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Aphids as Crop Pests
This page intentionally left blank
Aphids as Crop Pests
Edited by
Helmut F. van Emden
Emeritus Professor of Horticulture, University of Reading, UK
and
Richard Harrington
Plant and Invertebrate Ecology Division, Rothamsted Research, UK

CABI is a trading name of CAB International
CABI Head Office CABI North American Office

Nosworthy Way 875 Massachusetts Avenue
Wallingford 7th Floor
Oxfordshire OX10 8DE Cambridge, MA 02139
UK USA
Tel: +44 (0)1491 832111 Tel: +1 617 395 4056
Fax: +44 (0)1491 833508 Fax: +1 617 354 6875
E-mail: cabi@cabi.org E-mail: cabi-nao@cabi.org
Website: www.cabi.org
© CAB International 2007. All rights reserved. No part of this

publication may be reproduced in any form or by any means,
electronically, mechanically, by photocopying, recording or
otherwise, without the prior permission of the copyright owners.
A catalogue record for this book is available from the British Library,
London, UK.
A catalogue record for this book is available from the

Library of Congress, Washington, DC.
ISBN-13: 978 0 85199 819 0
Typeset by AMA DataSet Ltd, UK.

Printed and bound in the UK by Cromwell Press, Trowbridge.
The paper used for the text pages in this book is FSC certified. The FSC (Forest
Stewardship Council) is an international network to promote responsible management
of the world’s forests.
In Memoriam
François Leclant
Emeritus Professor of Entomology, ENSA, Montpellier, France
who died while this book was in preparation
He told me he particularly enjoyed writing his chapter,

since it was one of the few times he forgot his
terminal illness. His research was highly respected
internationally, he was a gifted teacher, and
a generous friend (HvE)
Contents
Preface xxi
List of Contributors xxv
1. Taxonomic Issues 1
Roger L. Blackman and Victor F. Eastop
Introduction 1
Interpreting variation in aphids 3
The taxonomy of pest aphids – what’s in a name? 4
The 14 aphid species of most agricultural importance 6
Acyrthosiphon pisum (pea aphid) 7
Aphis craccivora (cowpea aphid) 8
Aphis fabae (black bean aphid) 8
Aphis gossypii (cotton or melon aphid) 10
Aphis spiraecola (green citrus aphid or spiraea aphid) 11
Diuraphis noxia (Russian wheat aphid) 12
Lipaphis pseudobrassicae (mustard aphid) 13
Macrosiphum euphorbiae (potato aphid) 14
Myzus persicae (peach–potato aphid) 15
Rhopalosiphum maidis (corn leaf aphid) 17
Rhopalosiphum padi (bird cherry–oat aphid) 18
Schizaphis graminum (greenbug) 18
Sitobion avenae (grain aphid) 20
Therioaphis trifolii (spotted alfalfa and yellow clover aphids) 21
Conclusions 22
References 22
2. Population Genetic Issues: The Unfolding Story Using

Molecular Markers 31
Hugh D. Loxdale and Gugs Lushai
Introduction 31
vii
viii Contents
Application of molecular markers in aphidology 32

Detection of genetic variation 32
Molecular phylogenetics and karyotypes 34
Clonal studies 34
Life cycle 37
Adaptation to host plants 43
Insecticide resistance 48
Pathogenicity 48
Dispersal and geographic range 49
Geographic colonization 54
Conclusions 56
Acknowledgements 57
References 57
3. Life Cycles and Polymorphism 69

Iain S. Williams and Anthony F. G. Dixon
Introduction 69
Types of life cycle 69
Host-alternating aphids 70
Non-host-alternating aphids 72
Two-year life cycles 73
Holocyclic and anholocyclic life cycles 73
Factors determining the production of different morphs 73
Determination of sexual morphs 74
Determination of asexual winged morphs 75
Diapausing morphs 77
Colour polymorphism 77
The importance of different morphs for crop protection 78
Pest status 78
Virus transmission 78
Forecasting 79
The importance of polymorphism in
determining pest status 79
Conclusions 81
References 81
4. Host-plant Selection and Feeding 87

