ECOLOGY AND BEHAVIOUR OF THE LADYBIRD BEETLES (COCCINELLIDAE)

Edited by I. Hodek, H.E van Emden and A. Honek

©WILEY-BLACKWELL A John Wiley & Sons, Ltd., Publication CONTENTS

Detailed contents, ix 8. NATURAL ENEMIES OF LADYBIRD BEETLES, 375 Contributors, xvii Piotr Ccryngier. Helen E. Roy and Remy L. Poland Preface, xviii 9. COCCINELLIDS AND [ntroduction, xix SEMIOCHEMICALS, 444 ]an Pettcrsson Taxonomic glossary, xx 10. QUANTIFYING THE IMPACT OF 1. PHYLOGENY AND CLASSIFICATION, 1 COCCINELLIDS ON THEIR PREY, 465 Oldrich Nedved and Ivo Kovdf /. P. Mid'laud and James D. Harwood

2. GENETIC STUDIES, 13 11. COCCINELLIDS IN BIOLOGICAL John J. Sloggett and Alois Honek CONTROL, 488 /. P. Midland 3. LIFE HISTORY AND DEVELOPMENT, 54 12. RECENT PROGRESS AND POSSIBLE Oldrkli Nedved and Alois Honek FUTURE TRENDS IN THE STUDY OF COCCINELLIDAE, 520 4. DISTRIBUTION AND HABITATS, 110 Helmut /; van Emden and Ivo Hodek Alois Honek Appendix: List of Genera in Tribes and Subfamilies, 526 5. FOOD RELATIONSHIPS, 141 Ivo Hodek and Edward W. Evans Oldrich Nedved and Ivo Kovdf

Subject index. 532 6. DIAPAUSE/DORMANCY, 275 Ivo Hodek Colour plate pages fall between pp. 250 and pp. 251

7. INTRAGUILD INTERACTIONS, 343 Eric 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 Honek 1. PHYLOGENY AND CLASSIFICATION, 1 2.1 Introduction, 14 Oldrich Nedved and Ivo Kovdf 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 Classification of Subfamilies, 3 2.3.4 Cytogenetic changes, intraspecific 1.3.1 Morphologically based cytogenetic variation and classifications. 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 Significance and evolution. 25 1.3.5 Rejection of the monophyly of The Inheritance of Other Traits. 2 7 subfamilies, 5 2.5.1 Morphological characters: wing 1.4 Characteristics of the Subfamilies and polymorphism, 2 7 Tribes, 5 2.5.2 Life history characters: heritability. 1.4.1 Proposed classification. 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 Coccinellinae, 6 2.6.1 Sequence evolution. 29 1.4.5 Epilachninae. 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 ITS 1 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 identification. 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 liffect of temperature, Acknowledgements and Dedication, 44 photoperiod and References, 44 humidity, 90 3.5.9.3 FITectof food. 91 3.5.9.4 F.ffect of sexual 3. LIFE HISTORY AND DEVELOPMENT, 54 activity, 91 Oldrich Nedved and Alois Honek 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 Honek 3.4.3 Timing of pupation. 75 4.1 Introduction, 111 3.4.4 Places of pupation, 75 4.2 Sampling. Ill 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 flight, 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. 8 3 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, 20.1 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 Specificity, 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, 21.1 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 first 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-insect food (pollen, nectar, 5.4.2 Prey capture. 2 58 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 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. 3 79 (Prostigmata: 8.2.5 Social aphids with a soldier caste, 382 Podapolipidae), 411 3.3 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 Hostspecificityof 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 (Flymenoptera: 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 Conclud i ng note, 428 8.3.2.6 Metastemis 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 ]an Pettersson (Thomson) 9.1 Introduction, 445 (Hymenoptera: 9.2 Aposematism and Reflex Bleeding Eulophidae. Chemistry. 445 Tetrastichinae), 408 9.2.1 Reflex bleeding, 445 8.3.2.9 l^ediobius foveolatus 9.2.2 Reflex bleeding substances, 445 (Crawford) (Hymenoptera: 9.2.2.1 Experiments. 445 Eulophidae, 9.2.2.2 Identification. 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 modifications, 447 Plant Volatiles. 45 7 9.2.2.6 Species specific 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 /. P. Midland and James D. Harwood semiochemicals, 450 10.1 Introduction. 4 6 6 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 modified 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-specific 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-specificity of oviposition deterrence pheromone substances. 454 11. COCCINELLIDS IN BIOLOGICAL 9.5.3 Active substances in oviposition CONTROL, 488 deterrence pheromone /. P. Midland tracks, 455 11.1 Introduction, 489 9.5.4 Relation to other aphid enemies. 45 5 11.2 The Roles of Coccinellidae in Biological 9.5.5 Aphid abundance, 456 Control, 489 9.6 Egg and Pupa Protection, 456 .1.1.2.1 Prey specificity. 489 9.6.1 Protection of eggs. 456 1.1.2.2 Generalist coccinellids, 490 9.6.2 Protection of pupae, 456 11.2.3 Intraguilcl predation. 490 xvi Detailed contents

11.3 Scale Insects 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 I nvasive 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 E 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 1.1.5 Conservation. 498 AND SUBFAMILIES, 526 11.5.1 Alternative or supplementary Oldrich Nedved and Ivo Kovaf food. 499 Coccinellidae Latreille. I 807. 527 11.5.2 Hibernation refuges. 500 Chilocorinae Mulsant. I 846 [2:25], 527 11.5.3 Habitat management. 500 Coccidulinae Mulsant. 1846 [6:46]. 527 11.5.3.1 Strip-harvesting. 501 Coccinellinae Latreille. I 807 [5:94], 528 11.5.3.2 Floral diversity (non-crop Epilachninae Mulsant. I 846 [4:24]. 529 plants). 501 Exoplectrinae Crotch. I 874 [2:24], 529 11.5.3.3 Intercropping, 501 Microweiseinae Leng, I 920 [4:23], 529 11.5.3.4 Reduced tillage. 502 Ortaliinae Mulsant. 18 50 [2:14], 530 11.6 Ancillary Factors Influencing Biologic; Scymninae Mulsant. 1846 [11:5.1], 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 bet ween pp. 250 and pp. 251 SUBJECT INDEX

Note: Page numbers in bold face refer to tables, page numbers in italic face indicate figures and illustrations. Abbreviations used: LDT for lower development threshold: SET for sum of effective temperatures: SCP for super cooling point;

Aaron's rod see Verbascum thapsus Acyrthosiphon pisum 57, 58-65, 3 52, 344, 356-358. 363. Abgrallaspis cyanophylli 60, 87, 91, 74-75. 78. 115. 118. 120, 376. 377. 378, 391. 392, 190, 204 145. 147. 152-154. 157, 398.399,407.407,408. Abies balsamea 471 158. 760. 161, 161-163, 412.413.414,416.417, abundance of coccinellids 164. 168-9. 168, 170. 171, 417.420.422.425.449, absolute and relative 111 172. 174. 182, 188-195, 4 51, 454-458. 477. 498, and character of landscape 127 196, 206. 208, 218. 220, 504. 505. 508 on crops 129, 130 222-224.234,236,237. adaptation ability 157 and habitat fragmentation 238.240.297. 345.451, aphid prey 153 126-127 457.469, 534 capture of 238,239 link to aphid abundance 117, 118 ant attendance 236. 237 development on different 145, methods of estimating 468^72 consumption rates 260 3 47 reduced tillage 502, 503 essential prey 158,162,272, rejected/problematic 165—168 sampling methods 112-114 188-195 toxic, effects of 161,162, transgenic crops 122-123 larval period and pupal mass, effect 452 on trees 131. 132 on 161 augmentation 498 Abutilon theophrasti 227.227 nutritive value 196-197 Coccipolypus hippodamiae. infection Acacia 154.183.489 pea surface wax 217-218 by 423 Acalypha ostryaefolia 123.126,227, ratio between disturbed and colour pattern variation 21, 22, 227 consumed 240 2 J. 25 acarina visual ability 239 assortative mating 27 Coccipolipus 411-414 Adalia bipunctata 20. 21, 23-32, 23, evolution of 26 phoretic mites 345.411 30, 36. 37. 39, 55-57. 58. 68. geographic variation 23-24 accepted food/prey 143, 144, 145, 69. 72. 73. 77, 78, 79. 82-85. diapause regulation 290-291 146 87. 89. 90. 91.93,96. 113, dorm a ncy behaviour 313-314 Aconitum 165 117. 119-123. 121. 126, egg ca nnibalism 17 7-178.233 Acraea encedon 425 128. 129, 129-132. 144- essential foods 188 Acrocephalus schoenobaenus 379 146. 150. 151, 151-154. expansion in Japan 493-494 Acutaspis umbonifera 190 153. 159. 162, 163-169, inbreeding 40 Acyrthosiphon caraganae 58 7 65. 166, 172-174, 173, intraguild predation 350, Acyrthosiphon ignotum 58 179-180,205. 209. 211, i 51-352, 361 Acyrthosiphon kondoi 476 212.215. 216-217.224. larval attachment ability 218 Acyrthosiphon nipponicum see 230. 231. 236,291. 305, larval food consumption 202, Neoaulacorthum nipponicum 306. 308. 309, 311, 318. 203 241 320-322. 324-326. 329, preference for 452-453

Ecology and Behaviour of the Ladybird Beetles (Coccinellidae). First Edition. Edited by I. Hodek. H.F. van Emden, A. Honek. © 2012 Blackwell Publishing Ltd. Published 2012 by Blackwell Publishing Ltd.

532 Subject index 533

Adalia bipunctata, (continued) mean SCP 96 alfalfa lields male-killing bacteria, effect of ovarioles 78-82 alternative prey 1 70-172. 213 421. 422, 424, 424, 425, 425 oviposition 85-88 nutritional value 196-197 mtDNA and male-killing bacteria pre-oviposition period 77-78 strip-harvesting 501 30-31 size 78.95 alfalfa weevil see Hypera postica non-insect food 180.185 teneral development 76-77 alkaloids 376 olfaction 225 wings and flight 77 aphid toxicity 163-164,165 oviposition deterrence 228.229 aestivation diapause 276-277 egg and pupae protection 76. 456 larval tracks, chemical Afidenta misera 79 enemies and competitors 449 composition 455 Afissula rana 79, 316 host plant 167 search patterns 275. 220 Afissula sanscrita 79 larvae protection 357 sperm competition/paternity studies Aframomum melegueta 24-26 reflex blood 376, 377. 445-449 42-43 age of coccinellids alkanes. larval tracks 455 trophic egg laying 173. 177 growth curves 74 AUanionematidae 414 wing development 28 hatching rate 68 allelobiosis 458 Adalia conglomerate 131 oviposition rates 86, 87 alleloc hemicals 151.155,196 Adalia decempunctata 21. 23. 25, 27. at peak oviposition 86 and aphid toxicity 158. 160-161. 30. 39.42.43,45.46.48, pupal, respiration rate 75 164-165 59.79. 84, 85. 100. 129, and sexual activity 91 DIMBOA in wheat 164. 169 131. 136, 151. 161. 166, and variation in defensive chemistry evolution of adaptive resistance to 189. 196,231, 233.236. 447 166 238.248. 309. 339. 347, and willingness to mate 82 and prey rejection 165-166 365.376.377.378. 392, Agelastica coerulea 167 allelopathy 458 399.407,408.412.4.13. aggregations AUoncda dodecaspilota 79 420, 422. 440. 452. 454, chemical implicated in 451.453, allozymes 460 457 method of analysis 34 Adalia deficiens 387,391 dormancy 302, 305-309 population genetics 40 Adalia fasciatopunctata see Adalia monospecitic 308 sperm competition 42^3 bipunctata aggregative numerical response Alnus• japonica 167 Adalia flavomaculata see Lioadalia 211-213.227 Alternaria 180-181. 183, 184 flavomaculata Agistemus longisetus 490 alternative foods/prey 146, Adalia tetraspilota 79 Agonoscena pistaciae 195 170-174, 187, 302, 344 additional prey, aggregative response Agonwn dorsale 474 biological control 490 212-213 agricultural practices conservation 499-500 Adelges cooleyi 189 changes in, effects of 126-127 pollen 180 Adelges laricis 188 habitat management 500-502 altruistic behaviour 451 Adelges nordmannianae 189 Agropyron desertorum 207 Amaranthus hybridus 227, 227 Adelges nusslini see Adelges Aiolocaria hexaspilota 59. 79, 82, Ambh/seius andersoni 353 nordmannianae 90. 167, 189. 197. 197, 221, Arnl)lyseius fallacis 490 Adelges piceae 493 225. 259.308. 310. 311. Ambrosia artemisiifolia 152 Adelges tsugae 66, 153, 175. 311. 321,340.420,426, American hickory see Carpinus 193-196, 344.471.493 428.437 caroliniana adelgids dormancy sites 311 Aminellus see Cowperia control of 493 Aiolocaria mirabilis see Aiolocaria Aminellus sunmtraensis see Cowperia as prey 175 hexaspilota swnatraensis Adonia arctica see Hippodamia arctica air-currents, use of 302 Amphitetranychus viennensis 67 Adonia variegata see Hippodamia alarm pheromone 225, 239. 451. Amplified Fragment Length variegata 468 Polymorphism (AFLP) analysis Adonis ladybird see Hippodamia Alauda arvensis 379 34 variegata Alcaligenes paradoxus 421 Amiyasta kuelmiella see Ephestia adult(s) 76-91 alder leaf beetle see Agelastica coerulea kuehniella defence of 357-358 Aleurodicus cocois 169 Anayyrus australiensis 384 fecundity 88-90 Aleurodicus dispersus 490 Anaayrus 353, 384. 386, 471. 496 longevity 90-91 Aleurotuba jelinekii 190 Anwjyrus kamali 496 mating 82-85 Aleyrodes proletella 191 Amujyrus lopezi 471 534 Subject index

Anagyrus pseudococci 353 interaction with ladybirds 236- ant attendance 236-238. Anastatus 386 238. 359-360 502-503 Anatis halonis 174 semiochemical repellence 449 biological control of 492-493, Anatis labiculata 20, 132 Aonidiella aurantii 60. 190 503-504 Anatis mali 122. 185, .189. 208. Aonidiella orientalis 60, 189 colony age and oviposition 306.471 Aonidimytilus albus 60 227-228 Anatis ocellata 23. 56, 59. 79, 80, apex predators, coccinellids as defensive and escape behaviour 85, 102, 122, 128. 131. 146, 354-355 2)8-241 149. 151. 151, 189.214, Aphanogmus 387 dropping behaviour 118 221.223,256, 314. 355. aphid abundance/density 116 reduced size, effect on predation 389.392,399.406.407. and age of host plant 122 505 408 and climatic changes 115—116 reproduction 492 Anatis quindecimpunctata 130. 132 link to coccinellid abundance social (soldier) 382 anatomical state, diapause 117, J78 suppression of 467.472^73 fat body and digestive tract and oviposition 117.119—120, toxic 157-165 316-317 456 Aphis carduella 192 flight muscles 323 simulation of timing of peak Aphis cirsiiacanthoidis see Aphis fabae male gonads 321-323 7 26 cirsiiacanthoidis ovary and spermatheca 317—321 aphid alarm pheromone 225. 239, Aphis cruccivora cowpea aphid or Anechura harmandi 426 451.468 groundnut aphid 59-62, Anegleis cardoni 59, 68, 86. 87. 91 Aphidecta obliterate 21. 45, 52. 57. 64-65. 68. 74. 78.87. 117. Anethum graveolens 121 85. 95. 108, 120, 129. 131, 7 18. 147. 148. 160. 161, Angoumois grain moth see Sitotroija 151. 151. 154. 189.205. 167. 189-195. 222. 236. cerealella 229. 230. 263. 271. 273. 237. 238. 453. 504 Anisolemnia dilatata see Megalocaria 308. 314. 325. 327.328. Aphis cylisorum cytisorum 191 dilatata 329. 339. 370.399.408. Aphis fabae cirsiiacanthoidis 58 Anisolemnia tetrasticta 452 415.431.443.458.462. Aphis fabae 58. 157. 158. 258. 259. Anisolobus indicus 420 463. 493. 509 160. 167-168 Anisosticta bitriangularis 167. 306, aphid parasitoids AphisJai inosa 63-66. 87. 91, 308 avoidance of plants with 1 17. 144, 190-195,222. Anisosticta novemdecimpunctata 23, coccinellids 353 223.237. 361.469.480. 56, 79. 129. 151. 151, 233. detection of footprint 49 5 378,399.422,442 semiochemicals 455-456 Aphis ghnines 59 Anisosticta sibirica 392 aphid population cycle, phases of Aphis gossypii 59-65. 73. 75, 88, Anisotylus Timberlake see Homalotylus 466-467 91. 148. 154. 161. 162, 164, annual mercury see Mercurialis annua aphid population dynamics 472 168. 178-179. 190-196, Anopheles quadrimaculatus 474 demographic-based estimates 197. 205. 210, 236.239. antennae 473^74 345.469 chemoreceptors on 450. 453 multi-species combinations survival, and increased CO2 levels subfamily characteristics 5-10 472-173 197 Anthus campestris 379 population decline, confounding Aphis hederae 188 Anthus pratensis 379 factors 466 Aphis helianthi see Aphis carduella Anthus trivialis 379 aphid resistant crops Aphis jacobaeae 163-165. 190. antibiosis in plants 504-505 Bt toxins, effect on ladybirds 446 antibody-based analysis of predation 505-506 Aphis lalmrni see Aphis cytisorum 474-477 synergism with coccinellids cytisorum antipodean ladybird see Harmonia 504-505 Aphis nerii 61. 68, 147. 148. antipoda Aphidentula bisquadripunctata 316 lhl-163. 179, 190-194, anti-predator defences Aphidius colemani 171, 400. 456 446 ladybirds 3 76-377 Aphidius eadyi 455 Aphis pomi 58-65. 153-154. 155, of prey 117-118 Aphidius ervi 232, 450, 455, 472 168. 188-195 ants Aphidoletes aphidimyza 352. 361 Aphis puiricae 191-193 attendance on honeydew producers aphids Aphis sailibuci 144-146. 145, 153, 236.237. 359, 362, alarm pheromone 225.239.451. 158. 158-159, 160. 260, 379-382. 502-503 468 163. 164. 166. 188. 196 Subject index 535

