann) (Aphididae) – woolly Periphyllus californiensis (Shinji) (Aphididae) apple aphid Periphyllus lyropictus (Kessler) (Aphididae) xxviii Taxonomic glossary

Periphyllus testudinaceus (Fernie) (Aphididae) Agonoscena pistaciae Burckhardt et Lauterer (Psyllidae) Phorodon humuli (Schrank) (Aphididae) – damson – hop Aleurodicus cocois (Curtis) (Aleyrodidae) aphid Aleurodicus dispersus Russell (Aleyrodidae) – spiralling Phyllaphis fagi (L.) (Aphididae) whitefl y Phylloxera glabra (von Heyden) (Phylloxeridae) Aleurotuba jelinekii (Frauenfeld) (Aleyrodidae) Pineus pini (Macquart) (Adelgidae) Aleyrodes proletella (L.) (Aleyrodidae) Pseudoregma alexanderi (Takahashi) (Aphididae) Aonidiella aurantii (Maskell) (Diaspididae) – California Pseudoregma bambucicola (Takahashi) (Aphididae) red scale Pterocallis alni (deGeer) (Aphididae) Aonidiella orientalis (Newstead) (Diaspididae) Rhopalosiphum maidis (Fitch) (Aphididae) – corn leaf Aonidimytilus albus (Cockerell) (Diaspididae) – cassava aphid scale Rhopalosiphum padi (L.) (Aphididae) – bird cherry – oat Aspidiotus destructor Signoret (Diaspididae) – coconut aphid scale Schizaphis graminum (Rondani) (Aphididae) – Aspidiotus nerii Bouch é (Diaspididae) greenbug Asterolecanium sp. (Asterolecaniidae) Schizolachnus pineti (F. ) (Aphididae) Aulacaspis tegalensis (Zehntner) (Diaspididae) – sugar Schizolachnus piniradiatae (Davidson) (Aphididae) cane scale Sitobion akebiae (Shinji) (Aphididae) Aulacaspis tubercularis Newstead (Diaspididae) Sitobion avenae (F.) (Aphididae) – grain aphid Bemisia tabaci (Gennadius) (Aleyrodidae) Sitobion ibarae (Matsumura) (Aphididae) [Bemisia argentifolii Bellows] = Bemisia tabaci Symydobius oblongus (von Heyden) (Aphididae) Gennadius (Aleyrodidae) Thelaxes dryophila (Schrank) (Aphididae) Chionaspis alnus Kuwana (Diaspididae) [Therioaphis maculata (Buckton)] = Therioaphis trifolii Chionaspis salicis (L.) (Diaspididae) (Monell) (Aphididae) – spotted alfalfa aphid Chrysomphalus aonidum (L.) (Diaspididae) – Florida red Therioaphis trifolii (Monell) (Aphididae) – spotted scale alfalfa aphid Chrysomphalus bifasciculatus Ferris (Diaspididae) Toxoptera aurantii (Boyer de Fonscolombe) (Aphididae) Coccus hesperidum L.(Coccidae) – soft brown scale – black citrus aphid or tea aphid Coccus viridis (Green) (Coccidae) – green coffee scale Toxoptera citricidus (Kirkaldy) (Aphididae) – brown or soft green scale citrus aphid Dactylopius opuntiae (Cockerell) (Dactylopiidae) [Toxoptera graminum (Rondani)] = Schizaphis graminum Diaphorina citri Kuwayama (Psyllidae) – Asian citrus (Rondani) (Aphididae) – greenbug psyllid Tuberculatus annulatus (Hartig) (Aphididae) Diaspidiotus perniciosus (Comstock) (Diaspididae) Tuberolachnus salignus (J.F. Gmelin) (Aphididae) Dysmicoccus (Pseudococcidae) – willow aphid Eriococcus coriaceus Maskell (Eriococcidae) Uroleucon aeneum (Hille Ris Lambers) (Aphididae) Eulecanium caraganae Borchsenius (Coccidae) Uroleucon ambrosiae (Thomas) (Aphididae) Ferrisia virgata (Cockerell) (Pseudococcidae) – striped Uroleucon cichorii (Koch) (Aphididae) mealybug Uroleucon cirsii (L.) (Aphididae) Hemberlesia lataniae (Signoret) (Diaspididae) Uroleucon compositae (Theobald) (Aphididae) Heteropsylla cubana Crawford (Psyllidae) Uroleucon formosanum (Takahashi) (Aphididae) Icerya purchasi Maskell (Monophlebidae) – cottony Uroleucon jaceae (L.) (Aphididae) cushion scale Uroleucon nigrotuberculatum (Olive) (Aphididae) Lepidosaphes beckii (Newman) (Diaspididae) – citrus Vesiculaphis caricis (Fullaway) (Aphididae) mussel scale Lepidosaphes cornutus Ramakrishna Ayyar (Diaspididae) Hemiptera: other Sternorrhyncha Lepidosaphes ulmi (L.) (Diaspididae) – mussel scale or oystershell scale Abgrallaspis cyanophylli (Signoret) (Diaspididae) Maconellicoccus hirsutus (Green) (Pseudococcidae) Acutaspis umbonifera (Newstead) (Diaspididae) – pink hibiscus mealybug Taxonomic glossary xxix

Matsucoccus feytaudi Ducasse (Matsucoccidae) Chrysopa perla (L.) (Chrysopidae) Matsucoccus josephi Bodenheimer & Harpaz Chrysopa sinica (Tjeder) (Chrysopidae) (Matsucoccidae) Chrysoperla carnea (Stephens) (Chrysopidae) – green Matsucoccus matsumurae (Kuwana) (Matsucoccidae) lacewing Melanaspis glomerata (Green) (Diaspididae) Chrysoperla plorabunda Fitch (Chrysopidae) Monophlebulus pilosior (Maskell) (Monophlebidae) Chrysoperla rufi labris (Burmeister) (Chrysopidae) Orthezia urticae (L.) (Ortheziidae) Parlatoria blanchardi (Tergioni Tozzetti) (Diaspididae) Phenacoccus herreni Cox & Williams (Pseudococcidae) Lepidoptera Phenacoccus madeirensis Green (Pseudococcidae) Phenacoccus manihoti Matile - Ferrero (Pseudococcidae) Acraea encedon (L.) (Nymphalidae) – cassava mealybug [Anagasta kuehniella (Zeller)] = Ephestia kuehniella Phoenicococcus marlatti (Cockerell) (Phoenicococcidae) Zeller (Pyralidae ) Pinnaspis buxi (Bouch é ) (Diaspididae) Arctia (Arctiidae) – tiger moths Planococcus citri (Risso) (Pseudococcidae) – citrus Choristoneura pinus Freeman (Tortricidae) – jack pine mealybug budworm Planococcus minor (Maskell) (Pseudococcidae) Danaus plexippus (L.) ( Nymphalidae) – monarch Pseudochermes fraxini (Kaltenbach) (Eriococcidae) butterfl y Pseudococcus cryptus Hempel (Pseudococcidae) Ephestia kuehniella Zeller ( Pyralidae ) – Mediterranean Pseudococcus maritimus (Ehrhorn) (Pseudococcidae) fl our moth Pseudococcus viburni (Signoret) (Pseudococcidae) Galleria mellonella (L.) (Pyralidae) – wax moth Psylla alni (L.) (Psyllidae) Helicoverpa armigera (H ü bner) (Noctuidae) – cotton Psylla jucunda Tuthill (Psyllidae) bollworm Psylla mali (Schmidberger) (Psyllidae) – apple sucker Helicoverpa zea (Boddie) (Noctuidae) – corn earworm Psylla ulmi Forster (Psyllidae) Heliothis virescens (F.) (Noctuidae) – tobacco budworm Psylla uncatoides (Ferris et Clyver) (Psyllidae) Hyphantria cunea (Drury) (Arctiidae) – fall webworm Pulvinaria psidii Maskell (Coccidae) – green shield scale Ostrinia nubilalis (H ü bner) (Crambidae) – European Pulvinaria urbicola (Cockerell) (Coccidae) corn borer Pulvinaria vitis (L.) (Coccidae) – woolly vine scale Pectinophora gossypiella (Saunders) (Gelechiidae) – pink Rastrococcus invadens Williams (Pseudococcidae) bollworm Saissetia coffeae (Walker) (Coccidae) – helmet scale Phthorimaea operculella (Zeller) (Gelechiidae) – potato Saissetia oleae (Olivier) (Coccidae) – olive scale tuber moth Siphoninus phillyreae (Haliday) (Aleyrodidae) Pieris rapae (L.) (Pieridae) – small cabbage white Trialeurodes vaporariorum (Westwood) (Aleyrodidae) butterfl y – glasshouse whitefl y Plutella xylostella (L.) (Plutellidae) – diamondback Unaspis citri (Comstock) (Diaspididae) – citrus snow scale moth Unaspis euonymi (Comstock) (Diaspididae) Sitotroga cerealella (Olivier) (Gelechiidae) – Angoumois Unaspis yanonensis Kuwana (Diaspididae) grain moth Spodoptera litura (F.) (Noctuidae) – common cutworm Thysanoptera Trichoplusia ni (H ü bner) (Noctuidae) – cabbage looper or ni moth Thrips tabaci Lindeman (Thripidae) – onion thrips or Tyria (Arctiidae) – cinnabar moths tobacco thrips

Neuroptera Diptera

Chrysopa Leach (Chrysopidae) Anopheles quadrimaculatus Say (Culicidae) Chrysopa oculata Say (Chrysopidae) – golden - eyed Aphidoletes aphidimyza (Rondani) (Cecidomyiidae) lacewing Boettcheria latisterna Parker (Sarcophagidae) xxx Taxonomic glossary

Chetogena claripennis (Macquart) (Tachinidae) Hymenoptera Chrysotachina slossonae (Coquillett) (Tachinidae) Contarinia nasturtii (Kieffer) (Cecidomyiidae) – swede [Aminellus Masi] = Cowperia Girault (Encyrtidae) midge [Aminellus sumatraensis Kerrich] = Cowperia Cryptochaetum iceryae (Williston) (Cryptochaetidae) sumatraensis (Kerrich) (Encyrtidae) Culex quinquefasciatus Say (Culicidae) – southern Anagyrus Howard (Encyrtidae) house mosquito Anagyrus australiensis (Howard) (Encyrtidae) [Degeeria luctuosa (Meigen)] = Medina luctuosa Anagyrus kamali Moursi (Encyrtidae) (Meigen) (Tachinidae) Anagyrus lopezi (De Santis) (Encyrtidae) [Doryphorophaga doryphorae ] = Myiopharus doryphorae Anagyrus pseudococci (Girault) (Encyrtidae) (Riley) (Tachinidae) Anastatus Motschulsky (Eupelmidae) Drosophila melanogaster Meigen (Drosophilidae) [Anisotylus Timberlake] = Homalotylus Mayr Episyrphus balteatus (DeGeer) (Syrphidae) – (Encyrtidae) marmalade hover fl y Aphanogmus Thomson (Ceraphronidae) Euthelyconychia epilachnae (Aldrich) (Tachinidae) Aphidius colemani Viereck (Braconidae) [Exoristoides slossonae Coquillett] = Chrysotachina Aphidius eadyi Star ý (Braconidae) slossonae (Coquillett) (Tachinidae) Aphidius ervi Haliday (Braconidae) Helicobia rapax (Walker) (Sarcophagidae) Apis mellifera L. (Apidae) – European honey bee Lydinolydella metallica Townsend (Tachinidae) Aprostocetus Westwood (Eulophidae) [Lypha slossonae (Coquillett)] = Chrysotachina slossonae Aprostocetus esurus (Riley) (Eulophidae) (Coquillett) (Tachinidae) Aprostocetus neglectus (Domenichini) (Eulophidae)) Medina collaris (Fall é n) (Tachinidae) [Atritomellus Kieffer] = Dendrocerus (Megaspilidae) Medina funebris (Meigen) (Tachinidae) Austroterobia Girault (Pteromalidae) Medina luctuosa (Meigen) (Tachinidae) Aximopsis Ashmead (Eurytomidae) Medina melania (Meigen) (Tachinidae) Azteca instabilis (Smith) (Formicidae) Medina separata (Meigen) (Tachinidae) Baryscapus Foerster (Eulophidae) Megaselia (Phoridae) Baryscapus thanasimi (Ashmead) (Eulophidae) Myiopharus doryphorae (Riley) (Tachinidae) Brachymeria carinatifrons Gahan (Chalcididae) [Paradexodes epilachnae Aldrich] = Euthelyconychia Catolaccus Thomson (Pteromalidae) epilachnae (Aldrich) (Tachinidae) Centistes scymni Ferri è re (Braconidae) Phalacrotophora Enderlein (Phoridae) Centistes subsulcatus (Thomson) (Braconidae) Phalacrotophora berolinensis Schmitz (Phoridae) Centistina nipponicus (Belokobylskij) (Braconidae) Phalacrotophora beuki Disney (Phoridae) Cerchysiella Girault [ Zeteticontus Silvestri] (Encyrtidae) Phalacrotophora decimaculata Liu (Phoridae) Chartocerus subaeneus (Foerster) (Signiphoridae) Phalacrotophora delageae Disney (Phoridae) Cheiloneurus carinatus Compere (Encyrtidae) Phalacrotophora fasciata (Fall é n) (Phoridae) Cheiloneurus cyanonotus Waterston (Encyrtidae) Phalacrotophora indiana Colyer (Phoridae) Cheiloneurus liorhipnusi (Risbec) (Encyrtidae) Phalacrotophora nedae (Malloch) (Phoridae) Cheiloneurus orbitalis Compere (Encyrtidae) Phalacrotophora philaxyridis Disney (Phoridae) Chrysocharis johnsoni Subba Rao (Eulophidae) Phalacrotophora quadrimaculata Schmitz (Phoridae) Chrysonotomyia appannai (Chandy Kurian) Policheta unicolor (Fall é n) (Tachinidae) (Eulophidae) Pseudebenia epilachnae Shima & Han (Tachinidae) Coccidoctonus trinidadensis Crawford (Encyrtidae) Ravinia errabunda (Wulp) (Sarcophagidae) Conura Spinola (Chalcididae) [Sarcophaga helicis Townsend] = Helicobia rapax Conura paranensis (Schrottky) (Chalcididae) (Walker) (Sarcophagidae) Conura petioliventris (Cameron) (Chalcididae) [Sarcophaga latisterna (Parker)] = Boettcheria latisterna Conura porteri (Br è thes) (Chalcididae) Parker (Sarcophagidae) Cowperia Girault (Encyrtidae) [Sarcophaga reinhardii Hall] = Ravinia errabunda (Wulp) Cowperia areolata (Walker) (Encyrtidae) (Sarcophagidae) Cowperia indica (Kerrich) (Encyrtidae) Strongygaster triangulifera (Loew) (Tachinidae) Cowperia punctata Girault (Encyrtidae) Taxonomic glossary xxxi

Cowperia subnigra Li (Encyrtidae) [Homalotylus californicus (Say)] = Homalotylus Cowperia sumatraensis (Kerrich) (Encyrtidae) terminalis (Say) (Encyrtidae) Crematogaster Lund (Formicidae) Homalotylus cockerelli Timberlake (Encyrtidae) Crematogaster lineolata (Say) (Formicidae) Homalotylus ephippium (Ruschka) (Encyrtidae) Dendrocerus (Megaspilidae) Homalotylus eytelweinii (Ratzeburg) (Encyrtidae) Dendrocerus ergensis (Guesquiere) (Megaspilidae) Homalotylus ferrierei Hayat, Alam & Agarwal Dibrachys cavus (Walker) (Pteromalidae) (Encyrtidae) Dinocampus Foerster (Braconidae) Homalotylus fl aminius Dalman (Encyrtidae) Dinocampus coccinellae (Schrank) (Braconidae) Homalotylus hemipterinus (De Stefani) (Encyrtidae) [Dinocampus nipponicus Belokobylskij] = Centistina Homalotylus formosus Anis & Hayat (Encyrtidae) nipponicus (Belokobylskij) (Braconidae) Homalotylus himalayensis Liao (Encyrtidae) [Dinocampus terminatus (Nees)] = Dinocampus Homalotylus hybridus Hoffer (Encyrtidae) coccinellae (Schrank) (Braconidae) Homalotylus hyperaspicola Tachikawa (Encyrtidae) Dolichoderus bidens L. (Formicidae) Homalotylus hyperaspidis Timberlake (Encyrtidae) [Echthroplectis Foerster] = Homalotylus Mayr Homalotylus hypnos Noyes (Encyrtidae) Homalotylus indicus (Agarwal) (Encyrtidae) Elasmus Westwood (Eulophidae) Homalotylus latipes Girault(Encyrtidae) Encarsia sophia (Girault & Dodd) (Aphelinidae) Homalotylus longicaudus Xu & He (Encyrtidae) [Epidinocarsus lopezi (De Santis)] = Anagyrus lopezi (De Homalotylus longipedicellus (Shafee & Fatma) Santis) (Encyrtidae) (Encyrtidae) Eretmocerus mundus Mercet (Aphelinidae) Homalotylus mexicanus Timberlake (Encyrtidae) Eupelmus Dalman (Eupelmidae) Homalotylus mirabilis (Br è thes) (Encyrtidae) Eupelmus urozonus Dalman (Eupelmidae) Homalotylus mundus Gahan (Encyrtidae) Eupelmus vermai (Bhatnagar) (Eupelmidae) Homalotylus nigricornis Mercet (Encyrtidae) [Eupteromalus Kurdjumov] = Trichomalopsis Crawford Homalotylus oculatus (Girault) (Encyrtidae) (Pteromalidae) Homalotylus pallentipes (Timberlake) (Encyrtidae) Formica L. (Formicidae) Homalotylus platynaspidis Hoffer (Encyrtidae) Formica obscuripes Forel (Formicidae) Homalotylus punctifrons Timberlake (Encyrtidae) Formica polyctena Foerster (Formicidae) – European Homalotylus quaylei Timberlake (Encyrtidae) red wood ant Homalotylus rubricatus Sharkov (Encyrtidae) Formica rufa L. (Formicidae) – wood ant Homalotylus scutellaris Tan & Zhao (Encyrtidae) Formica rufi barbis F. (Formicidae) – red - barbed ant Homalotylus