Jan Pettersson, W. Fred Tjallingii and Jim Hardie
Introduction 87
Orientation and host-plant finding 87
Visual responses 88
Olfactory responses 88
Plant contact after landing 90
Plant penetration and feeding 93
Pathway phase 93
Phloem phase 99
Phloem feeding 101
Xylem drinking 102
Leaving a plant 103
Plant responses and predisposition to aphid feeding 104
Aphids and host plants – a challenging research field 106
Contents ix
Conclusions 106
References 107
5. Nutrition and Symbiosis 115

Angela E. Douglas and Helmut F. van Emden
Introduction 115
Microbial symbiosis in aphids 115
Diversity of microorganisms 115
Location of symbiotic microorganisms 116
Acquisition of symbiotic microorganisms 117
Significance of symbiotic microorganisms to aphids 117
Carbon nutrition 118
Sugars 118
Lipids 120
Nitrogen nutrition 121
Amino acids and their sources 121
The fate of amino acids in aphids 122
Other dietary sources of nitrogen 123
Minerals and micronutrients 123
Artificial diet 123
History of artificial diets for aphids 123
Recipe for the diet and practical procedures 124
Relation between the diet and plant phloem sap 125
Aphid performance on the diet 127
How useful is the diet for studies on aphid nutrition? 129
Conclusions and future prospects 129
Acknowledgements 130
References 130
6. Growth and Development 135

Caroline S. Awmack and Simon R. Leather
Introduction 135
Definitions 135
Uses of aphid growth and developmental rates 136
Measurement of aphid growth and development rates 136
Growth rates 136
Developmental rates 137
The intrinsic rate of increase, rm 138
Experimental techniques 138
Aphid cages 138
Disadvantages of aphid cages 139
Factors affecting aphid growth and development 140
Factors affecting the reliability of size x fecundity relationships 141
Difference between nymphs destined to be apterous and alate 143
Temperature 143
Population-scale factors 144
Conclusions 144
References 144
7. Aphid Movement: Process and Consequences 153

Michael E. Irwin, Gail E. Kampmeier and Wolfgang W. Weisser
Introduction 153
Pest status 153
x Contents
The nature of aphid movement 154

Conceptual framework for aphid movement 155
Modes of transport 155
Scales of displacement 155
Aphid life stages, morphs and active modes of transport 157
The migratory process 159
Events leading to take-off 160
Take-off and ascent 161
Horizontal translocation 163
Switching off the migratory urge 164
Appetitive dispersal 164
Stimuli that cause aphids to disperse in
an appetitive manner 164
Response to multiple perturbations 168
Landing and alighting cues 168
Incorporating aphid movement into Pest
Management strategies 169
Components of IPM programmes 170
Categories of aphids as agricultural pests 174
Conclusions 176
Acknowledgments 177
References 177
8. Predators, Parasitoids and Pathogens 187

Wolfgang Völkl, Manfred Mackauer, Judith K. Pell and
Jacques Brodeur
Introduction 187
Predators 188
Coccinellidae (ladybird beetles) 188
Syrphidae (hover flies) 192
Chrysopidae and Hemerobiidae (lacewings) 194
Cecidomyiidae (predatory midges) 196
Other arthropod predators 198
Parasitoids 199
General biology 199
Foraging behaviour and host finding 201
Patterns of resource use 203
Aphid pathogens 206
Biology of entomophthoralean fungi 207
Epizootiology of entomophthoralean fungi 208
Intraguild interactions, mutualistic ants and aphid symbionts 210
Intraguild predation and competition 210
The effects of mutualistic interactions with ants on
predation and parasitism 212
Aphid symbionts and resistance to parasitism 214
Conclusions 215
References 215
9. Chemical Ecology 235

John A. Pickett and Robert T. Glinwood
Introduction 235
Methods 236
Contents xi
Interactions between aphids 238