Aphis spiraecola 63, 66. 123. 146- associative learning 452—153 behaviour patterns in diapause 301 147. 149. 163, 176. 183. assorrative mating 27 dormancy phases 302-309 194.228.236,361-362 Asterolecanium sp. 163 individual species 309-316 Aphis spiraephaga 58 Atripkx sagittata 119 Bemisia argentifolii see Bemisia tabaci Aphis spiraephila 61. 64—65 Atritomellus see Dendrocerus Bemisia labaci 60, 87, 172. 189, Aphis urticata 190 attachment ability of larvae 218 196. 202 Apis mellifera 174, 179. 189, 193 augmentation of coccinellids Benthamidia florida 201 Apolinus lividigaster 79. 97. 183, 495-496 benzaldehyde 450 236.241. 243,266.297. dietary requirements 497-498 Berberis vulgaris 120 324. 333. 381.439 life stages for release 498 Beta vulgaris 121 diapause 297 mealybug destruction 496 Betula popnlifolia 152. 182 aposematic colouration 445 redistribution 496-497 Betulaphis brevipilosa 239. 239 anti-predator defence 376 source material for 497 Betulaphis quadrituberculata 188 chemical compounds 445-449 augmentative biocontrol 362 Bieri model, peak oviposition rate 86 enemies and competitors, impact on Aulacaspis tegalensis 60 big-eyed bug see Geocoris punctipes 449 Aulacaspis tubercularis 60 big-headed ant see Pheidole pupae 75 Aulacorthwn magnoliae see mcgaceplmla toxicity warning signal 358 Neoaulacorthum magnoliae biological control 150. 154. 222, apple see Malus pumila Aulacorthum solani 78. 153, 188 4S9. 524-525 apple powdery mildew see Podosphera Austroterobia 386 ancillary factors 502-509 sphaera leucotricha Aximopsis 388 ant-attendance 502-503 apple sucker see Psylla mali Axinoscymnus cardilobus 59. 75. 87. pesticides, selective use Aprostocetus esurus 388 88. 89. 90. 93, 96, 101. 189, 506-509 Aprostocetus neglectus 384 254 plant structure and chemistry Aprostocetus 384. 388. 409 Azteca instabilis 237,360. 381 504-506 Apus apus 379 Azya orbicjera 189. 23 7. 360. 381, seasonal cycles 503-504 Araneus diadematus 449 384, 442. 503 ants, disruptive impact of 362 Arctia 164 Azya trinitatis see Pseudoazya trinitatis and aphid population cycle 467 Argentine ant see Linepithema hwnile Azyini tribe 7. 9 augmentation of coccinellids Argentipilosini tribe 5 49 5-498 Arionater 477 Bacillus tliuringiensis 122, 123.421, coccinellid role 466. 489-491 Arion hortensis 477 505-506 Coccipolipus species, use in aromatic Apiaceae. conservation bacteria 41 5^14 501 general pathogenic 464 Col. maculata 152 arresting responses, plant volatiles male-killing 349,421-425 conservation 498-502 inducing 450,458 evolution of 38, 39 exotic introductions, targets of Artemisia tridentata 225 and failure of egg hatching 68 49 1-495 Artemisia vulgaris 117, 121, 126, molecular genetic studies facilitation and 361-362 501 29-3.1 intraguild interactions and 362 artificial diets 161.166,185-187 balsam fir see Abies balsamea intraguild predation and 361 and conservation 499-500 balsam woolly adelgid see Adelges large-scale field trials 477 artificial infestations, field cages 470 piceae Pediobhis foveolatus. use in artificial substrates for pupation 75 banding, chromosomes 14 410-411 Ascomycota, fungal pathogens 'barcode' DNA, species identification biological invasion 360, 361 415-418 29. 36 bird cherry-oat aphid see ash-gray lady beetle see Olla v-nigrum bark crevices, dormancy sites 314 Rlwpalosiphum padi Asian citrus psyllid see Diaphorina citri barn swallow see Hirundo rustica bird predaiors 3 77. 378. 379 Asian multi-colored ladybeetle see barred warbler see Sylvia nisoria bitter melon see Momordica charantia Harmonia axyridis Baryscapus 386. 409 bivoltinism Aspidimerini tribe 10 Baryscapus lhanasimi 386 A. bipunctata 320 Aspidiotus destructor 192. 236. 492 B-chromosomes and sex-ratio C. novemnotata 289 Aspidiotus nerii 60. 91. 163. 189. 15-16 C.s.brurki 286-287.287-288, 228 Beauveria bassiana 72, 359, 415, 288. 320 assemblages of coccinellids. 445. 498. 500 C. septenipunctata 278. 279. mechanism of 116-117 'beetle banks' 500 283 536 Subject index bivoltinism. (continued) Brachymeria carinatifrons 387 Calandrella cinerea 379 Har. axyridis 295 brachypterous individuals 77 California red scale see Aonidiella Hip. tredecimpunctata 292 Brassica campestris 207 aurantii black ant or garden ant see Lasius Brassica napus 58, 123. 165. 266. Callicaria superba 59. 71. 79 niger 225 Callipterinella calliptera 239 black bean aphid see Aphis fabae Brassica napus subsp. oleifera 58, Calocoris norvegicus 476 black citrus aphid see Toxoptera 123. 165, 266.225.452 Calotmpis procera 163. 190 aurantii Brassica nigra 166, 166. 452 Caltha palustris 313 black nightshade see Solanum nigrum Brassica oleracea 217 Calvia albida 79 black (or brown) mustard see Brassica Brassica oleracea Italica group 121 Calvia decemguttata 59. 79. 356 nigra Brevicoryne brassicae 61—64, 88, Calvia duodecimmaculata 59 black peach aphid see Brachycaudus 145. 148, 163. 165, 166. Calvia muiri 174, 189. 392 persicae 7 65-7 66. 166. 168, 7 68, Calvin quatuordecimguttata 37, 79, black redstart see Phoenicurus ochruros 190. 193, 225.451.468. 131, 313,412.455 blackcap see Sylvia atricapilla 490 Calvia quindecimguttata 37, 39, 55, black lady beetle see Rhizobius ventralis aphid alarm pheromone 225 59. 79,93, 102. 107. 128, blood-red lady beetle see Cycloneda control of 490 131, 132, 143, 146, 151, sanguinea release of toxic substance 154, 189 blue alfalfa aphid see Acyrthosiphon 451-452 essential prey 189 kondoi broad bean or faba bean see Viciafaba Calvia shiva 79 blue-headed wagtail see Motacilla flava broccoli see Brassica oleracea Italica Calvia shiva pasupati 79 blue tit see Cyanistes caendeus group Calvia shiva pinaki 79 bluethroat see Luscinia svecica brown apple mite see Bryobia Calvia shiva trilochana 79 body length 78.82 rubrioculus Campylomma verbasci 350, 353, body mass 73.74,82 brown citrus aphid see Toxoptera 532 body size 78 citricidus Canniibis sativa 152,182 defensive measure 356 brown mite see Bryobia rubrioculus cannibalism 175 dietary generalists and specialists brown planthopper see Nilaparvata non-sibling eggs 177-179 151-152 lugens siblingeggs 69.175-177,233. larva 73 Brumoides septentrionis 225 356 link to prey size & density Brunwides suturalis 59. 99, 189, Capiiophorus elaeagni 193 147-150 251, 507, 508 carbon dioxide concentration see also niche partition 345 (= Scymnus suturalis 39, 81. 129, CO2 concentration sibling egg cannibalism 176 308. 314) Aphis gossypii survival 197 and size of prey 119 Brumus quadripustulatus see and pupation timing 75 temperature affecting 9 5 Exochomus quadripustulatus Carduus crispus 216.217 and winter survival 331-332 Brumus suturalis see Brumoides Carinodulini tribe 5 'body size-dietary breadth' suturalis Carpinus caroliniana 152. 182 hypothesis 151-152 Brustiospora indicola 420 caryophyllene 451 body weight 78 Bryobia praetiosa Koch 167 cassava mealybug see Phenacoccus and egg size 56 Bryobia rubrioculus 167 manihoti and number of ovarioles 78, bryony ladybird see Henosepilachna cassava scale see Aonidimytilus albus 79-81, 82 argus catch per unit effort, sampling 522 Boettcheria latisterna 385 Budapest slug see Tandonia Caternaultiella rugosa 452 Brachiacantha quadripunctata 380, budapestensis Catolaccus 388 417,418 Bulaea lichatschovi 25, 142. 181. Caulnphijllum robustum 199 Brachiacantha ursina 130. 132, 306, 200, 246. 313. 378 Cavuriella konoi 58 380. 396.398 bullrush see Typha latifolia Centaurea jacea 216.217 Brachiacanthini tribe 10 butternut see Juglans cinerea Cemistes scymni 384 Brachycaudus helichrysi 153, 188, Cemistes subsulcatus 384 195 cabbage aphid see Brevicoryne brassicae Centistina nipponicus 383, 384, 391 Brachycaudus persicae 188-189, cabbage looper see Trichoplusia ni Centrosema pubescens 198. 299 195. 236 cage inclusion 470-471 Cephaloscymnini tribe 6 Brachycaudus prunicola 236 Calandra lark see Melanocorypha Ceratomegilla barovskii kiritschenkoi Brachycaudus tragopogonis 188 calandra 181 Subject index 537

Ceratomegilla notata 131, 181. chemical defences 38. 76, 355, 356 Chilocorus quadrimaculatus 385 281 chemo-mechanical defences, aphids Chilocorus renipustulatus 129. 190, food of 180-181 241 298, 320. 324, 325, 326. Ceratomegilla undecimnotata 59. 79. chemical markers, larvae 222. 226 327, 339.389,417 88,90, 96, 101. 106, 117. and oviposition deterrence Chilocorus rubidus 15, 79, 85, 105. 222, 131. 134. 146. 154, 228-233 190, 264.298.298. 324. 161, 162, 168. 169, 170. chemicals 325, 326.327, 328,329. 174. 189,220.223,229. see also allelochemicals: 339.420 230, 231. 249. 250. 255. semiochemicals Chilocorus similis 386.388 264, 266. 267, 268. 279. stimulating feeding behaviour Chilocorus spp., diapause 292.293.294.294,295, 199 298-299 298. 300,301, 302, 303. chemoreceptors on antennae 450, Chilocorus stigma 15. 16. 27. 18. 51. 305, 307. 308. 309, 310. 453 132. 168. 190,261,306. 311, 312, 314. 315. 316. Chenopodium 121 386,412. 417, 421, 509 317.328,320. 321, 322. cherry blackfly see Myzus cerasi B-chromosomes 16 323.324. 325. 326.327. Chetogena daripennis 385 chromosomal fusion 329.330.335, 336,338, Chilocorinae subfamily 5, 7-8, 9 polymorphisms 16-17 339. 341,372,378, 392, Chilocorini tribe 8 speciation and cytogenetic change 414,415. 420.453, 454. Chilocorus baylei see Chilocorus 18 455.460, 461,463. 500, malasiae Chilocorus tricydus 14, 17 507. 516 Chilocorus bijugus 79 Chinese jute see Abutilon theophrasti confused with C. septempunctata Chilocorus bipustulatus 60, 71, 73, Chinese lantern see Physalis alkekengi 311 79,93,99. 108, 109. 122. Chinese white pine see Pinus armandii diapause regulation 292-295 129, 131. 136. .138. 167. Chionaspis alnus 190 migratory and aggregation 168. 189. 220.221, 254. Chionaspis salicis 190 behaviour 309-310 272, 273.276.287,289. Chnootriba similis 384. 386. 427 Ceratovacuna lanigera 241-242, 382. 298.298, 299.299. 300, chorion, pores in 55 498 305. 314. 320. 322, 327. Choristoneura pinus 221 Cerchysiella 386 328.334. 336.337, 339, Chromaphis juglandicola 153. 188 Cervaphis quercus 205 341,386.387.388,389. chromatographic analysis 145. Cerylonid complex 2 404, 416.417,421.434, 165-166.225 Chaetosiphon fragaefolii 117 443.472,485,492, 513. chromosomal fusions 16-18 Chaitophorus capreae 145. 153, 153. 518 geographic distribution 17 188-189 Chilocorus braeti 79 chromosome numbers & banding 14 Chaitophorus leucomelas 188 Chilocorus cacti 384. 412 chromosomes & cytology 14 Chaitophorus versicolor see Chilocorus circumdatus 189 B-chromosomes 15-16 Chaitophorus leucomelas Chilocorus discoideus 384 cytogenetic changes 16-18 Chamerion angustifoliwn 211 Chilocorus distigma 384 sex determination 14-15 Chartocerus subaeneus 389 Chilocorus gerninus 299 chrysanthemum aphid see Cheilomenes lunata 189, 223. 233. Chilocorus hauseri 79 Macrosiphoniella sanborni 263,389.417 Chilocorus hexacyclus 14. 17 Chrysocharis johnsoni 384 Cheilomenes propinqua vicina Chilocorus infernalis 163 Chrysomela populi 189 386 Chilocorus inornatus 389 chrysomelids (leaf beetles) Cheilomenes sexmaculata see Chilocorus kuwanae 180. 190. 202, as prey 154,167,170-171 Menochilus sexmaculatus 220. 258. 261. 266.434. larval mortality/pupal weight Cheilomenes sulphurea 73,392 486, 503. 516 143 Cheilomenes vicina see Cheilomenes Chilocorus malasiae 190 tritrophic food chain 197 propinqua vicina Chilocorus nigripes 384 Chrysomphalus aonidum 60. 168, Cheiloneurus carinatus 387 Chilocorus nigritus 36, 60, 71. 78. 189 Cheiloneurus cyanonotus 387 79.86,87.88.90.91.92. Chrysomphalus bifasciculatus 190 Cheiloneurus liorhipnusi 387 93. 99, 103, 104, 105, 107. Chrysonotomyia appannai 384, 386 Cheiloneurus orbitalis 387 156. 157. 157. 190.203. Chrysopa 344,3 79 Cheiracanthium 478 204,224. 228,249,253. (7i rysopa oculata 161,352,456 chemical cues, plants as source of 265,267.492. 517 C'/irysopa perla 161, 252 120 Chilocorus orbus 14 Chrysopa sinica 225 538 Subject index