scymnivorus Tachikawa (Encyrtidae) Gelis Thunberg (Ichneumonidae) Homalotylus shuvakhinae Trjapitzin & Triapitsyn Gelis agilis (F.) (Ichneumonidae) (Encyrtidae) [Gelis instabilis (F ö rster)] = Gelis agilis (F.) Homalotylus similis Ashmead (Encyrtidae) (Ichneumonidae) Homalotylus sinensis Xu & He (Encyrtidae) Gelis melanocephalus (Schrank) (Ichneumonidae) Homalotylus singularis Hoffer (Encyrtidae) [Hemaenasioidea Girault] = Homalotylus Mayr Homalotylus terminalis (Say) (Encyrtidae) (Encyrtidae) Homalotylus trisubalbus Xu & He (Encyrtidae) Homalotyloidea dahlbomii (Westwood) (Encyrtidae) Homalotylus turkmenicus Myartseva (Encyrtidae) Homalotylus Mayr (Encyrtidae) Homalotylus vicinus Silvestri (Encyrtidae) Homalotylus affi nis Timberlake (Encyrtidae) Homalotylus yunnanensis Tan & Zhao (Encyrtidae) Homalotylus africanus Timberlake (Encyrtidae) Homalotylus zhaoi Xu & He (Encyrtidae) Homalotylus agarwali Anis & Hayat (Encyrtidae) Inkaka quadridentata Girault (Pteromalidae) Homalotylus albiclavatus (Agarwal) (Encyrtidae) Iridomyrmex Mayr (Formicidae) Homalotylus albifrons (Ishii) (Encyrtidae) Isodromus niger Ashmead (Encyrtidae) Homalotylus albitarsus Gahan (Encyrtidae) Lasius claviger (Roger) (Formicidae) Homalotylus aligarhensis (Shafee & Rizvi) (Encyrtidae) Lasius japonicus Santschi (Formicidae) – Japanese ant Homalotylus balchanensis Myartseva (Encyrtidae) Lasius niger (L.) (Formicidae) – black ant or garden ant Homalotylus brevicauda Timberlake (Encyrtidae) Lasius umbratus (Nylander) (Formicidae) xxxii Taxonomic glossary

[Lepidaphycus Blanchard] = Homalotylus Mayr Ooencyrtus epilachnae Annecke (Encyrtidae) (Encyrtidae) [Ooencyrtus epulus Annecke] = Ooencyrtus Linepithema humile (Mayr) (Formicidae) – Argentine camerounensis (Risbec) (Encyrtidae) ant Ooencyrtus polyphagus (Risbec) (Encyrtidae) [Lygocerus Foerster] = Dendrocerus (Megaspilidae) Ooencyrtus puparum Prinsloo (Encyrtidae) Lysiphlebus fabarum (Marshall) (Braconidae) Ooencyrtus sinis Prinsloo (Encyrtidae) Lysiphlebus testaceipes (Cresson) (Braconidae) Oomyzus Rondani (Eulophidae) [Mendozaniella Brethes] = Homalotylus Mayr Oomyzus mashhoodi (Khan & Shafee) (Eulophidae) (Encyrtidae) Oomyzus scaposus (Thomson) (Eulophidae) Merismoclea rojasi De Santis (Pteromalidae) Oomyzus sempronius (Erdoes) (Eulophidae) Mesopolobus Westwood (Pteromalidae) Ophelosia bifasciata Girault (Pteromalidae) Mesopolobus secundus (Crawford) (Pteromalidae) Oricoruna Boucek (Pteromalidae) [Mestocharis lividus Girault)] = Pediobius foveolatus Ophelosia crawfordi Riley (Pteromalidae) (Crawford) (Eulophidae) Oricoruna orientalis (Crawford) (Pteromalidae) Metastenus Walker (Pteromalidae) Pachyneuron Walker (Pteromalidae) Metastenus caliginosus Szel é nyi (Pteromalidae) Pachyneuron albutius Walker (Pteromalidae) Metastenus concinnus Walker (Pteromalidae) Pachyneuron altiscuta Howard (Pteromalidae) Metastenus indicus Sureshan & Narendran Pachyneuron chilocori Domenichini (Pteromalidae) (Pteromalidae) [Pachyneuron concolor F ö rster] = Pachyneuron Metastenus sulcatus (Dodd) (Pteromalidae) muscarum (L.) (Pteromalidae) Metastenus townsendi (Ashmead) (Pteromalidae) Pachyneuron muscarum (L.) (Pteromalidae) Microctonus Wesmael (Braconidae) [Pachyneuron siculum Delucchi] = Pachyneuron Monomorium minimum (Buckley) ( Formicidae ) – little muscarum (L.) (Pteromalidae) black ant Pachyneuron solitarium (Hartig) (Pteromalidae) Myrmica ruginodis Nylander (Formicidae) [Pachyneuron syrphi (Ashmead)] = Pachyneuron Myrmica rugulosa Nylander (Formicidae) albutius Walker (Pteromalidae) [Neoaenasioidea Agarwal] = Homalotylus Mayr Parachrysocharis Girault (Eulophidae) (Encyrtidae) Paratrechina Motschulsky (Formicidae) [Neotainania Husain & Agarwal] = Uga Girault Pediobius Walker (Eulophidae) (Chalcididae) Pediobius amaurocoelus (Waterston) (Eulophidae) [Nobrimus Thomson] = Homalotylus Mayr (Encyrtidae) [Pediobius epilachnae (Rohwer)] = Pediobius foveolatus Nothoserphus Brues (Proctotrupidae) (Crawford) (Eulophidae) Nothoserphus admirabilis Lin (Proctotrupidae) Pediobius foveolatus (Crawford) (Eulophidae) Nothoserphus aequalis Townes (Proctotrupidae) [Pediobius mediopunctata (Waterston)] = Pediobius Nothoserphus afi ssae (Watanabe) (Proctotrupidae) foveolatus (Crawford) (Eulophidae) Nothoserphus boops (Thomson) (Proctotrupidae) Pediobius nishidai Hansson (Eulophidae) Nothoserphus debilis Townes (Proctotrupidae) [Pediobius simiolus (Takahashi)] = Pediobius foveolatus Nothoserphus epilachnae (Pschorn - Walcher) (Crawford) (Eulophidae) (Proctotrupidae ) [ Perilitus americanus Riley] = Dinocampus coccinellae Nothoserphus