Sex pheromones 238
Alarm pheromones 240
Social interactions: aggregation, density regulation and avoidance 243
Interaction with plants 245
Host-plant semiochemicals 245
Non-host-plant semiochemicals 247
Aphid effects on the plant 248
Interaction with natural enemies 248
Responses of natural enemies to aphid-produced chemicals 248
Responses of natural enemies to aphid-induced plant signals 252
Conclusions 253
References 254
10. Insecticide Resistance 261

Stephen P. Foster, Gregor Devine and Alan L. Devonshire
Introduction 261
Diagnosis of resistance in aphids 262
Relevance of bioassays to field control 263
Biochemistry and molecular basis of resistance 263
Resistance mechanisms in Myzus persicae (peach–potato aphid) 264
Resistance mechanisms in other aphid species 268
Neonicotinoid resistance 271
Factors affecting the dynamics of insecticide resistance in the field 272
Selection pressures 272
Ecological factors 273
Pleiotropic effects of resistance 274
Aphids in which resistance poses a minor or potential threat 276
Conclusions 277
References 278
11. Coping with Stress 287

Jeffrey S. Bale, Katherine L. Ponder and Jeremy Pritchard
Introduction 287
Plant factors 287
Nutritional stress 288
Non-nutritional plant factors 289
How aphids cope with nutritional stress 292
Temperature stresses 293
Effects of temperature on development, reproduction and movement 293
Cold Stress 294
Coping with cold 296
Heat stress 300
A molecular approach to understanding aphid responses to stress 300
Conclusions 302
References 303
12. Population Dynamics 311

Pavel Kindlmann, Vojtgch Jarosík and Anthony F.G. Dixon
Introduction 311
xii Contents
Biological background 312

Aphid biology relevant to population dynamics 312
Biology of natural enemies relevant to aphid population dynamics 314
Theory of aphid population dynamics 316
Features of aphid population dynamics that
should be incorporated in models 316
Regression model 316
Regression model with stochasticity 317
Logistic model with variable ‘carrying capacity’ 318
Cumulative density model 318
Logistic model with variable carrying capacity and growth rate
affected by cumulative density 319
Comparison of the different population models 319
Natural enemies and aphid abundance 321
Practical problems 321
Field estimates of abundance and population growth rate 321
Trap estimates of abundance 323
Consequences for pest management 324
Conclusions 324
References 324
13. Feeding Injury 331

Sharron S. Quisenberry and Xinzhi Ni
Introduction 331
Injury classification 332
Asymptomatic injury 332
Symptomatic injury 334
Aetiological agents in Aphidoidea saliva 340
The biochemical and physiological bases for symptoms of injury 340
Basis of asymptomatic injury 340
Basis of symptomatic injury 340
Ecological modulation of aphid injury 342
Abiotic factors 342
Biotic factors 344
Ramifications of aphid injury research for crop protection 345
Conclusions on research frontiers in the understanding of
injury from aphid feeding 346
References 347
14. Transmission of Plant Viruses 353

Nikos I. Katis, John A. Tsitsipis, Mark Stevens and Glen Powell
Introduction 353
Virus–vector interactions 354
Aphids as effective virus vectors 357
Host selection related to virus acquisition and inoculation 357
Transmission modes of plant viruses by aphids 359
Non-persistent transmission 359
Semi-persistent transmission 362
Persistent transmission 366
Epidemiology 370
Biological factors 370
Contents xiii
Cultural practices 372

Weather 372
Disease forecasting 372
Disease management strategies 372
Use of genotypes resistant to the virus and/or to the aphid vectors 373
Elimination of virus sources 374
Prevention or reduction of virus spread 374
Conclusions 377
References 377
15. Chemical Control 391