Chrysoperla carnea 347. 356, 361, Coccinella algerica see Coccinella 420.422.453.477. 480. 501 septempunctata algerica 507-508 Chrysoperla plorabunda 352 Coccinella californica 130. 201. 202. aggregative numerical response Chrysoperla rufilabris 75. 35.1, 353 209. 292. 420 211-213 Chrysophtharta bimaculata 170, 495 Coccinella explanata 174 alternative prey 170. 171-172. chrysopid larvae, attack by 358 Coccinella hieroglyphica 24. 79, 129. 173. 174.209 Chrysotachina slossonae 384-385 143. 143. 154. 190.253. aphid prey Cinara palaestinensis 189. 195 306. 313.392. 399 larval development 158.159, cinnabar moths see Tyria Coccinella leonina transversalis 60, 163. 768 circadian rhythms see rhythmicity 68. 69. 70, 70. 86. 117. 118. larval survival 149. 159, 163. Cirsium arvense 120. 153. 211, 118. 126. 146. 147. 163. 166 224.236.458 176. 178, 179. 191. 191, artilicial diets 186 Cirsium kagamontanum 198, 232 227. 228, 228. 229. 247. consumption rates 160. 201, Cirsium kamtschaticum 199 249. 263. 297. 356. 371, 202.205 cis-jasmone. plant volatile 447. 450 392.405.412.414,420. diapause regulation 278-287 citrus mealybug see Planococcus citri 478.485, 506. 510 dormancy behaviour 311-312 citrus mussel scale see Lepidosaphes diapause regulation 297 essential foods 190-191 beckii dormancy behaviour 313 exploitative competitors 348 Citrus sinensis 66 Coccinella luteopicta 79 foraging behaviour 215-2.18 citrus snow scale see Unaspis citri Coccinella magnifica 23. 38. 51. 79. ant attendance 236.237 Cladosporium 780-182. 183, 784 90. 151. 159, 190. 226, 269. intensive search 220,221 classification, subfamilies 317. 3Z8. 322. 367, 369, sensory perception 222, 223. changes in 3—4 373.380,392.398.399, 224 future perspectives 10.521 408.412, 413.436.441 vagility 234. 235. 236 morphologically based 3 Coccinella monticola 24 intraguild predation 210, proposed 5.6 Coccinella nigrovittata 183 351-352 Clematis 200 Coccinella novemnotata 24. 60. 90. introduction to North America Cleobora mellyi 170. 186. 392, 407. 103. 152. 154. 185.289, 493 489,499 289.290. 307.317. 318, larval development on aphid prey climate 337.392. 397.420.433. 158. 159, 163. 168 change, predictions 115-116 519 Ian al survival on aphid prey 149. and melanism 26 diapause 289-290 159. 163, 166 microclimate, effects on host plants Coccinella quinquepunctata 16. 90, life history as affected by aphid prey 124-125 129. 131. 151. 312. 328. 148 and voltinism 90 378.392, 398.399.420 non-insect food 780.181,184 Clitostethus arcuatus 60. 93. dispersal 309 overall predator voracity 255 103. 108, 154, 190. 244. hibernation sites 312 oviposition and aphid density 245 testicular activity 32 8.322 226.228 Clitostethus oculatus 56, 60, 71. 76. Coccinella reitteri 181 oviposition deterrence 229. 230. 77. 85, 86. 87, 89,93,96. Coccinella repanda see Coccinella leonina 231, 232-233 102. 106. 190.219, 258, Coccinella septempunctata 21. 30, 32, plant cues attracting 225 266.484. 490 39. 61. 69-70. 79.93. 276. prey capture 238.239.240 cloudberry beetle see Galerucella 723. 125. 129-132. 143. substitute diets 185. 186, 187 sagittariae 147-149, 151. J55. 758, tritrophic studies 196-197 clover mite see Bryobia praetiosa 159. 760, 163, 164. 166. voliinism 281.286-287.320 cluster size, eggs 56-57.58—67 366. 168. 272, 173. 280. Coccinella septempunctata algerica CO2 concentration, effects of elevated 186. 190-191, 197.205. 392 75. 197 222. 215, 276. 217. 221. Coccinella septempunctata brucki 191. Coccidoctonus trinidadensis 387 224.225.225.230.237. 221.276,287. 323.392. Coccidophilus citricola 385 238.240.240. 276.278. 395.420.422 Coccidula rufa 129, 131, 378, 422. 279-282.285-286. 287. aulolysis of flight muscles 323 424 306. 308. 311-312. 378. bivoltine life cycle 286-287 Coccidula scutellata 378 321.323. 330.347-348, diapause regulation 287-289 Coccidulinae subfamily 4. 5, 8-9 358. 376, 388-389.392, Coccinella sinuatomarginata see Coccidulini tribe 9 397. 399.407.412.417, Coccinula sinuatomarginata Subject index 539

Coccinella transversalis see Coccinella negative effect on hosts 41 3 nature of colour patterns 19-20 leonina transversalis prevalence 413 significance and origin 25—27 Coccinella transversoguttala 24. 6.1. Coccus hesperidum 15 7. 190 temporal variation 25 122. 191, 344.412 Coccus viridis 1 54. 189. 237. 360. colour vision 223-224 Coccinella transversoguttata richardsoni 503 commensalism 345. 347 24. 191 coconut scale see Aspidiotus destructor common Australian lady beetle see Coccinella trifasciata 61. 130. 306. Coelophora biplagiata 62, 74. 174. Codoplwra inaequalis 392.420 192.498 common barberry see Berberis vulgaris Coccinella undecimpunctata aegypiiaca Coelophora bissellata 56. 62. 80 common chiffchaff see Phylloscopus see Coccinella undecimpunctata Coelophora duvaucelii 80 collybila menetriesi Coelophora inaequalis 21. 62, 69, common cuckoo see Cuculus canorus Coccinella undecimpunctata 39. 62. 149. 163. 392. 497 common cutworm see Spodoptera litura 80. 130. 151. 154. 171. 191. Coelophora mulsanti 192 common damsel bug see Nabis 205. 353.389. 391,392, Coelophora quadrivittata 21, 192 (Reduviolus) americoferus 397,399.412.422. Coelophora saucia 62, 68. 69, 82. common froghopper see Philaenus 507-508 178. 192 spwnarius importation to New Zealand 492 co-existence of intraguild prey common redstart see Phoenicians Coccinella undecimpunctata menetriesi 360-361 phoenicurus 191 cold-hardiness 328-332 common starling see Sturnus vulgaris Coccinellidae family Coleomeijilla maculata 62. 74. 152- common stonechat see Saxicola characteristics 3 15 3. 773. 182. 313.354. 388 rubicola monophyly 2-3 cannibalism, egg 177-178 common swift: see Apus apus Coccinellimermis see Hexamermis consumption common whitethroat see Sylvia Coccinellinae subfamily food deprivation and larval communis characteristics of 6—7 development 207 community of coccinellid species molecular analyses 4-5 prey density effects on 111 coccinelline 446.449 consumption 204. 207 composition of. factors determining Coccinellini tribe 6. 7 dormancy behaviour 313 114 Coccinula crotchi 174. 417. 422 egg prey 170, 173. 173 dominance, diversity and niches Coccinula quatuordecimpustulata 23. essential foods 192 127-128 25,80, 167. 312, 378.392. pollen as 152,181-182,233 local faunas 114-116 420 frequency of short flights 234 locality determinants 116-12 7 Coccinula redimita 80 laboratory diet for 185 habitat identification 128-132 Coccinula sinensis 392. 417. 422 larval and adult diets 157 competition 347-349 Coccinula sinuatomarginata 25 nutrient requirements 186-187 apparent 349 Coccipolipus 412 olfaction 224 exploitative 347 Coccipolipus africanae 412 oviposition 227 competitive displacement of native Coccipolipus arturi 412 deterrence 229.237 ladybirds 494-495 Coccipolipus benoiti 412 periodicity 218.279 competitive IGP hypothesis 351 Coccipolipus bifasciatae 412 vagility 234. 235 complex communities, coccinellid Coccipolipus cacti 412 Coleomegilla maculata fuscilabris impact 472-474 Coccipolipus camerouni 412 149 conditioning, colour vision 224, Coccipolipus chilocori 412 Coleomegilla maculata lengi 143. 226 Coccipolipus cooremani 412 182. 354. 532 conidia (fungal spores) 183. 7 84 Coccipolipus epilaclmae 412 Coleomegilla quadrifasciata 391 conservation 360-361.498-502. Coccipolipus hippodamiae 412 Colorado potato beetle see Leptinotarsa 524 Coccipolipus macfarlanei 412 decemlineala alternative or supplementary food Coccipolipus micraspisi 412 colouration 499-500 Coccipolipus oconnori 412 aposematic 358. 376 biocontrol approach 362 Coccipolipus solanophilae 412 eggs 55. 56 habitat management 500-502 Coccipolipus species (parasitic mite) pupae 76 hibernation refuges 500 411 coloured cards, sampling 113 ennspecific larval tracks in biological control 413-414 colour pattern variation 18. 43-44 egg clustering 454—455 hosts and distribution 412 genetic determination 20-23 oviposition deterrence 222. 229- life cycle 412-413 geographic variation 23—25 233.356.456 540 Subject index consumed food, conversion and crested wheatgrass see Agropyron C-value, genome size 14 utilization of 209-210 desertorum Cyanistes caeruleus 376 consumption quantity 201-202 critical photoperiod 276,284.290. Cycloneda ancoralis 192, 330 assays 467-468 291,298. 299 Cycloneda limbifer 192, 223. 229, growth and reproduction crops 229, 230. 231, 232,454, adult performance 208-209 see also alfalfa fields 455 larval development 207-208 agricultural practices 127 Cycloneda nmnda 120, 129, 130, physical factors aphid resistant cultivars 164. 132. 152, 173, 192, 305. daily consumption rate 202 169, 504-505 392 397.417.418 total food consumption coccinellid abundance 130 Cycloneda polita 130, 132 202-204 diversification 126 Cycloneda sanguinea 15, 71. 119. and prey density 204—207 food spray substitutes 187 149. 161, 162, 163. 174. Contarinia nasturtii 170,497 ladybird habitats 128-129 175. 175. 179. 185, 192. Conura 387 microclimate effects 124—125 210.224. 357, 360-361. Conura paranensis 387 sampling from 112-113 377. 587.392.407.412, Conura petioliventris 387 strip cutting 126 417 507-508 Conura porteri 387 timing of colonization 503-504 development time and survival rate co-occurring prey 207 weeds in, positive effects of 126 162 Cooley spruce gall adelgid see Adelges Crotalaria striata 183 Cydonia vicina nilotica see Cheilomenes cooleyi crowding propinqua vicina coriander see Coriandrum sativum and egg viability 68,83 Cymbopogon c.itratus 181, 183 Coriandrum sativum 121 fecundity, adverse effect on 88 Cynegetini tribe 7 corn earworm see Helicoverpa zea of larvae, effect on adult size 78 cytogenetic changes 16-18.7 9 corn leaf aphid see Rhopalosiphum and ovarian dysfunction 82 maidis and pupal development 75 Dactylopius opuntiae 178, 192 corpora allata (CA) 278. 327-328 and take-off 77 daily fecundity 58-67, 86. 87. 90 Cosmos 201 cryptically coloured pupae 75. 76 damson-hop aphid see Phorodon Cotoneaster integerrima 120 cryptic species, delimiting 41—42 humuli Cotoneaster tomentosus 120 Cryptochaetum iceryae 1 72 Danaus plexippus 170,205 cotton aphid see Aphis gossypii Cryptognatha nodiceps 492 dandelion see Taraxacum officinale cotton bollworm see Helicoverpa Cryptognatha signata 383 Declivitata spp. 412 armigera Cryptognatha simillima 384 defence(s) cottony cushion scale see Icerya Cryptognathini tribe 9 ant attendance 236-238 purchasi Cryptogonus ariasi 80 aphid 2 38-241 Coturnix coturnix 449 Cryptogonus kapuri 391, 396. plant 200 Coturnix japonicus 445 401 defensive mechanisms, coccinellids couch grass see Elytrigia repens Cryptogonus orbiculus 80 against ants 380 courtship behaviour. D. coccinellae Cryptogonus poslmedialis Kapur 80. against inlraguild predation males 394 316, 527-530 355-358 Cowperia areolata 383-384, 386- Cryptogonus quadriguttatus 80 against parasitization 394 387, 399-400 Cryptolaemus montrouzieri 36. 167, against Phalacrotophora 408 Cowperia indica 401 169, 185. 192, 218.222. aposematic colouration 358. Cowperia punctata 401 237. 353.383. 385-386, 376 Cowperia sp. (parasitoid) 383. 391. 391.401.492,496, 505 morphological adaptations 3 77 399,401. 401 control of mealybug and scale reflex bleeding 376-377 Cowperia subnigra 401 insects 496 Degeeria lucmosa see Medina luctuosa Cowperia sumatraensis 401 Cucujoidea 2 Delichon urbica 377,379 Cranophorini tribe 9 Cuculus canorus 379 Delphastus aitalinae 62, 68, 89. 91, cream-spot ladybird see Calvia Cucurbita maxima 126 93. 96.96. 192,202.219, quatuordecimguttata Culex quinquefasciatus 479 348. 353,495 cream-streaked ladybird see Harmonia Curinus coeruleus 174, 192. 217. Delphastus pusillus 56, 76, 85, 172. quadripunctata 223. 362.493.497. 175. 192,205, 361 creeping thistle see Cirsium arvense 507-508 Delphiniobiiiin junackianum 165 Crematogaster lineolata 76,456 cuticular cribriform plate, ductless Dendrocahmuis giganteus 163 Crematogaster 76. 381. 456 glands 454 Dendrocerus 388 Subject index 541