fuscipes Lin (Proctotrupidae) (Schrank) (Braconidae) Nothoserphus mirabilis Brues (Proctotrupidae) [Perilitus coccinellae (Schrank)] = Dinocampus Nothoserphus partitus Lin (Proctotrupidae) coccinellae (Schrank) (Braconidae) Nothoserphus scymni (Ashmead) (Proctotrupidae) Perilitus rutilus (Nees) (Braconidae) Nothoserphus townesi Lin (Proctotrupidae) Perilitus stuardoi Porter (Braconidae) Omphale Haliday (Eulophidae) [Perilitus terminatus (Nees)] = Dinocampus coccinellae [Omphale epilachni Singh & Khan] = Chrysonotomyia (Schrank) (Braconidae) appannai (Chandy Kurian) (Eulophidae) Pheidole megacephala (F.) (Formicidae) – big - headed Ooencyrtus azul Prinsloo (Encyrtidae) ant Ooencyrtus bedfordi Prinsloo (Encyrtidae) Phygadeuon subfuscus Cresson ( Ichneumonidae ) Ooencyrtus camerounensis (Risbec) (Encyrtidae) [Pleurotropis Foerster] = Pediobius Walker (Eulophidae) Ooencyrtus distatus Prinsloo (Encyrtidae) Pnigalio agraules (Walker) (Eulophidae) Taxonomic glossary xxxiii

Praon volucre (Haliday (Braconidae) [Watanabeia Masner] = Nothoserphus Brues Pristomyrmex pungens Mayr (Formicidae) (Proctotrupidae) Prochiloneurus aegyptiacus (Mercet) (Encyrtidae) Prochiloneurus nigrifl agellum (Girault) (Encyrtidae) Pseudocatolaccus Masi (Pteromalidae) Coleoptera (other than Coccinellidae) Quadrastichus ovulorum (Ferri è re) (Eulophidae) [Scymnophagus Ashmead] = Metastenus Walker Agelastica coerulea (Baly) (Chrysomelidae) – alder leaf (Pteromalidae) beetle [Scymnophagus mesnili Ferri è re] = Metastenus Agonum dorsale (Pontoppidan) (Carabidae) concinnus Walker (Pteromalidae) Chrysomela populi L. (Chrysomelidae) – poplar leaf Sigmoepilachna indica Khan, Agnihotri & Sushil beetle (Eulophidae) Chrysophtharta bimaculata (Olivier) (Chrysomelidae) Solenopsis invicta Buren (Formicidae) – red imported – eucalyptus leaf beetle fi re ant Diabrotica virgifera LeConte (Chrysomelidae) – western [Syntomosphyrus taprobanes Waterston] = Oomyzus corn rootworm scaposus (Thomson) (Eulophidae) Dicranolaius bellulus (Gu é rin - M é neville) (Melyridae) Syrphoctonus tarsatorius (Panzer) ( Ichneumonidae ) Dytiscus sp. (Dytiscidae) Tamarixia radiata (Waterston) (Eulophidae) Galeruca interrupta arminiaca Weise (Chrysomelidae) Tapinoma nigerrimum (Nylander) (Formicidae) Galerucella lineola (F.) (Chrysomelidae) Tetramorium caespitum (L.) (Formicidae) Galerucella pusilla Duftschmid (Chrysomelidae) Tetrastichus Haliday (Eulophidae) – golden loosestrife beetle [Tetrastichus coccinellae Kurdjumov] = Oomyzus Galerucella sagittariae Gyllenhal (Chrysomelidae) scaposus (Thomson) (Eulophidae) – cloudberry beetle Tetrastichus cydoniae Risbec (Eulophidae) Harpalus pennsylvanicus (De Geer) (Carabidae) Tetrastichus decrescens Graham (Eulophidae) Hypera postica (Gyllenhal) (Curculionidae) – alfalfa Tetrastichus epilachnae (Giard) (Eulophidae) weevil [Tetrastichus melanis Burks] = Oomyzus scaposus Laricobius nigrinus Fender (Derodontidae) (Thomson) (Eulophidae) Leptinotarsa decemlineata (Say) (Chrysomelidae) – Tetrastichus orissaensis Husain & Khan (Eulophidae) Colorado potato beetle [Tetrastichus sexmaculatus Chandy Kurian] = Oomyzus [Melasoma populi (L.)] = Chrysomela populi L. scaposus (Thomson) (Eulophidae) (Chrysomelidae) [Thomsonina Hellen] = Nothoserphus Brues Melolontha sp. (Scarabaeidae) (Proctotrupidae) Plagiodera versicolora (Laicharting) (Chrysomelidae) Trichogramma Westwood (Trichogrammatidae) Platynus dorsalis (Pontopiddan) (Carabidae) Trichogramma evanescens Westwood Pterostichus melanarius (Illiger) (Carabidae) (Trichogrammatidae) [Pyrrhalta luteola (Muller)] = Xanthogaleruca luteola Trichomalopsis Crawford (Pteromalidae) (Muller) (Chrysomelidae) Trichomalopsis acuminata (Graham) (Pteromalidae) Sitona discoideus Gyllenhal (Curculionidae) Trichomalopsis dubia (Ashmead) (Pteromalidae) Stenotarsus rotundus Arrow (Endomychidae) [Tripolycystus Dodd] = Metastenus Walker Tachyporus sp. (Staphylinidae) (Pteromalidae) Xanthogaleruca luteola (Muller) (Chrysomelidae) – elm Uga Girault [ Neotainania Husain & Agarwal] leaf beetle (Chalcididae) Uga colliscutellum (Girault) (Chalcididae) Uga coriacea Kerrich (Chalcididae) PARASITIC MITES Uga digitata Qian & He (Chalcididae) Uga hemicarinata Qian & Li (Chalcididae) Coccipolipus Husband (Podapolipidae) Uga javanica Kerrich (Chalcididae) Coccipolipus africanae Husband (Podapolipidae) Uga menoni Kerrich (Chalcididae) Coccipolipus arturi Haitlinger (Podapolipidae) Uga sinensis Kerrich (Chalcididae) Coccipolipus benoiti Husband (Podapolipidae) Wasmannia auropunctata (Roger) (Formicidae) Coccipolipus bifasciatae Husband (Podapolipidae) xxxiv Taxonomic glossary