Alan M. Dewar
Introduction 391
Choice of aphicide 391
Modes of action 393
Organophosphates and carbamates 393
Pyrethroids 393
Neonicotinoids 393
Pymetrozine 394
Diafenthiuron 395
Triazamate 395
Adjuvants and synergists 395
Application 395
Thresholds for control 398
Wheat and barley 398
Pea 399
Field bean 399
Brassicas 399
Cotton 400
Sugar beet 400
Top fruit 400
Efficacy 400
Poaceae 400
Soft Fruit 401
Top fruit 402
Solanaceae 403
Chenopodiaceae 405
Leguminosae 405
Brassicas 406
Lettuce 407
Other vegetables 407
Malvaceae 408
Others 409
Conclusions 410
References 411
16. Cultural Control 423

Steve D. Wratten, Geoff M. Gurr, Jason M. Tylianakis and
Katherine A. Robinson
Introduction 423
Overview 423
xiv Contents
Mulches 426
Row covers 427
Particle films 427
Sowing and planting date 428
Plant density 429
Crop-plant pruning 430
Irrigation and fertilizer management 430
Intercropping, living mulches and cover crops 431
Trap crops 434
Provision of resources for natural enemies 434
Provision of refuges for natural enemies 437
Conclusions 438
References 439
17. Host-plant Resistance 447

Helmut F. van Emden
Introduction 447
Types of host-plant resistance to aphids 447
Antixenosis 449
Antibiosis 449
Tolerance 450
What is the preferred type of resistance? 450
Mechanisms of host-plant resistance to aphids 450
Mechanisms of antixenosis 451
Mechansims of antibiosis 453
Mechanisms of tolerance 456
Further considerations 457
Yield drag or other fitness costs 457
Negative effects on natural enemies 457
Problem trading 459
Biotypes 459
Spread of virus 461
Interactions with other control measures 462
Conclusions 462
Acknowledgments 462
References 462
18. Biological Control 469

Wilf Powell and Judith K. Pell
Introduction 469
Parasitoids 471
Cereal aphids 471
Legume aphids 475
Aphids on orchard crops 476
Banana aphids 477
Aphids on forest and urban trees 477
Glasshouse aphids 478
Aphidoletes aphidimyza (predatory gall midge) 479
Aphids on field and orchard crops 484
Coccinellidae (ladybird beetles) 485
Contents xv
Aphids on field crops and amenity plants 486

Aphids on orchard crops 489
Lacewings 489
Aphids on field crops and amenity plants 493
Aphids on orchard crops and tree seedlings 494
Syrphidae (hover flies) 494
Entomopathogenic fungi 494
Aphids on field crops 497
Cotton aphids 498
Aphids in orchards 498
Aphids on vegetables 499
Conclusions 499
References 500
19. Monitoring and Forecasting 515

Richard Harrington, Maurice Hullé and Manuel Plantegenest
Introduction 515
Why monitor and forecast? 515
What should be monitored and forecast? 515
Where should monitoring and forecasting be carried out? 516
When should monitoring and forecasting be undertaken? 516
How can aphids be monitored and forecast? 516
Monitoring 516
Databases and data handling 524
Forecasting 524
Conclusions 531
References 531
20. Integrated Pest Management and Introduction to IPM Case Studies 537
Helmut F. van Emden
Introduction 537
IPM as the use of multiple control measures 538
The golden rules of IPM 538
Interaction between chemical and biological control 538
Interaction between chemical control and host-plant resistance (HPR) 539
Interaction between biological control and host-plant resistance 540
Three-way interaction between chemical control, host-plant
resistance and biological control 543
Interaction between cultural control and biological control 543
The IPM case studies 543
Conclusions 544
References 546
21. IPM Case Studies: Brassicas 549

Rosemary H. Collier and Stan Finch
Introduction 549
Biology of pest aphids in the UK 549
Integrated crop management 550
Chemical control 551
xvi Contents
Monitoring and forecasting 552

Sampling and decision-making 553
Biological control 554
Host-plant resistance 554
Cultural control 554
Crop covers 554
Increasing crop diversity 555
Executive summary 555
References 557
22. IPM Case Studies: Berry Crops 561