Dendrocerus ergensis 388 Dinocampus coccinellae (parasitoid) migration 302-305 Dendrocopos medius 379 383-384, 391 pre-diapause 301-302 Deroceras reticulatum 477 development 394-395 dormancy sites 282, 309-316 development flight performance, effect on emergence from 309 embryonic 56, 85 303-304 Doryphorophaga doryphorae see larval 72-73 host preferences and parasitization Myiopharus doryphorae pupal 75 rates 396-399 Drepanosiphum platanoidls 145, 153. temperature effects 91-9 7 host range and host suitability 153. 188-189.238 teneral 76-77 392-393. 396 drone honeybee powder, substitute development rate isomorphy 95 ladybird fertility, effect on 88 food 174 development rate and temperature ladybird ovaries, effect on 317 dropping behaviour 75, 82, 92. 93-94. 96 larval nutrition 395-396 aphid escape response 118. 238, development time life cycle 394 239.240,468 effect of CO2 on pupation 75 reproduction, mode of 391,394 coccinellid escape mechanism of larvae 72-73 semiochemical attraction 449 357. 358 and temperature 92 Dinocampus 383-384. 391 Drosophila melanogaster 28 Diabrotica virgifera 32 Dinocampus nipponicus see Centistina ductless glands, pheromones released diamondback moth see Plutella nipponicus by 454 xylostella Dinocampus terminatus see Dinocampus duration diapause/dormancy 2 76 coccinellae flight 77 see also individual species Diomus austrinus 89. 93. 95, 192 hatching 69 anatomical/physiological changes Diomus hennesseyi 143. 491 larval development 72, 73 anatomical state 316-323 Diomus pumilio 384 milling 82-83, 84 metabolic changes 323-327 Diomus seminulus 417 pupal stage 75 behaviour patterns 301 Diomus thoracicus 449 dusky lady beetle see Scymnus loewii dormancy phases 277. diploid chromosome numbers 14. D-Vac. vacuum sampling by 112 301-309 75 Dysaphis crataegi 64.194 individual species 309-316 dipterans. in IGP 352-353 Dysaphis devecta 192 endocrinological aspects 2 77—278 directional selection 28 Dysaphis plantaginea 62. 192 hibernation and aestivation direct observation 468. 469, 474, Dysmicoccus 194 276-277 475 Dytiscus sp. 241 research activity 523 disc equation, functional response termination/completion 277 206 (E)-(5-farnesene (EBF) 225. 447, 451 Diaphorina citri 174. 469 Discotomini tribe 6 early-season predation 477 Diaspidiotus perniciosus 60 dispersal flight, dormancy 302. 309 Echthroplectis see Homalotylus Dibrachys cavus 383,385-386. distribution 110-117,121-128. ECI (efficiency of conversion of 388. 405 132 ingested material) 209-210 Dicranolaius bellulus 478 vertical on host plant 722, 125 ecologically relevant temperatures 92 dietary complementation across life Diuraphis noxia 63, 66. 123. 146, ecophysiological regulation of stages 156-157 190. 192-195, 205. 206. diapause 278. 300-301 dietary generalists and specialists 466. 504 Adalia bipunctata 290-291 150-152 diversity, coccinellid 114 Apolinus lividigaster 297 diet(s) and geographical latitude Ceratomegilla undecimnotata see also food 114-115 292-295 combination and mixed 15 5-15 7 studies of 127-128 Chilocorus species 298-299 and egg cluster size 57. 58-67 tropical/subtropical areas 129 Cot cinella leonina 297 longevity, effect on 91 DNA-based gut content analysis Coccinella novemnotata 289-290 and reproductive output 78 144-145,478-481 Coccinella septempunctata 278 substitute and sprays 18 5-18 7 DNA sequence-based studies 36, Central Europe 2 78-282 digestion • 3 7-38 Mediterranean region 284-285 during dormancy 316-317 Dolichoderus bidens 76 Nearctic region 285-286 extra-intestinal 241 dominant species 127-128 Northern Europe 283-284 dill see Anethum graveolens dormancy phases 277 voltinism 286-287 dilution effect 350, 352. 353, 357 aggregations 305-309 Western Europe: France DIMBOA in wheat 164,169 emergence 309 282-283 542 Subject index ecophysiological regulation of elm leaf beetle see Xanthogaleruca Epilaclma nigrolimbata 412 diapause, (continued) luteola Epilaclma niponica see Henosepilachna Coccinella septempunctata brucki elytra niponica Central Japan 287-288 hydrocarbons 454 Epilaclma paenulata 76, 355. 447 Northern Honshu (Japan) 289 inheritance studies 21-22 Epilaclma philippinensis see Sapporo. Hokkaido (Japan) pattern, variation in 19-20 Henosepilachna 288-289 red morphs. mating disadvantage vigintisexpunctata Harmonia axyridis 295-297 26-27 Epilaclma pusillanima see Harmonia sedecimnotata 297-298 Elytrigia repens 120. 224, 458 Henosepilachna pusillanima Hippodamia convergens 292 embryonic development Epilaclma pustulosa see Henosepilachna Hippodamia tredecimpunctata 292 and egg size 56 pustulosa Illeis galbula 297 optimum microclimate for 85 Epilachna quadricollis 384 Propylea quatuordecimpunctata emergence from dormancy 309 Epilaclma septima see Henosepilachna 291 emigration 114. 211, 235. 357 septima Stethorus punctwn picipes Encarsia Sophia 353 Epilaclma sparsa orientalis 299-300 encounter with prey, effects of Henosepilachna edelweiss see Leontopodium alpinum 220 vigintioctopunctata egg cannibalism 68.69.175-179. endocrinological pathways, diapause Epilachna varivestis 73-74. 76. 187, 356 277-278 198, 200. 224. 384-389, egg-surface chemicals 455 endosymbiotic bacteria 29-31. 38. 405, 410-411 semiochemicals. role of 456 39, 41-42, 522 Epilachna vigintioctomaculata see egg production 88-90 enzyme-linked immunosorbent assay Henosepilachna effect of food restriction 209 (ELISA) technology 474. vigintioctomaculata multiple matings 82-83 475.477.478.479 Epilachna vigintisexpunctata 62. 384. and ovariole number 5 7 Eocaria muiri see Calvia muiri 411 rate of 86 Ephestia kuehniella 57. 58. 73. 153. Epilachna yasutomii see Henosepilachna egg protection behaviour 256 yasutomii oviposition on highly pubescent Epidinocarsus lopezi see Anagyrus lopezi Epilachninae subfamily 5. 7 plants 227 Epilaclma 'chrysomelina' see feeding stimulants 452 species-specific semiochemicals Henosepilachna argus or food of 198.200 456 Henosepilachna elaterii or pupal defence 76 egg retention 84 Henosepilachna Epilachnini tribe 7 egg(s) 55 vigintioctopunctata Episyrphus balteatus 347.356, 358 cluster size 56-68 Epilaclma admirabilis 72. 90. 405, Epiverta chelonia 7 defence of 356 411.452 Epivertini tribe 7 hatching rate 68-69 Epilaclma bisquadripunclata see Eremochilini tribe 7 meanSCP 96 Aphidenlula bisquadripunctata Eretmocerus mundus 353 morphology 55-56 Epilaclma boisduvali see Henosepilachna Eriococcus coriaceus 195, 241 nutrient content 233 boisduvali Eriopis connexa 126. 164, 164. 205, size 56 Epilaclma borealis 76, 385 224, 330. 391, 392.417. trophic eggs 69 Epilaclma canina 400 505 egg teeth 56 Epilaclma cucurbitae see Henosepilachna Eriosoma lanigerum 193. 205. eighteen spot ladybird see Myrrha sunibana 285 octodecimguttata Epilaclma defecta 384 Erysiphe cichoracearum 66. 91 Elasmus 388 Epilaclma dregei 316. 384 Erysiphe polygoni 124 Elatobium abietinum 189, 205 Epilaclma dumerili 80 Erythrina corallodendron 167 elder aphid see Aphis sambuci Epilaclma eckloni 384 essential foods/prey 746. 187. 196 elder see Sambucus nigra Epilaclma enneasticta see list of 188-196 electroantennograms (EAG) 224, Henosepilachna enneasticta psyllids 174 225.453 Epilaclma eusema 385, 387 Eucallipterus tiliae 58. 153. 153, electrophoretic methods 327-328 Epilaclma karisimbica 412 157. 177. 188-189. 195. eleven spot ladybird see Coccinella Epilaclma marginella 385 238-239 undecimpunctata Epilaclma marginicollis 80 eucalyptus leaf beetle see ELISA technology 474,475.477, Epilaclma mexicana 384. 387 Chrysophtharta bimaculata 478,479 Epilaclma mystica 80 Euceraphis betulae 58,238.239 Subject index 543

Euceraphis punctipennis 1 53. 168. 192.313-314. 329, firebug see Pyrrhocoris apterus 188-189 377. 389, 393, 420 firethorn see Pyracantha coccinea eugregarines. protozoan pathogens Exochomus troberti see Parexochomus fireweed see Chamerion angustifolium 419^2] I roberti first i ibosomal internal transcribed Eulecanium caraganae 190 Exoplectrinae subfamily 7 spacer (ITS 1) region 31-33 Euonymus japonicus 202 Exoplectrini tribe 7 fitness 56, 86 Eupelmus 385-386, 388 Exoristoides slossonae see Chrysotachina adaptive food preferences Eupelmus urozonus 386 slossonae 452-453 Eupelmus vermai 385 extinction 115 cannibalism 176 Euphorbia 200 extralloral nectaries 1 83. 499. fitness compensation 424 Eupteromalus Kurdjumov sec 504 five spot ladybird see Coccinella Trichomalopsis Crawford eyed ladybird see Anatis ocellata quinquepunctata Eurasian blackbird see Turdus merula eyespotted lady beetle see Anatis mali Elavobacterium 68, 176. 422-423 European black slug see Arion ater flies, dipteran parasitoids 406-408 European corn borer see Ostrinia facilitation, predator 361—362 flight 77 nubilalis facultative diapause 276. 281. 292, frequency of short 234 European earwig see Eorficula 293, 316 habitat switching 12 7 auricularia faeces, oviposition deterrence 231. in the landscape 234-235 European honey bee see Apis 232 migratory 303-305 mellifera fall webworm see Hyphantria cunea flightlessness 77 European red mite see Panonychus fat body flightless mutants, release of 498 ulmi diapause 316-317 flight muscles, diapause 304, 321. European red wood ant see Formica site of synthesis of aposematic 323 polyctena substances 445 floral nectar, sugar as substitute 187 Eurotia ceratoides 200 fat reserves during diapause Floridii red scale see Chrysomphalus euryphagous species see generalist 323-325 aonidwn species 150-152 faunas of coccinellid communities fluorescent in situ hybridization (FISH) Euthelyconychia epilachnae 384. 114 techniques 15 387-388 climatic changes, effects of follicles, ovarioles 57.78.82 evolutionary history studies 115-116 food phylogenetics 37-38 geographic differences 114-115 egg cluster size 57.58-67 phylogeographic studies 38-42 invasion & extinction 115 fecundity link 88 population genetics 38-41 fecundity 87. 88-90 larval instars 73 sequence evolution 29-33 daily and lifetime 58-67 longevity link 91 techniques 33-37 diet affecting 88.91 and oviposition period 88 exclusion cages, predator impact study oviposition rate 86 preferences, evolution of 38 469-470 temperature, effect of 96 and pre-oviposition period 78 Exochomus childreni 174, 192, feeding habits 2 food consumption 361 feeding studies, laboratory 467-468 functional response 204—207 Exochomus concavus see Parexochomus females growth and reproduction troberti concavus finding oviposition site 226-232 207-209 Exochomus flavipes 85, 178, 192. food consumption, effect on physical factors 202-204 313.384.387.389 reproduction 208-209 food conversion 209-210 Exochomus flaviventris 143, 169. preference for melanic morphs food intake 241-242 169.222,384.404 27. 83-85 food range 142-144 Exochomus fulvimanus 412 size/weight 75, 78 methods for detection of 144-145 Exochomus lituratus see Priscibrumus Ferrisia virgata 59. 189 food-related behaviour lituratus fertilization success 42-43 food intake 241-242 Exochomus melanocephalus see fescue aphid see Metopolophium foraging behaviour 213-238 Parexochomus melanocephalus festucae prey capture 238-241 Exochomus nigromaculatus see Ficedula parva 379 food-related semiochemicals Parexochomus nigromaculatus field cages, predator impact studies 449-452 Exochomus quadripustulatus 16, 27. 469-470 associative learning 452—453 32, 57.62. 74. 80. 86.87, fifteen spotted lady beetle see Anatis phagostimulants 452 87.96. 131-132, 161, 164, labiculata plant volatiles 449-451 544 Subject index food-related semiochemicals. fourteen spot ladybird see Propylea Genista 167 (continued) quatuordecimpunctata genome size 4, 14 prey alarm pheromones 451 foxglove aphid see Aulacorthum solani Geocoris punctipes 118 toxic substances in prey French bean see Phaseolus vulgaris geographic variation 451-452 fruit, damaged, food source 123 in colour patterns 23-25 food relationships 142 fruit tree spider mite see diapause expression 332 food-related behaviour 213-242 Amphitetranychus viennensis latitude and ladybird diversity quantitative aspects 201-213 functional response 204-207, 114-115 summary 242 471 giant bamboo see Dendrocalamus food specificity 142-201. 746. 777, fungal pathogens yiganteus 214, 242. 523 Hesperomyces species 308-309, 'gin traps' 357, 377 alternative food 746, 147. 150, 416-418 glacial lady beetle see Hippodamia 170-174. 172. 180. 187. Hypocreales 415-416 glacialis 214, 242. 302, 344 Nosematidae 418-419 glasshouse potato aphid see cannibalism 175-179 fungal spores, food source 180-184 Aulacorthum solani essential foods 1 87-196 glasshouse whitefly see Trialeurodes food range 142-145 Galeruca interrupta arminiaca 167 vaporariorum generalist vs. specialist species Galerucella lineola 195 glassy-winged sharpshooter see 150-155 Galerucella pusilla 152 Homalodisca vitripennis lower qu ality prey 157-169 garden pea see Pisum sativum Glochidion ferdinandi 183 mixed and combined diet garden slug see Arion hortensis glossy leaves, biocontrol 504 155-157 garden warbler see Sylvia borin glucosinolates 165-166. 451-452 mycophagous Coccinellidae. food of Galerucella sagittariae 143. 190 glycogen reserves, diapause 200-201 Galleria mellonella 328 325-326 non-aphid prey 169-175 gas chromatography-mass glycosides 160, 162, 169. 241 non-insect food 180-185 spectrometry (GC-MS) 448. golden-eyed lacewing see Chrysopa nutritional suitability 145-147 456 oculata phytophagous Coccinellidae. food of gean or wild cherry see Primus avium golden loosestrife beetle see Galerucella 198-200 gel electrophoresis 34.145. 478,480 pusilla prey size-density hypotheses Gelis agilis 388 gonads. diapause/dormancy 317, 147-150 Gelis instabilis see Gelis agilis 321-322 substitute diets and supplements Gelis melanocephalus 388 gorse spider mite see Tetranychus 185-187 Gelis 388 lintearius tritrophic studies 196-198 generalist (euryphagous) species grain aphid see Sitobion avenae food sprays 187 150-152 grains of paradise see Aframomum footprint semiochemicals, aphid Adalia bipunctata 153 melegueta parasitoid response to Coleomegilla maculata 152-153 green apple aphid see Aphis pomi 455-456 Harmonia axyridis 153 green citrus aphid see Aphis spiraecola foraging behaviour 213-214 Hippodamia species 154 green coffee scale see Coccus viridis ants, interaction with 236-238 other generalists 154 green lacewing see Chrysoperla carnea first instars 233 role in biological control 489, 490 green shield scale see Pulvinaria psidii indirect factors 215-218 genetically modified (GM) crops greenbug see Schizaphis graminum diurnal periodicity 218 122-123, 505-506 Gregarina barbarara 420,421 plant structure 215-217 genetic correlations between traits Gregarina californica 420 plant surface 217-218 28-29 Gregarina chilocori 420 larger scale 23 4-2 3 6 genetic markers see molecular genetic Gregarina coccinellae 420 oviposition site, finding 226-233 markers 29 Gregarina dasguptai 420 research progress 520-525 genetic studies 14, 43-44 Gregarina 419,420.421 senses, role of 218-226 chromosomes & cytology 14-18 Gregarina fragilis 420 Forficula auricularia 472 colour pattern variation 18—27 Gregarina hyashii 420 Formica 379 future research 521-522 Gregarina katherina 420 Formica obscuripes 380 host plant use by phytophagous Gregarina ruszkowskii 420 Formica polyctena 221,380. 449 species 198-199 Gregarina straeleni 420 Formica rufa 449 inheritance of other traits 27-29 gregariousness. defensive mechanism Formica rufibarbis 236 molecular genetics 29^3 307-308, 357 Subject index 545 grey field slug see Deroceras reticulatum 122-123. 129, 130. 132. Harmonia quadripunctata 80, 151. grey garden slug see Deroceras 149. 153. 755. 760. 161. 393,399.412,420,423 reticulatum 168, 170. 172, 173. Harmonia sedecimnotata 80, 193, guilds 344 173-174. 275. 277. 782. 297 spatial 344, 352. 363 185, 186. 193. 203. 205. diapause 29 7-298 temporal 344-345. 352. 361. 207. 208. 209. 222, 223. Harmonia yedoensis 63 363 228,232.295.296-297. Harpalus pennsylvanicus 345, 347, thermal 344 306. 310. 331. 349.350. 472 gustatory sense 142, 226, 242 352. 354. 377. 386, 389. hatching rate 68-69 gut content analysis 144-145. 392, 397.407,412.417. and lifetime fecundity 69 180-181.478-481 422,457.494.495. 507- and mating duration 82 508. 532. 535 and multiple matings 82-83 habitat management 500-501. 524 alternative food, effect on egg laying and polyandry 83 floral diversity, non-crop plants 777 hatching synchrony 56,68-69 501 artificial diet 185. 786 hawthorn-parsnip aphid see Dysaphis intercropping 501-502 colour pattern variation 21-22. crataegi reduced tillage 502 22,23-24.25.27 hawthorn spider mite see strip-harvesting 501 diapause regulation 295-297 Amphitetranychus viennensis habitat preferences and displacing native ladybirds hazel aphid or filbert aphid see semiochemicals 457-458 494-495 Myzocallis coryli attractive plants 458 dormancy behaviour 3.1.0—311 heather ladybird see Chilocorus avoided plants 458 essential foods 193 bipustulatus plant-plant interactions 458 flightless strain 498 Helianthus animus 499. 499. 535 plant stand traits 457-458 food consumption, effect on growth Helicobia rapax 385 role of plant volatiles 457 207-208 Helicoverpa armigera 170, 478. habitats 110-132 intraguild predation 210. 350. 202 artificial substrates as 75 351-352 Helicoverpa zea 1 70. 182, 202. 477, crops 126.128-129.130 introduction of 494 502 fragmentation 126-127.234 invasion, genetic studies 40-4.1 Heliothis virescens 152, 477 movement among 234-236 routes 42 helmet scale see Saissetia coffeae niches 128 olfaction 222. 223. 224-225 Hemaenasioidea see Homalotylus sampling methods 111—114 overall predator voracity 255 Hemberlesia lataniae 60 spatial guild partition 344 oviposition deterrence 229. hemipterans (non-aphid) as prey switching between 127 231-232 174-175 trade-off in selection of 357 pest status 495 hemiptera-tending ants 379-382 trees 129,131.132 sperm competition/paternity studies Hemisarcoptes 411 tropics 129. 132 42,43 Hemisarcoptes cooremani 347. 412 wild herbs 129.131 survival rates of larvae fed on hemlock woolly adelgid see Adelges hairs, pupal defence 76. 357. 456 aphids 147, 149 tsugae Halmus chalybeus 57 top predator 354 Henosepilachna argus 56 Halyzia hauseri see Macroilleis hauseri trophic egg laying 173,177 Henosepilachna bifasciata 412 Halyzia sanscrita 80 unsuitable aphid diet 160-161 Henosepilachna boisduvali 76 Halyzia sedecimguttata 39, 131. vagility 234,235 Henosepilachna dodecastigma 80 392,399 vision 223-224 Henosepilachna elaterii 21,22, 198, Halyzia straminea 80 wing polymorphism 28 387,412 Halyzia tschitscherini 314 Harmonia breiti see Harmonia Henosepilachna cjuttatopustulata 400 Halyziini tribe 6, 7 expalliata Henosepilachna indica 80 hand-picking/hand-shaking, sampling Harmonia conformis 154, 193,205. Henosepilachna niponica 85-86. 232, methods 112 311, 393. 407 332. 404. 405, 427 haricot bean see Phaseolus vulgaris dormancy behaviour 311 Henosepilachna ocellata 80. 400 harlequin ladybird see Harmonia Harmonia dimidiate 80, 162, 193, Henosepilachna processa 80 axyridis 205, 229. 230. 393 Henosepilachna pusillanima 76, 80 Harmonia antipoda 392 Harmonia eucharis 80, 407 Henosepilachna pustulosa 42. 405. Harmonia axyridis 21, 22, 32, 41. Harmonia expalliata 407 420, 427 63, 80, 87. 93. 123, Harmonia octomaculata 393 Henosepilachna septima 427 546 Subject index