Coccipolipus cacti Husband(Podapolipidae) Hexamermis Steiner (Mermithidae) Coccipolipus camerouni Husband (Podapolipidae) Howardula Cobb (Allantonematidae) Coccipolipus chilocori Husband (Podapolipidae) Mermis Dujardin (Mermithidae) Coccipolipus cooremani Husband (Podapolipidae) Parasitilenchus coccinellinae Iperti & van Waerebeke Coccipolipus epilachnae Smiley (Podapolipidae) (Allantonematidae) Coccipolipus hippodamiae (McDaniel & Morrill) (Podapolipidae) MOLLUSCA Coccipolipus macfarlanei Husband (Podapolipidae) Coccipolipus micraspisi Husband (Podapolipidae) Arion ater (L.) (Arionidae) – European black slug Coccipolipus oconnori Husband (Podapolipidae) Arion hortensis (F é russac) (Arionidae) – garden slug Coccipolipus solanophilae (Cooreman) (Podapolipidae) Deroceras reticulatum (M ü ller) (Limacidae) – grey fi eld Hemisarcoptes Lignieres (Hemisarcoptidae) slug or grey garden slug Hemisarcoptes cooremani Thomas (Hemisarcoptidae) Tandonia budapestensis (Hazay) (Milacidae) – Budapest Leptus ignotus (Oudemans) (Erythraeidae) slug

ARACHNIDA (other than parasitic mites) AVES Agistemus longisetus Gonzalez (Stigmaeidae) Acrocephalus schoenobaenus (L.) (Sylviidae) – sedge Amblyseius andersoni (Chant) (Phytoseidae) warbler Amblyseius fallacis (Garman) (Phytoseidae) Alauda arvensis L. (Alaudidae) – sky lark Amphitetranychus viennensis (Zacher) (Tetranychidae) Anthus campestris (L.) (Motacillidae) – tawny pipit – hawthorn spider mite or fruit tree spider mite Anthus pratensis (L.) (Motacillidae) – meadow pipit Araneus diadematus (Clerck) (Araneidae) Anthus trivialis (L.) (Motacillidae) – tree pipit Bryobia praetiosa Koch (Tetranychidae) – clover mite Apus apus (L.) (Apodidaae) – common swift Bryobia rubrioculus (Scheuten) (Tetranychidae) Calandrella cinerea (Gmelin) (Alaudidae) – red - capped – brown mite or brown apple mite lark Cheiracanthium (Miturgidae) Coturnix coturnix L. (Phasianidae) – quail Clubiona reclusa O.P. - Cambridge (Clubionidae) Coturnix japonicus Temminck & Schlegel (Phasianidae) [Metatetranychus ulmi (Koch)] = Panonychus ulmi – Japanese quail (Koch) (Tetranychidae) Cuculus canorus L. (Cuculidae) – common cuckoo Misumenops tricuspidatus (F.) (Thomisidae) Cyanistes caeruleus (L.) (Paridae) – blue tit Panonychus mori Yokovama (Tetranychidae) Delichon urbica (L.) (Hirundinidae) – house martin Panonychus ulmi (Koch) (Tetranychidae) – red spider Dendrocopos medius (L.) (Picidae) – middle spotted mite or European red mite woodpecker Tetranychus evansi Baker & Pritchard (Tetranychidae) Ficedula parva (Bechstein) (Muscicapidae) – red - headed Tetranychus lintearius Dufour (Tetranychidae) – gorse fl ycatcher spider mite Hippolais icterina (Vieillot) (Sylviidae) – icterine Tetranychus mcdanieli McGregor (Tetranychidae) warbler [Tetranychus telarius (L.)] = Tetranychus urticae Koch Hirundo rustica L. (Hirundinidae) – barn swallow (Tetranychidae) Luscinia luscinia (L.) (Muscicapidae) – thrush Tetranychus urticae Koch (Tetranychidae) – two - spotted nightingale spider mite Luscinia svecica (L.) (Muscicapidae) – bluethroat Thanatus Koch (Philodromidae) Melanocorypha calandra (L.) (Alaudidae) – Calandra lark Motacilla fl ava L. (Motacillidae) – blue - headed wagtail NEMATODES Muscicapa striata (Pallas) (Muscicapidae) – spotted fl ycatcher [Coccinellimermis Rubtzov] = Hexamermis Steiner Oenanthe oenanthe (L.) (Muscicapidae) – northern (Mermithidae) wheatear Taxonomic glossary xxxv

Parus major L. (Paridae) Microsphaera alphitoides Griffon & Maublanc Passer domesticus (L.) (Passeridae) – house sparrow (Erisyphaceae) Passer montanus (L.) (Passeridae) – tree sparrow Microsphaera pulchra Cooke & Peck (Erisyphaceae) Phoenicurus ochruros (Gmelin) (Muscicapidae) – black Neotyphodium lolii Latch et al . (Clavicipitaceae) redstart Nosema coccinellae Lipa (Nosematidae) Phoenicurus phoenicurus (L.) (Muscicapidae) – common Nosema epilachnae Brooks, Hazard & Becnel redstart (Nosematidae) Phylloscopus collybita (Vieillot) (Phylloscopidae) Nosema henosepilachnae Toguebaye & Marchand – common chiffchaff (Nosematidae) Phylloscopus trochilus (L.) (Phylloscopidae) – willow Nosema hippodamiae Lipa & Steinhaus (Nosematidae) warbler Nosema tracheophila Cali & Briggs (Nosematidae) Saxicola rubicola (L.) (Muscicapidae) – common Nosema varivestis Brooks, Hazard & Becnel stonechat (Nosematidae) Sitta europaea L. (Sittidae) – wood nuthatch Tubulinosema hippodamiae Bjornson, Le, Saito & Wang Sturnus vulgaris L. (Sturnidae) – common starling (Nosematidae) Sylvia atricapilla (L.) (Sylviidae) – blackcap Oidium monilioides (Nees) Link (Erisyphaceae) Sylvia borin (Boddaert) (Sylviidae) – garden warbler [Paecilomyces farinosus (Holmskiold ex S. F. Gray) Sylvia communis Latham (Sylviidae) – common Brown & Smith] = Isaria farinosa (Holmskiold ex S. whitethroat F. Gray) Fries (Cordycipitaceae) Sylvia curruca (L.) (Sylviidae) – lesser whitethroat [Paecilomyces fumosoroseus (Wize) Brown & Smith] = Sylvia nisoria (Bechstein) (Sylviidae) – barred warbler Isaria fumosorosea Wize (Cordycipitaceae) Turdus philomelos (Turton) (Turdidae) – song thrush Pandora neoaphidis (Remaudi è re & Hennebert) Turdus merula L. (Turdidae) – Eurasian blackbird Humber (Entomophthoraceae) Phyllactinia moricola (Hennebert) Homma FUNGI (Erisyphaceae) Podosphaera leucotricha (Ellis & Everhart) E.S. Salmon Alternaria Nees (Pleosporaceae) (Erisyphaceae) – apple powdery mildew Beauveria bassiana (Balsamo) Vuillemin Podosphaera tridactyla (Wallroth) de Bary (Cordycipitaceae) (Erisyphaceae) – plum powdery mildew Cladosporium Link (Davidiellaceae) Puccinia Pers. (Pucciniaceae) Erisyphe polygoni DC. (Erisyphaceae) – powdery Saccharomyces fragilis Kudrjawzew mildew (Saccharomycetaceae) Erysiphe cichoracearum DC (Erisyphaceae) Sphaerotheca castagnei L é verrier (Erisyphaceae) – hop Hesperomyces Thaxter (Laboulbeniaceae) powdery mildew Hesperomyces chilomenis (Thaxter) Thaxter Sphaerotheca cucurbitae (Jaczewski) Z.Y. Zhao (Laboulbeniaceae) (Erisyphaceae) Hesperomyces coccinelloides (Thaxter) Thaxter Sphaerotheca pannosa (Wallroth) de Bary (Laboulbeniaceae) (Erisyphaceae) – rose powdery mildew Hesperomyces hyperaspidis Thaxter (Laboulbeniaceae) [Verticillium lecanii (Zimmerman) Hesperomyces virescens Thaxter (Laboulbeniaceae) Viegas] = Lecanicillium lecanii (Zimmerman) Zare & Isaria farinosa (Holmskiold ex S. F. Gray) Fries Gams (Cordycipitaceae) (Cordycipitaceae) Isaria fumosorosea Wize (Cordycipitaceae) Lecanicillium lecanii (Zimmerman) Zare & Gams PROTISTS (Cordycipitaceae) [Lecanicillium longisporum (Zimmerman) Zare & Gams Anisolobus indicus Haldar, Ray & Bose (Gregarinidae) – as ‘ Vertalec ’ ] = Lecanicillium lecanii (Zimmerman) Brustiospora indicola Kundu & Haldar Zare & Gams (Cordycipitaceae) (Brustiophoridae) Metarhizium anisopliae (Metschnikoff) Sorokin Gregarina Dufour (Gregarinidae) (Clavicipitaceae) Gregarina barbarara Watson (Gregarinidae) xxxvi Taxonomic glossary

Gregarina californica Lipa (Gregarinidae) Betula populifolia Marshall (Betulaceae) Gregarina chilocori Obata (Gregarinidae) Brassica campestris L. (Brassicaceae) Gregarina coccinellae Lipa (Gregarinidae) Brassica napus L. (Brassicaceae) – rape Gregarina dasguptai Mandal, Rai, Pranhan, Gurung, Brassica napus L. subsp. oleifera DC Metzger Sharma, Rai & Mandal (Gregarinidae) (Brassicaceae) – oil seed rape Gregarina fragilis Watson (Gregarinidae) Brassica nigra (L.) Koch (Brassicaceae) – black (or Gregarina hyashii Sengupta & Haldar (Gregarinidae) brown) mustard Gregarina katherina Watson (Gregarinidae) Brassica oleracea L. Italica group (Brassicaceae) Gregarina ruszkowskii Lipa (Gregarinidae) – broccoli Gregarina straeleni Theodorides & Jolivet Brassica oleracea L. (Brassicaceae) (Gregarinidae) Calotropis procera (Aiton) Aiton (Apocynaceae) Caltha palustris L. (Ranunculaceae) Cannabis sativa L. (Cannabaceae) – Indian hemp Carduus crispus L. (Asteraceae) BACTERIA Carpinus caroliniana Walter (Betulaceae) – American hickory Alcaligenes paradoxus Davis (Alcaligenaceae) Caulophyllum robustum Maximowicz (Berberidaceae) Bacillus thuringiensis Berliner (Bacillaceae) Centaurea jacea L. (Asteraceae) Flavobacterium Bergey (Flavobacteriaceae) Centrosema pubescens Bentham (Fabaceae) Rickettsia da Rocha - Lima (Rickettsiaceae) Chamerion angustifolium (L.) Holub (Onagraceae) Spiroplasma Saglio, Lhospital, Lafl eche, Dupont, Bove, – fi reweed Tully & Freundt (Spiroplasmataceae) Chenopodium L. (Chenopodiaceae) Wolbachia Hertig (Rickettsiaceae ) Cirsium arvense (L.) Scopoli (Asteraceae) – creeping thistle Cirsium kagamontanum Nakai (Asteraceae) PLANTS Cirsium kamtschaticum De Candolle (Asteraceae) Citrus sinensis (L.) Osbeck (Rutaceae) – sweet orange Abies balsamea (L.) Miller (Pinaceae) – balsam fi r Clematis L. (Ranunculaceae) Abutilon theophrasti Medikus (Malvaceae) – Chinese Coriandrum sativum L. (Apiaceae) – coriander jute or Indian Mallow Cosmos (Asteraceae) Miller (Fabaceae) Cotoneaster integerrima Medikus (Rosaceae) Acalypha ostryaefolia Riddell (Euphorbiaceae) Cotoneaster tomentosus Lindley (Rosaceae) Aconitum L. (Ranunculaceae) Crotalaria striata De Candolle (Fabaceae) Aframomum melegueta (Roscoe) K. Schum Cucurbita