Rufus Isaacs and J.A. Trefor Woodford
Introduction 561
Aphid IPM in raspberry 561
Aphid IPM in blueberry 565
Aphid IPM in strawberry 568
Aphid IPM in grape 568
References 569
23. IPM Case Studies: Cotton 573

Jean-Philippe Deguine, Maurice Vaissayre and the late François Leclant
Introduction 573
Biological and ecological characteristics of Aphis gossypii 573
Infestation 573
Damage and its economic importance 575
Control and its evolution 576
The limits of chemical control of A. gossypii 576
Evolution towards integrated control 576
IPM of Aphis gossypii in the USA 576
IPM of Aphis gossypii in Brazil and elsewhere in South America 577
IPM of Aphis gossypii in South-east Asia and China 578
IPM of Aphis gossypii in Africa and Madagascar 578
References 581
24. IPM Case Studies: Leafy Salad Crops 587

G. Mark Tatchell
Introduction 587
What are leafy salad crops? 587
The IPM challenge 587
The aphids 588
Components for aphid IPM 588
Chemicals and resistance to insecticides 588
Reducing insecticides 590
Modifying aphid behaviour 591
IPM in practice 591
Assurance and accreditation schemes 591
Practical control 591
Contents xvii

References 594
25. IPM Case Studies: Grain 597

Hans-Michael Poehling, Bernd Freier and A. Michael Klüken
Introduction 597
Population dynamics 598
Damage 599
Direct damage 599
Indirect damage 599
Control possibilities 600
Forecasting systems 600
Monitoring and decision-making systems 600
Habitat management practices 604
Conclusions 605
References 606
26. IPM Case Studies: Seed Potato 613

Edward B. Radcliffe, David W. Ragsdale and Robert A. Surányi
Introduction 613
Certification 613
Vector species 614
Economic thresholds 616
Mathematical modelling 616
IPM tactics 616
Monitoring aphid flight 616
References 621
27. IPM Case Studies: Sorghum 627

Gerald J. Michels, Jr. and John D. Burd
Introduction 627
A short history of Schizaphis graminum on sorghum 627
Current greenbug management practices 629
Putting the pieces together 633
The ‘current model’ and the future 633
Conclusions 634
References 634
xviii Contents
28. IPM Case Studies: Cucurbits 639

Susan E. Webb
Introduction 639
Aphid-vectored viruses affecting cucurbits 639
Management options for aphids and aphid-vectored
viruses in cucurbits 641
Cross-protection 643
IPM programmes for cucurbits that include
aphid management 645
References 646
29. IPM Case Studies: Deciduous Fruit Trees 651

Sebastiano Barbagallo, Giuseppe Cocuzza, Piero Cravedi and
Shinkichi Komazaki
Introduction 651
Apple and pear aphids 651
Damage and virus transmission 651
Monitoring and economic thresholds 654
Chemical and supervised control 655
Stone-fruit tree aphids 655
References 660
30. IPM Case Studies: Tropical and Subtropical Fruit Trees 663
Sebastiano Barbagallo, Giuseppe Cocuzza, Piero Cravedi and
Shinkichi Komazaki
Introduction 663
Citrus aphids 663
Tropical fruit tree aphids 669
Supervised chemical control 673
References 674
Contents xix
31. Decision Support Systems 677

Jonathan D. Knight and Deborah J. Thackray
Introduction 677
What is a Decision Support System? 677
Are aphids suitable subjects for the development of DSSs? 678
Examples of aphid and virus DSSs 678
Development and delivery of DSSs 679
Case studies 679
New Zealand Decision Support System 681
Western Australian Decision Support System 683
Conclusions 685
References 686
Taxonomic Glossary 689

Index 699
The colour plate section can be found following page 356
Colour plates sponsored by Syngenta and BASF