Henosepilachna species 198, 799 biological control, use in 490, Homalotylus albidavatus 383-384. Henosepilachna sumbana 64 492. 497 386-387. 399-400 Henosepilachna vigintioctomaculata consumption of mummies 7 73 Homalotylus albifrons 383-384, 19. 29. 80. 85.400.404, diapause regulation 292 386-387, 399-400 405.416.421,427.427 drinking extra-lloral nectar 499 Homalotylus albitarsus 383-384. Henosepilachna vigintioctopunctata egg masses laid 231 386-387, 399-400 64, 80.80, 299.384.386. essential foods 193 Homalotylus aligarhensis 383-384, 388, 399,400.405.410. food deprivation 207 386-387, 399-400 412.427.427 frequency of short flights 234 Homalotylus balchanensis 383-384. Henosepilachna vigintisexpunctata migration and aggregation 387. 399-400 384.411 314-316 Homalotylus brevicauda 391, 396, Henosepilachna yasutomii 85, 198, redistribution of overwintering 400. 402-403 427 496^97 Homalotylus californicus see herbaceous stands 128 survival rates of larvae fed on Homalotylus terminalis sampling from 112 aphids 147. 149 Homalotylus cockerelli 391. 396. wild herbs 129,131 total egg production as function of 400. 402-403 herbivore-stressed plants, volatiles food provisioning 7 56 Homalotylus ephippium 383-384, released from 450. 459 Hippodamia glacialis 130 387. 399-400 heritability studies 28-29 Hippodamia parenthesis 130. 132, Homalotylus eytelweinii 383-384. Hesperomyces chilomenis 417 193. 306.308.393. 387. 399-400 Hesperomyces coccinelloides 418 397 Homalotylus ferrierei 399-400 Hesperomyces hyperaspidis 402-403. Hippodamia quinquesignata 22, 193. Homalotylus flaminius 383-384, 417 316.422.423 387. 399-400 Hesperomyces species 84.308, diapause behaviour 3.16 Homalotylus formosus 383-384, 416-418,417.428 subnivean hibernator 329 387. 399-400 Hesperomyces virescens 84, 308. Hippodamia septemmaculata 80 Homalotylus hemipterinus 383-384. 416 Hippodamia sinuata 22. 130. 132, 387. 399-400 Heteropsylla cubana 192. 217, 223 193.420 Homalotylus himalayensis 469 heterospecific aggregations 308 Hippodamia tredecimpunctata 25, 26. Homalotylus hybridus 400 Hexamermis 415 64.80. 225. 129-130, 151. Homalotylus lujperaspicola 406 hibernation 276 193. 292. 306. 308. 378. Homalotylus hyperaspidis 391, 396. see also diapause/dormancy 389.393.420 400. 402-403 mating frequency and longevity diapause regulation 292 Homalotylus hypnos 388-389, 91 Hippodamia variegata 20, 23. 24. 25. 391.400-401.402-404, movements prior to 12 7 3 3. 36.40,64, 77.80.87. 428 mycophagous and phytophagous 90. 117, 121. 222. 126-129. Homalotylus indicus 399-400 species 316 129. 130-131. 145. 146, Homalotylus latipes 399. 400. refuges 500 151. 162, 170. 177. 194. 409 second 90 211. 235. 307, 309. 312, Homalotylus longicaudus 383—384. Hibiscus 345.496 330. 353. 360. 377.378, 387, 399-400 hieroglyphic ladybird see Coccinella 386. 389, 393. 398-399, Homalotylus longipedicellus 402 hieroglyphica 414.415.420.423.478. Homalotylus 383-384. 386-387. Hippodamia arctica 393 493 399-400 Hippodamia caseyi 316 hippodamine 448. 449. 456 Homalotylus mexicanus 391. 396. Hippodamia convergens 22, 29, 64, Hippolais icterina 379 400. 402-403 93. 223-225, 130. 132, 149, Hirundo rustica 379 Homalotylus mirabilis 400 256. 173-174. 186. 193. hogbrake see Ambrosia artemisiifolia Homalotylus mundus 383-384. 207.237,234.292.303. Homalodisca vitripennis 480 387. 399-400 305.306-308.314-315. Homalotyloidea dahlbomii 387 Homalotylus nigricornis 383-384. 376,387,393. 396.397, Homalotylus afflnis 383-384. 386- 387. 399-400 412.417,420. 497. 499. 387. 399-400 Homalotylus oculatus 399-400, 503. 507-508, 535 Homalotylus africanus 383-384. 409 aggregation, olfactory cues 307 386-387, 399-400 Homalotyluspallentipes 391. 396, artificial diet 186 Homalotylus agarwali 383-384, 400. 402-403 assassin bug preying on 497 386-387, 399-400 Homalotylus platynaspidis 400 Subject index 547

Homalotylus punctifrons 391. 396. humidity llleis koebelei 200. 207 400. 402-403 and food consumption diet of 200. 202 Homalotylus quaylei 39.1, 396. 400. 203-204 immigration 114. 115. 117. 211. 402-403 and hatching rate 68 235 Homalotylus rubricatus 383-384. longevity effect 91 immunolabelling 477-478 387. 399-400 hump earwig see Anechura harmandi impaction traps 113 Homalotylus scutellaris 383-384. Hyaliodes vitripennis 207, 353 inbreeding studies 40 387, 399-400 Hyalopterus pruni 58-59, 63. 66. Indian hemp see Cannabis sativa Homalotylus scymnivorus 406 91. 147. 147. 153, 165-168. Indian Mallow see Abutilon theophrasti Homalotylus shuvakhinae 400-401, 188. 190-191, 193. 236. Indian ricegrass see Oryzopsis 402 241. 252.267 hymenoides Homalotylus similis 405 hybridization 85 induction of diapause 280-281. Homalotylus sinensis 383-384. hybrid sterility 17-18 282-283.287 387, 399-400 hydrocarbons on elytra 454 critical photoperiod 290,291.299 Homalotylus singularis 400 hydroxamic acid DIMBOA 164 lack of food 293 Homalotylus sp. 400-401, Hypera postica 170. 171-172. 210. stage sensitivity 289. 290 402-403, 404 315,490 industrial melanism 25.26 Homalotylus terminalis 383-384, Hyperaspidini tribe 10 inheritance 387, 399-400 Hyperaspis aestimabilis 387 colour patterns 20-23 Homalotylus trisubalbus 383-384. Hyperaspis bigeminata 306 experimental evidence 21-22 387, 399-400 Hyperaspis binotata 132 life history characters 28-29 Homalotylus turkmenicus 401 Hyperaspis campestris 71, 80 wing polymorphism 27-28 Homalotylus vicinus 381 Hyperaspis congressis see Hyperaspis Inkaka quadridentata 385 Homalotylus yunnanensis 383-384. conviva innate capacity for increase. 387. 399-400 Hyperaspis conviva 380 temperature dependent 96 Homalotylus zhaoi 383-384. 387. Hyperaspis desertorum 194 insect growth regulators (IGRs) 509 399-400 Hyperaspis lateralis 194, 384 insecticides 506-509 honeydew Hyperaspis notata 143,194. insidious flower bug see Orius arrestant effects 120.222 491 insidiosus cardenolides in 162 Hyperaspis pantherina 194 instars 71-72 and old prey colony 22 7-228 Hyperaspis raynevali 143, 194 and body size 73. 74. 150 honeydew producers, ant attendance Hyperaspis reppensis 80. 378. 380 food consumption 159 236. 237. 359, 362. Hyperaspis senegalensis hottentotla foraging of first 233 379-382. 502-503 143, 194 survival when fed B. brassicae 166 honeysuckle see Lonicera Hyperaspis senegalensis 143, 194. tolerance to extreme temperatures peridymenum 389 96-97 hop powdery mildew see Sphaerotheca Hyperaspis sphaeridioides 126 integrated pest management (IPM) castagnei Hyperaspis undulata 132. 306 programmes 506 host plants Hyperomyzus carduellinus 63. 193 insecticides used in 507-508 age. effects of 122.228.294 Hyperomyzus lactucae 154, 190 integumentary system chemical cues/stimuli 120, hyperparasitoids 387-389 pheromone release 454 199-200. 224-226 Hyphantria cunea 178. 197 pupal defence 76 co-existence of several ladybird Hypocreales, fungal pathogens intercropping, ladybird diversity species 121-122 415-416 501-502 food of herbivorous Coccinellidae hypsotactic orientation/responses interguild effects 362-363 198-200 302, 303.305.310 inter-simple sequence repeat (ISSR) genetically modified 122-123 hypsotact ic species 310-311 analysis 35. 41-42 microclimate effect 124-125 interspecific mating 85 preference for type 120-121 Iceryapurchasi 165. 167, .1 72 intraguild interactions 344, 346 resistance 169.200.524-525 control of 491.509 ant-ladybird interactions structure 215-217 icterine warbler see Hippolais icterina 359-360 surface 123-124.217-218 llleis bielawskii 405 biological control 361-362 house martin see Delichon urbica llleiscincta 80,393.405 commensalism 345. 347 house sparrow see Passer domesticus llleis galbula 155. 183, 297, 324 competition 347-349 Howardula 414 diapause 297 conservation 360-361 548 Subject index intraguild interactions, (continued) Juglans cinerea 152 Uranicilliwn longisporum see interguild effects 362-363 juniperus virginiana 120 Lecanici/Jiimi lecnnii male-killing bacteria 349 juvenile hormone (JH) 327-328 Lcis dimidiate see Harmonia dimidiata mutualism 345-347 Leis see Harmonia niche partitioning 344-345 kairomones 451 Lemnia biplagiata see Coelophora intraguild predation (IGP) 172. 349 kidney-spot ladybird see Chilocorus biplagiata and biocontrol 361 renipustulatus lemon grass see Cymbopogon citratus and biocontrol limits 490-491 kin recognition, semiochemicals Lcontopodium alpinum 181 on coccinellids 355.358-359 448 Lepidaphycus see Homalotylus defensive mechanisms lepidopterans as prey 169-170 355-358 labial palp, chemoreception 223 Lepidosaphes beckii 168 coccinellids as predators lacewings. IGP 349. 352 Lepidosaphes cornutus 60 benefits to coccinellids 350 Laingia psammae 191 Lepidosaphes ulmi 167 of coccinellids 351-352 landscapes Leptinotarsa decemlineata 62, 144, general rules 349-350 diversity and character 125—127 152, 170, 192.202,205. hypotheses 350-351 landscape scale 211 326,490 on intraguild parasitoids 353 larger scale foraging 234-236 Leptothea galbula see llleis galbula on intraguild pathogens 354 Lapsana communis 180, 180 l.cptus ignotus 411 of non-coccinellids 352-353 larch adelgid see Adelges laricis lesser whitethroat see Sylvia curruca top predators 354-3 5 5 larch ladybird see Aphidecta obliterata Leucaena 217 definition of 349 Laricobius nigrinus 353, 358 life history characters 55 nitrogen shortage hypothesis 210 larva(e) 71-73 adult 76-91 vulnerability to 350-351.355 attachment ability 218 aphid prey effects 148 intrinsic rate of population increase body size 73 egg 55-70 89 consumption of 178-179 inheritance of 28—29 food and humidity affecting 91 defence 356-357 larva 71-73 temperature sensitivity 96 development 72-73 pupa 73-76 introductions of coccinellid species instars 71-72 life span see longevity 115, 362 mean SCP 96 life tables 471-472 aphidophagous 492-493 morphology 71 li fetime fecundity 5 8-67, 8 8 coccidophagous 491-492 parasitization of 391 and egg hatching rate 68, 69 competitive displacement response to semiochemicals higher for once-mated females 83 494-495 450-451 temperature effects 89 invasive species 493-494 larval development light traps, sampling 114 inundative biological control 362, aphid prey 158. 159. 163, 168 l.igustrum 183 498 food consumption/deprivation lime aphid see Eucallipterus tiliae invasive species 115, 360. 493-494 207-208 Lindorus lophanthae 36, 81, 94. 492 genetic diversity studies 40-41 larval tracks, oviposition deterrence l.inepithema humile 162. 237. 237 invertebrate predators 3 7 7-3 79 222.229-233. 356.454. Iioadalia flavomaculata 194. 393 lridomyrmex 381 456 Liosomaphis berberidis 58 lsaria farinosa 415 active substances in 455 Lipaphispseudobrassicae 59, 61. 68, Isaria fumosorosea 415—416 Lasius claviger 380 88,91. 147. 164, 168. Isodromus niger 386 Lasius japonicus 237 190-195.207 isothiocyanates 165-166. 225, 452 Lasius niger 85. 236, 380-381. lipids. diapause 323-325 isozymes analysis 34, 41 404, 449, 503 little black ant see Monomoriwn Italian rye grass see Lolium Lasius umbratus 380 minimum multiflorum leaf-curling plum aphid see local faunas 114 ITS-1 (first ribosomal internal Brachycaudus helichrysi climatic changes 115—116 transcribed spacer) region, learned response geographic differences 114—115 gene sequencing 31-33 colour perception 224,226 invasion and extinction 115 intensive search 220 location of communities, jack pine budworm see Choristoneura prey selection 452-453 determinants of 116 pinus leaves host plant 120-124 Japanese ant see Lasius japonicus and biological control 504 landscape 125-127 Japanese quail see Coturnix japonicus feeding on 184-185,197 microclimate 124-125 Jauravia quadrinotata 80 Lecanicillium lecanii 415 prey 116-120 Subject index 549

Lolium multiflorum 183 evolutionary rationale 423 mealy plum aphid see Hyalopterus Lolium perenne 123, 183, 183-184 molecular genetic studies 29-3] pruni longevity phylogenetic studies 38. 39 mealybug destroyer see Cryptolaemus food affecting 91 and sibling cannibalism 176 montrouzieri sexual activity affecting 83, 91 male size 78 mealybugs (Pseudococcidae) 169, temperature effects 88. 89. and female mating preference 222.237 90-91. 96 84-85 biological control of 496.503 voltinism 90 pupal stage 75 Medicago sativa alfalfa (or often Longiunguis donacis see Melanaphis Malus pumila 120 lucerne in UK) 197.501 donacis mandibles 241 Medina collaris 384-385 Lonicera periclymenum 120 Tytthaspis sedecimpunctata 183, Medina funebris 384-385 lower development threshold (LDT) 200 Medina luctuosa 386 92 Manihot esculenta 169, 169 Medina melania 384-385 for pre-imaginal development 92, Manikara zapota 496 Medina separata 384-385 93-94 mark-recapture method, sampling Mediterranean flour moth see Ephestia relationship with SET 92. 95 114, 522 kuehniella thermal window 95 marmalade hover fly see Episyrphus Megalocaria dilatata 57. 64. 80, 85, lupin aphid see Macrosiphum balteatus 194. 382 albifrons mating(s) Megaselia 385 Lupinus luteus 163 duration 82-83, 84 Megoura viciae 119, 145. 161. Lupinus mutabilis 163 frequency 82-83 162-163, 164. 168. 191. Luscinia luscinia 379 and fecundity 88 193 Luscinia svecica 379 and longevity 91 Melanaphis donacis 191 luteolin 7-0-glucoside 199-200 genetic studies 42-43 Melanaspis glomerate 60 Lydinolydella metallica 385, 387 hybridization 85 melanic (dark) morphs Lygocerus see Dendrocerus male selection 83-85 environmental conditions Lygus 475, 477. 504 inbreeding studies 40 23-27 Lygus hesperus 477 and melanism 83-85 inheritance 19-23 Lygus lineolaris 475 advantage for melanic morphs mating advantage 26-27,42 Lypha slossonae see Chrysotachina 26-27 melanism slossonae female choice 83-85 and mate preference 83-85 LysiphlebusJabarum 171. 353 multiple 68 melanization extent, pupae 76 Lysiphlebus testaceipes 173, 353. preference 84 winter survival, effect on 504 prior to dispersal 309. 322 331-332 refractory periods 77 Melanocorypha calandra 379 Maconellicoccus hirsutus 192. 496 refusal to mate 83 Melasoma populi see Chrysomela populi Macroilleis hauseri 194 semiochemicals implicated in melegueta pepper see Aframomum Macronaemia hauseri 393 antennae chemoreceptors melegueta Macrosiphoniella artemisiae 58, 191, 453 Melolontha sp. 302 194 glands, pheromone release melon aphid see Aphis gossypii Macrosiphoniella sanborni 191 454 Mendozaniella see Homalotylus Macrosiphum albifrons 163 hydrocarbons on elytra 454 Menochilus quadriplagiatus see toxicity 163 sex pheromones 453 Menochilus sexmaculatus Macrosiphum euphorbiae 118, 174. sperm competition 83 Menochilus sexmaculatus 64. 68. 70, 205. 224. 468 willingness 82 75, 78, 80. 82, 83. 86. 90. Macrosiphum ibarae see Sitobion ibarae Malsucoccus feytaudi 222 91. 117. 118, 123. 146. 147. Macrosiphum rosae 153, 188, 193, Matsucoccus josephi 189 163. 174. 176. 178. 179. 497 Matsucoccus matsumurae 222, 451 183. 186,210,215.228, maize (corn in USA) see Zen mays maxillary palps 229.231, 356.389.393. malaise traps 113-114 perception of larval chemical 405. 405. 405. 410. 414. male gonads, diapause 321—323 marker 222, 223. 226 421.423,424,425.446, male-killing bacteria 349.421 prey contact and recognition 455. 507. 508 diversity of 421.422-423 223 essential foods 194 evolutionary dynamics 423-424 meadow pipit see Anthus pratensis and first chemically defined ODS evolutionary implications meadow spittlebug see Philaenus 231,232 424-425 spumarius oviposition 228-229 550 Subject index