maxima Duchesne () – squash (Zingiberaceae) – grains of paradise or melegueta or pumpkin pepper Cymbopogon citratus (De Candolle) Stapf (Poaceae) Agropyron desertorum (Link) Schultes (Poaceae) – lemon grass – crested wheatgrass Dendrocalamus giganteus Munro (Poaceae) – giant Alnus japonica (Thunberg) Steudel (Betulaceae) bamboo Amaranthus hybridus L. ( Amaranthaceae) Elytrigia repens (L.) Nevski (Poaceae) – couch grass Ambrosia artemisiifolia L. (Asteraceae) – hogbrake Erythrina corallodendron L. (Fabaceae) Anethum graveolens L. (Apiaceae) – dill Euonymus japonicus Thunberg (Celastraceae) Artemisia tridentata Nuttall (Asteraceae) – sagebrush Euphorbia L . (Euphorbiaceae) Artemisia vulgaris L. (Asteraceae) – wormwood Eurotia ceratoides Meyer (Chenopodiaceae) Atriplex sagittata Borkhausen (Chenopodiaceae) Genista L. (Fabaceae) Benthamidia fl orida (L.) Spach (Cornaceae) Glochidion ferdinandi (M ü ller Argiovensis) Bailey Berberis vulgaris L. (Berberidaceae ) – common (Euphorbiaceae) barberry Helianthus annuus L. (Asteraceae) – sunfl ower Beta vulgaris L. (Chenopodiaceae) – sugar beet and Hibiscus L. (Malvaceae) garden beet Juglans cinerea L. (Juglandaceae) – butternut Taxonomic glossary xxxvii

Juniperus virginiana L. (Cupressaceae) – red cedar or Pyracantha coccinea M.H. Roemer (Rosaceae) Virginian pencil cedar – fi rethorn Lapsana communis L. (Asteraceae) – nipplewort Quercus rubra L. (Fagaceae) – red oak Leontopodium alpinum Cassini (Asteraceae) – edelweiss Raphanus sativus L. (Brassicaceae) – radish Leucaena Bentham (Fabaceae) Robinia pseudoacacia L. (Fabaceae) Ligustrum L. (Oleaceae) Rosa multifl ora Thunberg (Rosaceae) Lolium multifl orum Lamarck (Poaceae) – Italian rye Rubus occidentalis L. (Rosaceae) grass Salix L. (Salicaceae) Lolium perenne L. (Poaceae) – rye grass Sambucus nigra L. (Caprifoliaceae) – elder Lonicera periclymenum L. (Caprifoliaceae) Sambucus racemosa L. subsp. sieboldiana (Miquel) H. – honeysuckle Hara (Caprifoliaceae) Lupinus luteus L. (Fabaceae) – yellow lupin Schizopepon bryoniaefolius Maximowicz Lupinus mutabilis Sweet (Fabaceae) (Cucurbitaceae) Malus pumila Miller (Rosaceae) – apple Senecio jacobaea L. (Asteraceae) – ragwort Manihot esculenta Crantz (Euphorbiaceae) Senecio vulgaris L. (Asteraceae) – groundsel Manikara zapota (L.) P. Royen (Sapotaceae) – sapodilla Sinapis alba L. (Brassicaceae) – white (or yellow) Medicago sativa L. (Fabaceae) – alfalfa (or often lucerne mustard in UK) Solanum japonense Nakai (Solanaceae ) Mercurialis annua L. (Euphorbiaceae) – annual mercury Solanum nigrum L. (Solanaceae ) – black nightshade Momordica charantia L. (Cucurbitaceae) – bitter melon Solanum tuberosum L.(Solanaceae ) – potato Morus australis Poir. (Moraceae) Solidago canadensis L. (Asteraceae) Nerium oleander L. (Apocynaceae) – oleander Spartium junceum L. (Fabaceae) – Spanish broom Nitraria L. (Nitrariaceae) Spiraea (Rosaceae) Onoclea sensibilis L. (Onocleaceae) Styrax L. (Styracaceae) Oryzopsis hymenoides (Roemer & Schultes) Piper Symphoricarpos rivularis Sucksdorf ( Caprifoliaceae) (Poaceae) – Indian ricegrass Talinum triangulare (Jacquin) Willdenow Paederia foetida L. (Rutaceae) (Portulacaceae) Pastinaca sativa L. (Apiaceae) – parsnip Tamarix L. (Tamaricaceae) Phaseolus vulgaris L. (Fabaceae) – French bean or Tanacetum vulgare L. (Asteraceae) – tansy haricot bean Taraxacum offi cinale Weber (Asteraceae) – dandelion Physalis alkekengi L. (Solanaceae ) – Chinese lantern Trichosanthes kirilowii Maximowicz (Cucurbitaceae) Picea schrenkiana Fischer & Meyer (Pinaceae) Tripleurospermum maritimum (L.) Koch (Asteraceae) Pinus armandii Franchet (Pinaceae) – Chinese white Triticum L. (Poaceae) – wheat pine Tsuga Carri è re (Pinaceae) Pinus sylvestris L. (Pinaceae) – Scots pine Typha latifolia L. (Typhaceae) – bullrush Pisonia L. (Nyctaginaceae) Ulmus (Ulmaceae) Pisum sativum L. (Fabaceae) – garden pea Urtica dioica L. (Urticaceae) – stinging nettle Pittosporum tobira Aiton ( Pittosporaceae) Verbascum thapsus L. (Scrophulariaceae) – Aaron ’ s rod Populus L. (Salicaceae) Vicia faba L. (Fabaceae) – broad bean or faba bean Prunus avium (L.) L. (Rosaceae) – gean or wild cherry Vicia sativa L. (Fabaceae) – vetch or tare Prunus cerasus L. (Rosaceae) – morello cherry or Vigna unguiculata (L.) Walpers subsp. cylindrica (L.) amarelle cherry Verdcourt (Fabaceae) Prunus persica (L.) (Rosaceae) – peach or nectarine Withania somnifera (L.) Dunal (Solanaceae ) Batsch Zea mays L. (Poaceae) – maize (corn in USA) Pterostyrax hispidus Siebold & Zuccarini (Styracaceae) Zingiber offi cinale Roscoe (Zingiberaceae) – ginger