Many aphids show considerable colour variation; mostly only one colour variant is
illustrated.
Preface
Following the publication in 1998 of Thrips as Crop Pests, edited by Trevor Lewis, CABI
commissioned Aphids as Crop Pests as a second in the series. However, in prefacing this
book, we wish to pay tribute to another predecessor, Aphids: Their Biology, Natural Ene-
mies and Control, edited by Albert Minks and Paul Harrewijn and published in three vol-
umes by Elsevier between 1987 and 1989. This comprehensive and impressive work has
been the standard reference text for aphidologists for nearly 20 years, and we would not
presume to make it redundant with the publication of Aphids as Crop Pests. Therefore, in
planning the content of our book, we have concentrated on the applied aspects of
aphidology and have not sought to match the comprehensive coverage of morphology,
physiology and ecology found in Aphids: Their Biology, Natural Enemies and Control. As
far as aphid biology is concerned, we have instead emphasized areas such as current taxo-
nomic issues (Chapter 1), host selection and feeding (Chapter 4), nutrition and symbionts
(Chapter 5, which includes full instructions for successful long-term rearing of Myzus
persicae on synthetic diet), chemical ecology (Chapter 9) and reactions to stress (Chapter
11) where there have been major advances in the last fifteen years. We would especially
direct the reader to Chapter 2 on Population Genetics; molecular markers were hardly
available when Minks and Harrewijn’s book was being put together. We have therefore
given a large page allocation to this topic. In similar vein, molecular methods have enabled
many substantial recent advances in our understanding of the mechanisms of aphid resis-
tance to insecticides (Chapter 10). We also felt that life cycles and polymorphism (Chapter
3), Growth and Development (Chapter 6) and aphid movement (Chapter 7) were areas
particularly relevant to aphids as pests.
We have been greatly encouraged by the readiness with which the potential authors
we first contacted agreed to participate. Having recruited scientists we felt were at the
forefront of their subject, it made sense to leave the selection of any co-authors to them.
The result is that Chapter 12 (Population Dynamics) has been brought up to date with a
strong modelling component, and that Chapter 8 on the natural enemies of aphids has
involved four leading specialists – one for each of the major taxa. This led us to expand the
page allocation for this chapter also.
The more identifiably applied part of the book starts with two chapters on the injury
caused to crops by aphid feeding (Chapter 13) and by the transmission of plant viruses
(Chapter 14). Both these chapters considerably update previous reviews on these topics.
xxi
xxii Preface
Separate chapters then cover the four main building blocks of pest management. Chap-
ter 15 on Chemical Control has much new material with the advent of neonicotinoid com-
pounds, which are especially effective against aphids, and Chapter 16 (Cultural Control)
highlights how far conservation biological control has come into practice. Chapter 17 cov-
ers Host Plant Resistance and Chapter 18 deals with Biological Control. This latter chapter
describes practical interventions, and avoids making a stronger case for the method than is
realistic. The involvement of a specialist on aphid pathogens enables these important tools
to receive adequate treatment.
Chapter 19 on Monitoring and Forecasting is again very different from what could
have been written on this topic in the 1980s, since advances in information technology
have enabled huge databases to be feasible and have made possible rapid transmission of
data and effective international networks.
Chapter 20 introduces the concepts and potential of IPM, pointing out that it origi-
nated in California with work on aphids. Specialists on aphid problems in selected crop-
ping scenarios then review the status of IPM in a series of Case Studies (Chapters 21–30).
We felt that the diversity of cropping-systems, pests and natural enemies, chemicals avail-
able etc. in different continents made a world-wide coverage for each crop too complex a
proposition, and we encouraged authors to concentrate on the region in which they them-
selves worked. Imposing a template for these chapters would have clouded the very differ-
ent issues that relate to aphids in these case studies, and we have taken the alternative
approach towards consistency of ending each case study with an “Executive Summary”,
initially written by the editors on the basis of the text supplied by the chapter author(s) but
then mutually agreed with them. The final Chapter on Decision Support Systems is another
topic which has only developed strongly since the 1980s.