Mercurialis annua 183 Microweiseinae basal subfamily 4, 5 morello cherry or amarelle cherry see Merismoclea rojasi 385. 405 Microweiseini tribe 5 Primus cerasus Mermis 414.415 middle spotted woodpecker see morphology Mermithidae 415 Dendrocopos medius egg 55-56 meroistic telotrophic ovarioles 78 migration 302-303 inheritance Mesopolobus secundus 406 Mindarus abietinus 189. 471 colour patterns 19-27 Mesopolobus 385-386.406 Misumenops tricuspidatus 358, 361 wing polymorphism 27—28 Mestocharis lividus see Pediobius mites larva 71 foveolatus Coccipolipus 411-414 pupa 74-75 metabolic changes related to diapause commensalism 345 mortality glycogen 325-326 intraguild predators 353 after consumption of sinigrin 166 lipids 323-325 phoretic 411 eggs used as diet 170. 173 metabolic rate 3 2 6-3 2 7 mitochondrial DNA (mtDNA) embryonic 85 water 326 29-31. 36, 37 larvae fed on aphids 143, 145 metallic blue lady beetle see Curinus mixed and combined diets 155-157 risk in newly hatched first instars coeruleus complementation across life stages 233 Metarhizium anisopliae 415. 498 156-157 stiige specific 72 Metastenus caliginosus 406 prey specialization through temperature extremes 96-97 Metastenus concirmus 383. 405-406 selection 157 toxic and rejected prey 158-167 Metastenus indicus 402. 406 prey switching 157 unsuitable host plants 200 Metastenus species (parasitoids) mixed plant stands, arresting effect Morns australis 201 405-406 458 'mosaic dominance' hypothesis 23 Metastenus sulcatus 405 modifier genes, colour pattern Motadlla flava 379 Metastenus townsendi 384. 405-406 variation 22-23 moulting of larva 70. 71, 72 Metastenus 405 molecular genetic markers 29 mullein bug see Campylomma verbasci Metatetranychus ulmi see Panonychus isozymes for delimiting cryptic Mullerian mimicry 27 ulmi species 41 Mulsantina hudsohica 306 methionine 155 [SSR, sperm competition 41-42 Mulsantina picta 122, 130, 132, methylalkylpyrazines. reflex blood mitochondrial DNA (mtDNA) 344. 423 377 29-31 multiple allele effect 22 methyl linolenate 452 species-specific 33-37 multi-species aggregations/ methyl salicylate 225. 448. 450 studies using 34—35 combinations 308 methyl tricosane 448. 454 molecular genetic studies 4-5. 29 aphid suppression 472-473 Metopolophium dirhodum 147. 156. future research in 521-522 multivoltinism 276 159. 191, 195 gut-content analyses 144-145. A. bipunctata 290.314 Metopolophium festucae 475, 475 358.478-481. 524 C. septempunctata 281. 283, 284. mexican bean beetle see Epilachna paternity and sperm competition 286. 320 varivestis 42-43 Har. axyridis 295 Micraspis allardi 80 phylogenetic studies 37-38 Hip. convergens 292 Micraspis discolor 64. 68. 88. 147, population genetic & potential multivoltinism 286-287 194.393, 507 phylogeographic studies mummies, feeding on 171-172,173 microarray chips, species 38-42 Must icapa striata 379 identification 36 sequence evolution 29-33 mussel scale see Lepidosaphes ulmi microclimate 124-125 species identification 33-37 musiard aphid see Lipaphis Microctonus 394 Momordica charantia 410 pseudobrassicae microhabitats 233 monarch butterfly see Danaus mutual interference 207 Microlophium carnosum 58. 145, plexippus mutualism 345-347. 350, 351 150. 153, 153. 188. Monocorynini tribe 9 ant attendance 236-238. 190-191,213.238 Monomorium minimum 237 379-382 micropyles. egg 55. 55 Monophlebulus pilosior 195, 242 ants and honeydew producers microsatelliteanalysis 35. 36,40.43 monophyly 359. 362 Microsphaera alphitoides 184, 200 Coccinellidae family 2—3 lacewings 352 Microsphaera pulchra 201 other subfamilies 4 mycophagous Coccinellidae microsporidia 418-419 of six subfamilies 3 food of 200-201 Microweisea sp. 384 rejection of 5 hibernation 316 Subject index 551

Myiopharus doryphorae 385 Neomyzus circumflexus 145. 188, Nosema tracheophila 419 myrmccophilous ladybirds 360. ' 238 Nosema varivestis 418 380-382 neonicotinoids. insecticides Nosematidae (fungal pathogens) myrmecophily 449 506-509 418-419 Myrmica ruginodis 236 Neophyllaphis podocarpi 191. 193 Nothoserphus admirabilis 405 Myrmica rugulosa 381 Neotainania see Uga Nothoserphus aequalis 405 Myrrha octodecimguttata 80. 122. Neotyphodium lolii 123 Nothoserphus afissae 404. 405 128. 149. 151, 194. 304. Nephaspis oculatus see Clitostethus Nothoserphus hoops 404, 405 308. 393.399.419.420, oculatus Nothoserphus debilis 405 421,449 Nephus bilucernarius 194. 503 Nothoserphus epilachnae 418 dormancy, responses 307 Nephus bisignatus 89. 94. 96 Nothoserphus fuscipes 405 hibernation sites 314 Nephus flavifrons 132 Nothoserphus mirabilis 404-405 Myzia oblongoguttata 80, 129. 129. Nephus guttulatus 384 Nothoserphus partitus 405 131. 151, 151, 314.389, Nephus includens 65. 89. 94, Nothoserphus scymni 384. 405 393. 399,408 194 Nothoserphus species (parasitoid) Myzia subvittata 132 Nephus kiesenwetteri 389 404. 405, 405 Myzocallis boerneri 153. 188-189 Nephus ornatus 389 Nothoserphus townesi 405 Myzocallis carpini 188 Nephus quadrimaculatus 314. 385, 'no-till' agriculture 502, 503 Myzocallis castanicola 153. 188 474 Noviini tribe 8 Myzocallis coryli 153. 188-189 Nephus redtenbacheri 80 N-oxides, Aphis jacobaeae 163-164 Myius cerasi 147. 153. 153. 188, Nephus soudanensis 389 n-pentacosane 448. 455. 456 191 Nerium oleander 161-162,446 n-tricosane 456 Myzus persicae 58, 61-65, 68, 76, neuropterans. IGP 352 numerical response 88. 466. 471 78,87.91, 117. 120. 143. n-heptacosane 456 aggregative 204,227. 145. 148. 156, 159, 162- ni moth see Trichoplusia ni 347-348 163, 165, 165-166. 166. niche differentiation 128 additional prey, importance of 168. 768. 170-171. 179. niche overlap 229,231 212-213 188-194.202,205. 207, niche partitioning 344-345. 472 modelling of 211 210. 222.224.236.238, Nilaparvata lugens 195 temporal and spatial patterns 293.450-451,467-490. nine-spotted lady beetle see Coccinella 211 498 novemnotata vegetation influence 211-213 Myzus persicae nicotianae 120. 191. nipplewort see Ixipsana communis nutrition 202.450 Nitraria 200 food suitability 145-147 Nobrimus see Homalotylus 400 food unsuitability 164 Nabis (Reduviolus) americoferus 472 non-fertile/non-hatching eggs 68, intraguild predation 210 natality 86. 87 69 non-abundant nutrient concept native ladybirds, competitive non-melanic (light) morphs 19 154-155 displacement of 115.171. balanced selection 26 nutrients in egg yolk 55 493.494-495 female preference 2 7 plants providing 196-197 natural enemies 3 76-428 paternity and sperm competition nutritional IGP hypothesis 351 impact on populations 425-428 studies 42 nutritional induction of diapause parasites and pathogens 411-425 non-myrmecophilous coccinellid 292.293-294 parasitoids 383-411 species, ant interaction Nysius huttoni 476 predation 3 76-382 359-360 nectar, feeding on 180. 183, 215. non-sibling cannibalism 175 obligatory diapause 2 76 499-500 egg consumption 177-178 Oenanthe oenanthe 379 Neda marginalis 407 kin recognition 179 Oenopia billieti 80 nematodes 414 larvae consumption 1 78-179 Oenopia conglohata 23, 65, 80, 131, Allantonematidae 414 population consequences 179 154. 195, 308. 314. 330, Mermithidae 415 northern wheatear see Oenanthe 378. 393,412,413. 414. Neoaenasioidea see Homalotylus oenanthe 420 Neoaulacortluun magnoliae 160, 260, Nosema coccinellae 418-419 Oenopia kirbyi 80 196 Nosema epiladmae 418 Oenopia lyncea 65 Neoaulacorthum nipponicum 241 Nosema henosepilachnae 418 Oenopia sexareata 81 Neocalvia anastomozans 407 Nosema hippodamiae 419 Oidium monilioides 183. 200 552 Subject index oil seed rape see Brassica napus subsp. orange ladybird see Halyzia oxygen consumption, diapause oleifera sededmguttata 326-327 oleander aphid see Aphis nerii Orcus australasiae 205. 384 oystershell scale see Lepidosaphes ulmi oleander aphids, toxicity 162-163 Orcus chalybeus see Halmus chalybeus oleander see Nerium oleander Oricoruna 385 Pachyneuron albutius 389 olfaction 224-226 Oricoruna orientalis 385 Pachyneuron altiscuta 389 early studies 220-222 Orius insidiosus 118. 358. 480 Pachyneuron chilocori 389 host plant odours 120 Ortaliinae subfamily 5. 8, 8-9 Pachyneuron concolor see Pachyneuron olfactory cues, larval response to Ortaliini tribe 8. 9 muscarum 451^152 Orthezia urticae 194 Pachyneuron muscarum 389 perception distance 222 Oryzopsis hymenoides 206 Pachyneuron siculum see prey odour detection 119,120 Ostrinia nubilalis 152. 207. 358. Pachyneuron solitarium 389 receptors in aphidophagous species 480. 490 Pachyneuron syrphi see Pachyneuron 223 Ouchterlony plate approach, antibody albutius olfactometers 222. 223. 224, 225 analysis 476 Pachyneuron 389 experiments 120. 450, 453, 458 ovarian development 82. Puccilomyces farinosus see Isaria olive scale see Saissetia oleae 317-321 farinosa Olla abdominalis see Olla v-nigrum ovarioles 78-82 Paccilomyces fumosoroseus see Isaria 011a v-nigrum 15, 20. 22. 65, 90, and cluster size 57. 82 fumosorosea 94,96, 130, 132, 143, 149, and egg weight 56 Paederia foetida 241 163. 172, 173, 174-175. and female weight & size 78. pair formation 84 179. 185, 195.235, 361, 79-81. 87 Palaeoneda auriculata 81 391,393,407,408,416. overwintering survival 308, Palconeda miniata see Palaeoneda 417, 507-508 328-331 auriculata Omphale epilachni see Chrysonotomyia oviposition 85-88. 119 Pandora neoaphidis 354 appannai alternative foods, effects on 1 71 Pania luteopustulaia 81 Omphale 386 animal protein, importance of Panonychus mori 67 onion thrips or tobacco thrips see 156 Panonychus ulmi 152.167 Thrips tabaci deterrence 228-233.454-456 Parachrysocharis 386 Onodea sensibilis 313 diurnal periodicity 218.229 Paradexodes epilachnae see oocytes 57.82.317-320 effect of food restriction 209 Euthelyconychia epilachnae Ooencyrtus azul 384 essential foods, effect on 746, Puranaemia vittigera 195 Ooencyrtus bedfordi 384 187. 196 parasitic mites, Coccipolipus Ooencyrtus camerounensis 384, 426 period 86, 88 411-414 Ooencyrtus distatus 387 place 85-86 Parasitilenchus coccinellinae 414 Ooencyrtus epilachnae 384 « and prey stage 156, 157 parasitization 390-391 Ooencyrtus epulus see Ooencyrtus rate 86, 87 population impact of 426-428 camerounensis peak of 86 parasitized aphids, inferior prey Ooencyrtus polyphagus 387 rhythmicity 86 171-172. 173. 353 Ooencyrtus puparwn 3 84 sites, search for 226-228 parasitoids 88, 383 Ooencyrtus sinis 384 stimulated by host plant odours iint behaviour towards 237.360 Oomyzus 409 120 avoidance of coccinellid tracks Oomyzus mashhoodi 384 substitute diets 185-186 232 Oomyzus scaposus 384,388-389. substrates 85-86 general characteristics 390-391 391,408-409,428 termination of 117 intraguild Oomyzus sempronius 386 toxic aphid, negative effect on ants attacking 360 oosorption 82, 317 162 coccinellids preying on 353 food restriction, effect of 209 trophic egg laying 177 predation on coccinellids by Ophelosia bifasciata 385 oviposition deterrence pheromones/ 358 Ophelosia crawfordi 388 semiochemicals 117, 229, primary 383-386 opportunistic IGP hypothesis 351 231. 356, 358.454 review of important parasitoids optimum temperature active substances in larval tracks 391-411 development rate 92,97 455 Cowperia 399.401.401 hatching rate 68 aphid abundance 456 Dinocampus coccinellae 391-399 thermal window 95 species-specificity 454-455 Homalotylus 400-404 Subject index 553 review of important parasitoids. Pentilia insidiosa 384 Phorodon humuli 58, 188. 191 (continued) Pentiliini tribe 10 photoperiodic activation 277. 281, Metastenus 405-406 percentage fertility 68-69 282 Nothoserphus 404-405 Perilitus americanus see Dinocampus precocious 295 Oomyzus scaposus 408-409 coccinellae photoperiodic response 276. 283, Pediobius foveolatus 409-411 Perilitus coccinellae see Dinocampus 288-289. 295, 300 Phalacrotophora 406-408 coccinellae photoperiods Uga species 399.400 Perilitus rutilus 394 see also diapause/dormancy secondary 387-389 Perilitus stuardoi 387,391 critical 276.284,290.291.298, semiochemical attraction 449 Perilitus lerminatus see Dinocampus 299 Parastethorus nigripes 195 coccinellae egg cluster size 5 7 Paratrechina 381 periodicity see rhythmicity hatching rate 68 parenthesis lady beetle see Hippodamia Periphyllus californiensis 193 longevity effects 90-91 parenthesis Periphyllus lyropictus 188 Phthorimaea operculella 59 Parexochomus melanocephalus 313 Periphyllus testudinaceus 153 Phygadeuon subfuscus 388 Parexochomus nigromaculatus 89, 94, pest control Phyllactinia moricola 201 167 augmentation 495-498 Phyllaphis fagi 153, 188 Parexochomus troberti 393, 412. 471 pesticides, selective use 506-509 Phylloscopus collybita 379 Parexochomus troberti concavus 393, phagostimulants 452 Phylloscopus trochilus 379 412 Phalacrotophora berolinensis Phylloxera glabra 146, 195 Parlatoria blanchardi 60, 195 384-385,406 phylogenetic studies 3 7-38 parsnip see Pastinaca saliva Phalacrotophora beuki 384-385. 406 phylogeny 5-10 Parus major 376 Phalacrotophora decimaculata phylogeographic studies 38-42 Passer domesticus 377 384-385. 406 Phymatosternus lewisii 360. 381 Passer montanus 377,379 Phalacrotophora delageae 3 84-3 8 5, Physalis alkekengl 199 Pastinaca sativa 126 406 phytophagous Coccinellidae paternity studies 42-43 Phalacrotophora 75, 384, 391, 406, food of 198-200 pathogens 407, 408. 426, 428 hibernation 316 bacteria 421-425 Phalacrotophora fasciata 391 population impact of natural fungal 415-419 Phalacrotophora Indiana 3 84—3 8 5, enemies 426-428 intraguild 406 Picea schrenkiana 314 coccinellids preying on 354 Phalacrotophora nedae 384—385. Pieris rapae 170 commensalism 345 406 pine ladybird see Exochomus predation on coccinellids by Phalacrotophora philaxyridis quadripustulatus 359 384-385.406 Pineus pini 188 nematodes 414—415 Phalacrotophora quadrimaculata pink bollworm see Pectinophora protozoan 419-421 384-385. 406 gossypiella PCR see polymerase chain reaction Pharoscymnus anchorago 420. pink hibiscus mealybug see 29, 145 421 Maconellicoccus hirsutus pea aphid see Acyrthosiphon pisum Pharoscymnus numidicus 388 Pinnaspis buxi 190 peach or nectarine see Prunus persica Pharoscymnus ovoideus 388 Pinus armandii 132 peach-potato aphid see Myzus persicae Phascolus vulgaris 126 Pinus sylvestris 153.449 Pectinophom gossypiella 63.477 Pheidole megacephala 362. 503 Pisonia 496 Pediobius amaurocoelus 388 Phenacoccus herreni 194 Pisum sativum 124.724 Pediobius epilachnae see Pediobius Phenacoccus madeirensis 192 Pittosporum tobira 227.228 foveolatus Phenacoccus manihoti 169. 194. Plagiodera versicolora 197.197 Pediobius foveolatus 384. 388-389. 222,404,471 Planococcus citri 65, 167. 192. 222. 391,409-411.418.426 phenotypic plasticity 522 496 Pediobius mediopunctata see Pediobius pheromones 117.222,224.225, Planococcus minor 59,189 foveolatus 239. 240. 241, 454 plant antibiosis 504-505 Pediobius nishidai 384 Philaenus spwnarius 476 plants Pediobius simiolus see Pediobius Phoenicococcus marlatti 194 see also crops: host plants foveolatus Phoeuicurus ochruros 379 spatial guild partition 344 Pediobius 409 Phoenicurus phoenicurus 379 plant stress 457.458 pentatomids, IGP 358 phoretic mites 345,411 plant structure 215-217.504 554 Subject index plant surfaces 123-124.217-218 Poriini tribe 9 alternative 146. 147. 150, 2 77. plant volatiles post-mortem analysis of predation 180. 187, 214. 242 avoidance of 458 474 an I-attended 360 conservation 499 antibody-based 474-477 anti-predator defences 117-118 herbivore-induced 449-450 prey-specific DNA detection essential 746, 158. 162, 772. larval response 450-451 478-481 188-195 link to patch preference 457 protein marking 477-478 preference 118-119 receptors for food-related 450 post-oviposition period 86 quantifying coccinellid impact related to aphid density 456 potato aphid see Macrosiphum 466-467 Platynaspidini tribe 8. 9. 10 euphorbiae assays of consumption Platynaspis luteorubra 55,81, 131, potato ladybird see Epilachna dregei 467-468 237. 241, 381,404 potato see Solanum tuberosum complex communities Platynus dorsalis 404 potato tuber moth see Phthorimaea 472-474 Pleurotropis see Pediobius operculella indirect impacts 468 plum powdery mildew see Podosphaera powdered food, preference for 152 post-mortem analysis 474-481 tridactyla powdery mildew see Erysiphe polygoni statistical approaches 471-472 Plutella xylostella 124. 170. 504 Praon volucre 455-456 traditional approaches Pnigalio agraules 386 precipitin test, predation analysis 468-471 Podisus maculiventris 145. 354. 354. 474, 475 prey abundance see aphid abundance/ 532 precoccinelline 446.449 density Podosphaera leucotricha 194. 200 predation 376-382 prey alarm pheromones 451 Podosphaera tridactyla 201 see also intraguild predation (IGP) prey capture 238-241 poisoning by toxic aphids anti -predator defences 3 76-377 prey choice niches 128 157-165 chemical protection against 56. prey contact Policheta unicolor 385 76 approach direction 239 polished lady beetle see Cycloneda hemiptera-tending ants 379-382 intensive search behaviour 220. munda post-mortem analysis 474 221 pollen, supplementary/alternative antibody-based 474-477 by maxillary palps 223 food 123,180-184 prey-specific DNA detection prey density pollinivory 152.180-182.185, 478-481 aggregative numerical response 200,282 protein marking 477-478 211 pollution and melanic morphs 24. social aphids with a soldier caste aphid infestation 227-228 25. 26 382 effect on fecundity 88 polyandry 93 studies quantifying 524 functional response, consumption polyazamacrolide alkaloids 456 traditional study of impact 204-207 pupal defence 76 468-469 and ladybird vagility 234-236 polyclonal antibodies, trophic cage inclusion 470-471 minimum threshold 117 relationships 474-477 field cages 469-470 and oviposition activity 456 polymerase chain reaction (PCR) 29. manual removal 471 prey size-density hypothesis 145 selective exclusion 469 147-150 gut-content analysis 478-481 predator facilitation 345 risk to larvae of low 233 species identification 35,36 and biocontrol 361-362 prey discovery, quantifying 471 polyphagous (euryphagous) species see predators of coccinellids prey handling, associative learning generalist species 1 50-1 54 ants 379 452-453 polyvoltine (multivoltine) species 90 invertebrate 377 prey-induced plant chemicals, poplar leaf beetle see Chrysomela vertebrate 377 attraction to 120. 218-226 populi pre-diapause phase 301-302 prey preference 118—119 population density preferential mating 83-85 prey quality and fecundity 88 pre-oviposition period 77-78 beneficial effects of 119 and larval development 73 prepupal stage 73-74 iind larval development 207-208 population genetic studies 38-42 prey 116 problematic'prey 167-169 populations of ladybirds, impact of sec also food, see also aphids rejected prey 746.165-168 natural enemies 425-428 age. effect on oviposition toxic prey 746.157-165,452 Populus 313 227-228 prey selection 153. 207 Subject index 555 prey sex pheromones 451 Pseudococcus cryplus 503 quantitative aspects, food relations prey size-density hypothesis Pseudococcus maritimus 237 201 147-151 Pseudococcus viburni 23 7 aggregative numerical response prey specialization through selection Pseudoregma alexanderi 196 211-213 157 Pseudoregma bambucicola 194. 196, food consumption prey specificity 489-490,523 382 consumed food, conversion and prey substitutes 169-175 Pseudoscymnus kurohime see utilization of 209-210 prey suitability 144,145-147. Sasajiscymnus kurohime effect of physical factors 173 Pseudoscymnus tsugae see 202-204 prey switching 157. 204. 206 Sasajiscymnus tsugae effect of prey density: functional prey toxicity 157-165,451-452 Psylla alni 59 response 204-207 primary parasitoids 383-386,390 Psylla jucunda 1 54, 193 effects on growth and Priscibrumus lituratus 393 Psylla mali 59,154,189 reproduction 207-209 Priscibrumus uropygialis 81, 393 Psylla ulmi 59 Quercus rubra 152 Pristomyrmex pungens 380. 419 Psylla uncatoides 195 quiescence 276.277,295 'problematic'prey 167-169 psyllids 174 quinolizidiue alkaloids 163 Prochiloneurus aegyptiacus 387 Psyllobora confluens 66. 9.1 Prochiloneurus nigriflagellum 384 Psyllobora vigintiduopimctata 25,81. radioimmunoassay. vitellogenin prolegs, prey detection 223 131, 201. 312, 316.393. synthesis 328 promiscuity 82 399. 417 radish see Raphanus sativus Propylea dissecta 65, 70, 70. 72. 94, Psyllobora vigintimaculata 132, 200, ragwort see Senecio jacobaea 148, 195.205.291 306,417.418 Random Amplification of Polymorphic Propylea japonica 22. 39, 65-66, Pterocallis alni 153,188 DKA (RAPD) 33, 34. 36 144, 158. 174. 195, 228, Pterostichus melanarius 479 Raphanus sativus 121 232,393.423 Pterostyrax liispidus 198 Rastrococcus invadens 496 Propylea quatuordecimpunctata 151, Puccinia 180. 183 Ravinia errabunda 385 180, 184. 195,355 Pullus auritus see Scymnus auritus red cedar see juniperus virginiana diapause regulation 291 Pullus mediterraneus see Scymnus red imported fire ant see Solenopsis dispersal from dormancy site marinus invicta 309 Pullus subvillosus see Scymnus red oak see Quercus rubra hibernation behaviour 313.321 subvillosus red spider mite see Panonychus ulmi movement on plant surfaces Pulvinaria psidii 192, 496 red-barbed ant see Formica ruflbarbis 217 Pulvinaria urbicola 496 red-capped lark see Calandrella cinerea seasonal distribution of flights Pulvinaria vitis 152 red-headed flycatcher see Ficedula 306 pupa(e) parva protandry 77 colouration 76 redistribution of ladybirds, biological protection defence 76. 357 control 496-497 eggs 56. 85 mean SCP 96 reflex bleeding 376-377,445 pupae 74. 76 morphology 74-75 chemical substances 445-449 protective IGP hypothesis 350-351 prepupal stage 73-74 enemies and competitors 449 protein marking, post-mortem protection, chemical defence 456 molecular genetic studies 29,42 predation analysis 477-478 thermal melanism 76 refractory period, mating 77 protein supplements 187 timing of pupation 75 refuges protogyny 77 pupation place 75-76 hibernal ion 500 protozoan pathogens 419-421 Pyracantha coccinea 201 landscape-scale 127 Primus avium 120 pyrazines 377.455 regression models 471 Prumis cerasus 153 Pyrrhalta luteola see Xanthogaleruca rejected prey 165-167 Primus persica 226 luteola relative growth rate (RGR) 73 Pseudebenia epilachnae 384 Pyrrhocoris apterus 327 release of coccinellids, augmentation Pseudoazya trinitatis 236, 384 pyrrolizidine alkaloids 163-164. 495-498 Pseudocatolaccus 386 165,445, 446 reproduction Pseudochermes fraxini 192 and food consumption 208-209 Pseudococcidae (mealybugs) 169. Quadrastichus ovulorum 385 interaction of larval and adult diet 222.237 quail see Coturnix coturnix 156-157 556 Subject index reproduction, (continued) Rodolia fumida 385 scotophase interspecific 85 Rodolia guerini 81 life events 70 and longevity 91 Rodolia iceryae 143. 386 oviposition 86. 299 net reproduction rate 89, 96 Rodolia occidentalis 385 pupation 75 temperature, effects on 96 Rosa multiflora 201 Scots pine see Pinus sylvestris reproductive isolation rose aphid see Macrosiphum rosae SCP see supercooling point cytogenetic changes 16,18 rose-grain aphid see Metopolophium Scymnillini tribe 10 hybridization 85 dirhodum Scymninae subfamily 5. 9-10 reproductive output 90 rose powdery mildew see Sphaerotheca Scymnini tribe 9,10 larval diet linked to 78 pannosa Scymnodes lividigaster see Apolinus reproductive rate 86,87 rosy apple aphid see Dysaphis lividigaster net reproductive rate 89,91. plantaginea Scymnophagus see Metastenus 96 rosy leaf-curling aphid see Dysaphis Scymnophagus mesnili see Metastenus reproductive success 42-43 devecta concinnus research, future trends 521-525 Rubus occidentalis 313 Scymnus (Nephus) quadrimaculatus see resilience, univoltinism 300 Russian wheat aphid see Diuraphis Nephus quadrimaculatus resistant host plants 91,164, noxia Scymnus abietis 72.131 200 rye grass see Lolium perenne Scymnus aeneipennis see 'lagloba and biocontrol 504—506 aeneipennis DIMBOA in wheat 164, 169 Saccharomyces fragilis 120 Scymnus apetzi 131. 383, 401. effect of prey feeding on 169 sagebrush see Artemisia tridentata 420 GM crops 122-123 Saissetia coffeae 190 Scymnus ater 81 respiration rate 75. 287, 304. 326, Saissetia oleae 168. 192, 472 Scymnus auritus 128, 131. 146. 327 Salix 153 195 Restriction Fragment Length Sambucus nigra 153, 158, 160 Scymnus cocdvora 195 Polymorphism (RFLP) analysis Sambucus racemosa subsp. sieboldiana Scymnus creperus 205 33,34.36 160 Scymnus dorcatomoides 405 Rhopalosiphum maidis 148. 1 54, sameness Scymnus flavifrons Blackburn 168. 188,212, 212, rule of 154-155 (Australia) 132 225-226 sampling methods 111-114.469. Scymnus flavifronsMelsheime r (North Rhopalosiphum path 188 471. 522 America) see Nephus flavifrons rhythmicity non-destructive 29 Scymnus frontalis 66. 77, 81, 128. eclosion 70 quadrat sampling 112 129, 131. 195,330,330. hatching 69, 70 sapodilla see Manikara zapota 378, 505 mating 70, 82 Sarcophaga helicis see Helicobia rapax Scymnus guttulatus see Nephus moulting 70 Sarcophaga latisterna see Boettcheria guttulatus oviposition 70.86,27 9 latisterna Scymnus haemorrhoidalis 81. 129 pupation 70, 75 Sarcophaga reinhardii see Ravinia Scymnus hilaris 174 Rhyzobius litura 155,183,195. errabunda Scymnus hoffmanni 66, 85 313. 387.423,424 Sasajiscymnus kurohime 241-242, Scymnus impexus 383 Rhyzobius lophanthae see Lindorus 382 Scymnus interruptus 66, 81, 131, lophanthae Sasajiscymnus ningshanensis see 236. 380 Rhyzobius ventralis 195, 241 Scymnus ningshanensis Scymnus kiesenwetteri see Nephus ribosomal DNA (rDNA) gene Sasajiscymnus tsugae 66, 77. 94. kiesenwetteri association with sex chromosomes 195, 358.471.493 Scymnus lacustris 122,344 15 Saxicola rubicola 379 Scymnus levaillanti 66, 71. 88. 89. ITS1 region 32 scale insects, biocontrol 491-492, 94, 195. 210 Rickettsia 30-31.30.38,39,68, 496 Scymnus loewii 130 176,421,422-423,424. scarce seven spot ladybird see Scymnus louisianae 55-56,66. 424, 534 Coccinella magnifica 85-86, 87 Robinia pseudoacacia 160 Schizaphis graminum 157, 497. 501, Scymnus marginicollis 66, 121 Rodatus major 195. 242, 381 504 Scymnus marinus 66, 94, 97. 195 Rodolia cardinalis 94. 154. 164. 167. Schizolachnus pineti 189 Scymnus mediterraneus see Scymnus 172,466,491, 497, 508. Schizolachnus piniradiatae 122, 167 marinus 509, 535 Schizopepon bryoniaefolius 198 Scymnus morelleti 387 Subject index 557

Scymnus nigrinus 81, 122. 314, hibernation and aggregation 457 Siphoninus phdlyreae 60, 190 380.405 mating and sex pheromones Sitobion akebiae 78 Scymnus ningshanensis 471, 493 453-454 Sitobion avenue 61. 77, 88. 116 Scymnus ornatus see Nephus ornatus olfactory response to 224—226 Sitobion ibarae 193. 222, 451 Scymnus otohime 384 oviposition 454-456 Sitona discoitleus 393 Scymnusposticalis 196, 228. 228, oviposition-deterring 229, 231 Sitotroga cerealella 63,169 360 Senecio jacobaen 165,446 Sitta europaea 153 Scymnuspyrocheilus 228. 229. sensitive stage, pupal melanization sixteen spot ladybird see Tytlhaspis 356 76 sedecii n punctata Scymnus quadrillion 389 sensory orientation/perception size Scymnus rubromaculatus 81. 131 218-220. 226 see also body size Scymnus sinuanodulus 55. 55, 66, earlier observations 220-222 egg 56 71. 74, 76, 90. 94, 196 effect of encounter 220 egg clusters 56-68 diapause 300 honeydew, arrestant 222 larval instar 73 Scymnus smithianus 399, 401 olfaction 224-226 sky lark see Alauda arvensis Scymnus soudanensis 389 sensory receptors 223 small cabbage white butterfly see Scymnus subvillosus 66, 88, 89. 89, vision 223-224 Pieris rapae 91,94.96. 131, 159, 196. septate eugrega rines 419-421 smoke pollution and melanism 26 241 sequence evolution 29-33 social aphids. soldier caste 382 Scymnus suturalis 81, 129, 308, ITS-1 region 31-33 soft brown scale see Coccus hesperidum 314 mitochondrial DNA (mtDNA) soft green scale see Coccus viridis Scymnus syriacus 66. 94 29-31 Solatium japoncnse 198 Scymnus tardus 417 Serangiini tribe 5 Solatium nigruin 198. 7 99 searching (foraging) behaviour Serangium parceselosum 66. 72. 91, Solatium tubeiosum 198, 452 213-236 155. 196.353,416 Solenopsis invicta 236,502 intensive 211.214.220 serological assays 144,145 Solidago canatltnsis 126 seasonal changes predator-prey research 474-477 song thrush see Turdus philomelos in frequency of colour morphs 25. seven spot ladybird see Coccinella Sospita vigintiguttata 407 83-84 septempunctata southern house mosquito see Culex and thermal properties of morphs sex determination 14-15 quinqiwfasciiitus 26-27 sex differences, antennal receptors sowthistle aphid see Hyperomyzus and habitat switching 127 453 lactucae seasonal cycles of crop colonization by sex pheromones soybean aphid see Aphis glycines coccinellids 503-504 prey attractant 222,224 Spanish broom see Spartium junceum second hibernation 90, 321 of prey, response to 451 spartein 16 5. 167 sedge warbler see Acrocephalus promoting coccinellid mating Spartium junceum 167 schoenobaenus 453 spatial guild partition 344 selection sex ratio 7 7 spatial matching, predator-prey directional 28 sexual activity 211 selective exclusion, predation impact longevity, effect on 91 spatial niches 128 measure 469 of melanic forms 84 specialist species 142-143, self-selection model, optimal diets sexual maturation 76-77 150-152, 154 155 sexual selection 27 non-abundant nutrient concept Selvadiini tribe 9. 10 shifting habitat hypothesis 360 154-155 Semiadalia undecimnotata see Shirozuellini tribe 6 role in biological control 490 Ceratomegilla undecimnotata shrubs, sampling from 113 speciation 16-18 semiochemicals 214. 445 sibling egg cannibalism 175-177. species identification aggregation 307 233 molecular genetic studies 3 3-3 7 aposematism and reflex bleeding Sidis see Nephus pupal colouration providing 76 445-449 Sidnia kinbergi 476 species specific chemistry egg and pupa protection 456 Sigmoepilaclma indica 385-386 oviposition deterrence substances food-related 449—£53 Sinapisalba 58. 123, 165, 765-266, 454-45 5 future challenges 459-460 225.452 reflex bleeding substances future research 523-524 Singhikaliini tribe 6 448-449 habitat preferences 4 5 7-4 5 8 sinigrin 451-452 spermatheca 296. 328, 320 558 Subject index spermatophores 425 Stenotarsus rotundus (Endomychidae) substitute diets 185-187 nuptial gift 83 276 substitute prey 169-185 protein source 1 73-174 Stethorini tribe 9-10 substrates sperm competition 42^3.83 Stethorus bifidus 66. 196, 206. 206 foraging (host plant) 206-207 Sphaerotheca castagnei 184. 200 Stethorus gilvifrons 67. 90. 96, 167. oviposition 8 5-86. 119-120 Sphaerotheca cucurbitae 200, 201 196, 498 pupation 75-76 Sphaerotheca pannosa 200,202 Stethorus japonicus 67. 94, 187. sucrose, alternative diet 169-1 70. spider-mite destro3'er see Stethorus 196. 300 185,222 punctum diapause 300 sugar beet and garden beet see Beta spiders 354, 358. 377-378, 449 Stethorus loxtoni see Parastethorus vulgaris Spiladelpha barovskii kiritschenkoi see nigripes sugar cane scale see Aulacaspis Ceratomegilla barovskii Stethorus madecassus 143 tegalensis kiritschenkoi Stethorus picipes see Stethorus punctum sugar cane woolly aphid see spined soldier bug see Podisus picipes Ceratovacuna lanigera maculiventris Stethorus punctillum see Stethorus sugar spraying 187,222 Spiraea 66 pusillus suitability of food/prey 144, 145- spiraea aphid see Aphis spiraecola Stethorus punctum 145, 196. 218. 147. 175 spiralling whitefly see Aleurodicus 224. 493 food combinations 155,156 dispersus Stethorus punctum picipes 143. 221. Sukunahikonim tribe 5 Spiroplasmataceae 30. 30, 68, 176, 299. 450, 499 sulphur dioxide pollution 26 421.422,424-425 diapause 299-300 sum of effective temperatures (SET) Spodoptera litura 3 79 Stethorus pusillus 56. 81, 82, 89. 92 spores, fungal 180 90.94, 96, 131. 143, 167 relationship with LDT 92, 93-94. collection of by larvae 183 .183, 196.217.218,299. 95 found in gut 2 80 313-314. 322.353.378. teneral pre-oviposition period 96 spot pattern, variation in 19-20, 417.4/8.490.493, 533 sunflower see Hclianthus annuus 20, 21-22. 22, 24. 25 Stethorus species, biocontrol 490 sunshine hours and melanism 26 spotted alfalfa aphid see Therioaphis Stethorus tridens 196.205 supercooling point (SCP) 96, trifolii Stethorus vegans 490 328-329 spotted flycatcher see Muscicapa Sticholotidinae subfamily outdoor and indoor overwintering striata characteristics 5-6 331 spotted lady beetle see Coleomegilla division of 3-4 supergenes 22 maculata sticky cards, sampling 113 supernumerary (B) chromosomes sprays of food substitutes 187.222. stinging nettle see Urtica dioica 15-16 499 strawberry aphid see Chaetosiphon supplementary food 123 spruce aphid see Elatobium abietinum fragaefolii supranivean hibernators 328. squash beetle see Epilachna borealis strawberry bug see Calocoris 329-330 squash or pumpkin see Cu'curbita norvegicus survival maxima strip cutting/harvesting 126, 501 on aphid prey 149. 162 starvation 202 striped ladybird see Myzia on artificial diets 2 86 aphid defensive measures causing oblongoguttata hibernating on higher ground 241 striped mealybug see Ferrisia virgata 305 foraging activity, effect on 218. Strongygaster triangulifera 386 overwintering 308,312. 221 Sturnus vulgaris 379 328-330, 331 and ovarian development 78, 82 Styrax 382 on plant foods 299 pollen feeding preventing 181 Subcoccinella vigintiquatuorpunctata and temperature extremes temporary, effect on reproduction 28,39.67. 76.81. 312.378. 96-97 209 385-386. 389.456 swarms 235-236 statistical approaches, predation subfamilies (of Coccinellidae) sweeping, sampling method 112 471-472 characteristics of 5-10 sweet orange see Citrus sinensis steelblue lady beetle see Halmus classification changes 3-5 swede midge see Contarinia nasturtii chalybeus parasitoids of 390 sycamore aphid see Drepanosiphum stenophagous species see specialist proposed classification 5. 6 platanoidis species 154 subnivean hibernators 328, 329 Sylvia atricapilla 379 Subject index 559

Sylvia borin 379 pupal colouration 76 thermal guild partition 344-345 Sylvia communis 379 pupal size 74 thermal melanism, pupae 76 Sylvia curruca 379 pupation timing and 75 thermal microhabitats 75 Sylvia nisoria 379 and rate of natural increase 143 thermal window 9 5 Symphoricarpos rivularis 120 reproduction, effect on 96 thermoregulatory basking 125 Symydobius oblongus 143 storage, developmental stages 332 thirteen spot ladybird see Hippodamia synchronization of hatching 56. tolerance to extreme 96-97 tredecimpunctata 68-69 temporal guild partition 344 Thomsonina see Nothoserphus Synhannonia conglobata see Oeiiopin temporal niches 128 three-banded lady beetle see Coccinella conglohata temporal patterns, prey density 211 trifasciata Synona obscura 405 temporal variation, colour pattern three-striped lady beetle see Brumoides Synonycha grandis 57. 67. 163. 196. polymorphism 25 suturalis 382,498 ten spot ladybird see Adalia Thrips tabaci 75. 174 Syntomosphyrus taprobanes see decempunctata thrush nightingale see Luscinia Oomyzus scaposus teneral development 76-77 luscinia syrphid larvae, attack by 358 thermal requirements 95-96 tiger moths see Arctia Syrphoctonus tarsalorius 388 terpenoids. coccinellid avoidance of tillage, reduction of 502, 503 458 tobacco budworm see Heliothis Tachyporus sp. 2 77.288 testicular follicles, diapause 28 i. virescens take-off, flight behaviour 77, 304 310.328, 321-323 top predators, coccinellids as Talitium triangulare 169 tethered flight 303.315-316 354-355 Tamarix 200,361.470 Tetrabrachini tribe 9 total food consumption 202-204 Tamarixia radiata 361.470 Tetramoriwn caespitum 381 toxicity. pesticides 506-509 Tanacetum vulgare 126 Tetranychus evansi 119. 196, 205, toxic prey 146,157-165 Tandonia budapestensis 477 224 toxic substances in prey 451-452 tansy see Tanacetum vulgare Tetranychus lintearius 119, 200. toxins, insect-resistant GM crops Tapinoma nigerrimum 380 224,299.498. 506 505-506 Taraxacum offlcinale 501 Tetranychus mcdanidi 196, 490 Toxoptera aurantii 225, 450 tarnished plant bug see Lygus Tetranychus telarius see Tetranychus Toxoptera citricidus 146. 163. 196 lineolaris urticae control of 494 Tasmanian ladybird see Cleobora mellyi Tetranychus urticae 67. 152, 1S7. Toxoptera graminum see Schizaphis tawny pipit see Anthus campestris 196.354, 532 graminum tea aphid see Toxoptera aurantii Tetrastkhus coccinellae see Oomyzus tracks, larval, oviposition deterrence Telsimiini tribe 5, 8 scaposus 229-233 temperature Tetrastichus cydoniae 386 trade-offs, genetic 28-29 adult size 78 Tetrastkhus decrescens 385 transgenic plants 122-123 and development 91-97 Tetrastichus epilachnae 385 transverse lady beetle see Coccinella determining course of events in Tetrastichus 409 transversoguttata adult life 95-96 Tetrastichus orissaensis 386 traps for sampling 113—114 relationship between LDT and Tetrastichus melanis see Oomyzus tree pipit see Anthus trivialis SET 92-95 scaposus tree sparrow see Passer montanus and diapause 276-2 79, 280, Tetrastichus sexmaculatus see Oomyzus trees 285.289 scaposus age of 122 effects of low 328-332 Thalassa saginata 76 and coccinnelid abundance 131. extremes, tolerance to 96-97 Thanatus 354, 532 132 fecundity 88, 89. 90. 96 Thea vigintiduopunctata see Psyllobora habitats 129 and food consumption 202-203 vigintiduopunctata sampling from 113 and frequency of short flights 234 Thelaxes dryophila 189 Trialeurodes vaporariorutn 66, 174. hatching rate varying with 68 thelytokous parthenogenesis 391. 195 life span decrease with 90-91 406 triangular fecundity function 86, and melanism 26 Therioaphis maculata see Therioaphis 87 mating advantage 26-27 trifolii Trichogramma evanescens 170, 353 and oviposition 233 Therioaphis trifolii 193.211 Trichogramma 358. 363, 386 oviposition period 88 thermal constants 92 Trichomalopsis acuminata 386 560 Subject index

Trichotnalopsis 386, 389 Uga sinensis 400 Virginian pencil cedar see Juniperus Trichomalopsis dubia 389 Uga species 399. 400, 400 virginiana trichomes 200,217.227,353.504 Uhnus 314 vision 222. 22 '.-224. 239 Trkhoplusia ni 170. 353. 479 Unaspis citri 492 visual counting, sampling method Trichosanthes kirilowii 201 Unaspis euonymi 190. 202 112 Tripleurospermum maritimum 117. Unaspis yanonensis 190 vitellogenin synthesis 327-328 121 univoltine species 90 volatiles, plant 200. 224-226. Tripolycystus see Metastenus univoltinism 449-451 Triticum 153 Chilocorus spp. 298 link to aphid density 456 tritrophic studies 196-198 C. septempunctata 279, 280. 281. link to patch preference 457 trophic eggs 69, 177 283.284. 285 voltinism 90. 300-301. 307 trophic pathway 475. 476. 477. Hip. convergens 292 see also bivoltinism: multivoltinism: 481 potential multivoltines 286-287 univoltinism Tsuga 153.175 resilience 300 voracity 255. 489 Tuberculatus annulatus 58, 153. upper temperature threshold 92 quantifying 467-468 158 Uroleucon aeneum 158.191 Tuberolaclmus salignus 153. 188 Uroleucon umbrosiae 163 walnut aphid see Chromaphis Tubulinosema hippodamiae 418-419 Uroleucon cichorii 217. 238 juglandu ola Turdus merula 379 Uroleucon cirsii 153.188.191. warm acclimation 326 Turdus philomelos 379 195 Wasmannia auropunctata 449 turnip aphid see Lipaphis Uroleucon compositae 61. 148. 195 Watanabeia see Nothoserphus 404 pseudobrassicae Uroleucon formosanum 195 water content, diapause 326 twelve spotted ladybeetle see Uroleucon jaceae 195.217 water ladybird see Anisosticta Coleomegilla maculata lengi Uroleucon nigrotuberculatum 126 novemdecimpunclata twenty four spot ladybird see Uroleucon species 148. 216, 217, wax moth see Galleria mellonella Subcoccinella 238 waxy surfaces vigintiquatuorpunctata Urtica dioica 11 7. 121, 126. 150. aphid prey I 65. 166 twenty-spotted lady beetle see 153.501 and biological control 504 Psyllobora vigintimaculata host plants 124,206,217-218 twenty two spot ladybird see vacuum sampling. D-Vac 112 larval protection 236,237, Psyllobora vigintiduopunctata vagility. interspecific differences in 241 twice-stabbed lady beetle see 234 weeds, positive effect of 1 20. 121. Chilocorus stigma variegated lady beetle see Hippodamia 126 two spot ladybird see Adalia bipunctata variegata weight two-spotted spider mite see vedalia beetle see Rodolia cardinalis adults 78-82 Tetranychus urticae vegetation diversity and aggregative egg clusters 57 Typha latifolia 152 numerical response 213 eggs 56 Tyria 164 Verania see Micraspis larvae 73 Tytthaspidini tribe 6-7 Verbascum thapsus 312 pupae 74-75 Tytthaspis sedecimpunctata 81. 131. vertebrate predators 377 western corn n >otworm see Diabrotica 142. 155. 183. 784.200. vertical distribution of coccinellids on virgifera 241. 309. 312, 378. 393. host plant 222, 125 western plant bug see Lygus hesperus 399. 420 Verticillium lecanii see Lecanicillium wheat bug see Nysius huttoni diet of 200-201 lecanii wheat see Triticum dormancy behaviour 312-313 Vesiculaphis caricis 191 white (or yellow) mustard see Sinapis mandible 783 vetch aphid see Megoura viciae alba Tytthaspis trilineata see Coccinella vetch or tare see Vicia sativa wild herb habitats 127. 128. 129. nigrovittata Vibidia duodecimgul tata 81,313, 131 316. 378 willow aphid see Tidierolachnus Uga colliscutellum 400 Vicia faba 58, 88, 91. 123, 160- salignus Uga coriacea 400 161. 765. 169, 196. 197, willow warbler see Phylloscopus Uga digitata 400 210,225.237.452 trochilus Uga hemicarinata 400 Vicia sativa 160 window traps 113,305 Uga javanica 400 Vigna ungukulata subsp. cylindrica wing polymorphism 2 7-28 Uga menoni 400 160 wings and wiuglessness 77 Subject index 561

Withania somnifera 410 woolly vine scale see Pulvinaria vitis Y-tube olfactometers 222,223,224, Wolbachia 30, 30, 38, 39, 68, 176, wormwood see Artemisia vulgaris 225 349.421,422-423, 424-425 Xanthogaleruca luteola 170 (Z)-jasmone 450 wood ant see Formica rufa Xy parachute (Xyp) system 14-15 (Z)-pentacos-12-ene, in larval tracks wood nuthatch see Sitta europaea 231, 455 woolly apple aphid see Eriosoma yellow lupin see Lupinus luteus Zagloba aeneipennis 236 lanigerum yolk, eggs 55.56.82 'lea mays 152,182,282,227