Inevitably, various problems forced a few of our original authors to withdraw and so
we are especially grateful to those who only started on their chapters later and to the other
authors for their patience when they saw these chapters delaying production of the book. We
are also indebted to our authors for their invariably positive response to our comments and
suggestions for changes; they all readily accepted that any large group of virtuoso performers
needs a conductor!
The editors of a multi-author book face many problems, in particular that of repetition:
take, for example, resistance to insecticides. Although our book has a chapter specifically
on this topic, it also is a phenomenon of population genetics, it governs what chemicals
can be used and so is relevant to the Chemical Control chapter and many of the IPM Case
Studies. It would be tidy to eliminate duplication, but we are not so naïve as to think any-
one is going to read our book from start to finish. Rather, each chapter needs to stand on its
own as a comprehensive review. We have used cross references to other chapters as a par-
tial solution, but have frequently left some discussion of a topic covered in another chapter
where a cross reference alone would fail to make the point(s) the author had intended.
Another problem is the nomination of pesticides for aphid control, particularly in the
IPM Case Studies. What products are permissible for use varies from country to country
and changes almost weekly even within countries! Individual experts have therefore
referred to products in relation to their own experience and environment. Mention of a
compound should not be taken to mean it can be used elsewhere, nor necessarily after
2006.
The chapters are followed by a “Taxonomic Glossary”. Here we have listed the Latin
names of species mentioned in the book under the appropriate taxon, and then in alphabet-
ical order together with the taxonomic authority and any generally accepted English com-
mon name(s). Such a glossary had to be compiled in order to get consistency between
chapters in the citing of Latin names, and including it in the volume has meant we could
avoid adding the taxonomic authority and Latin names of well-known crops in the individ-
ual chapters. In compiling the glossary we have encountered different practices in
Preface xxiii
different taxa (e.g. insects, fungi, plants etc.) in how authorities and subspecies are cited.
Thus entomologists tend to assume that a trinomial indicates a subspecies whereas bota-
nists still insert “subsp.” before the last name. We decided to follow the accepted practice
for each taxon rather than impose uniformity. We are extremely grateful to the specialists
who advised on current taxonomic usage, Dr Anne Baker (mites), Dr Roger Blackman
(aphids), Mr Barry Bolton (ants), Dr Roland Fox (other fungi and bacteria), Dr Francis
Gilbert (syrphids), Dr Simon Gowen (nematodes), Mr Paul Hillyard (spiders), Dr Stephen
Jury and Mr Ronnie Rutherford (plants), Dr Ivo Kovar and Dr Mike Majerus (coccinellids),
Dr Martin Luff (carabids), Professor Tim New (lacewings), Dr Judith Pell (entomopatho-
gens) and Mr Nigel Wyatt (Diptera). Another specialist who provided valuable advice was
Professor Roger Plumb in relation to the nomenclature of plant viruses.
Generous sponsorship from BASF and Syngenta has made it possible to include a sec-
tion of colour plates of the aphids selected by the authors of Chapter 1 as the most serious
aphid pests worldwide. The section also includes colour plates of representatives of impor-
tant groups of natural enemies. We are extremely grateful for this encouragement and sup-
port from the agrochemical industry. The colour plates come from a variety of sources, and
we greatly appreciate the permission of copyright holders to reproduce their images with-
out paying a royalty. Many others (scientists and publishers) have also allowed us to use
their material in the various chapters; their colour images have then been converted to
grayscale. Our colleagues have without exception been most helpful in allowing us to use
copyright material – in every case we have acknowledged the source in the legends to
colour plates and other figures.
Part of the value of this book is in the comprehensive bibliographies accompanying
each chapter. In order not to restrict authors in reviewing their topic, the list of references
was not included in their page allocation. The work of editing the references was thus con-
siderable and we are grateful to Ms Berit Pederson and Mr Greg Bentley for assistance with
this chore.
We have both been researching on aphids for many years, and have become good
friends in the process. We have enjoyed working together on this book and have both some-
how found time to edit all chapters independently, though in each case one of us has taken
the lead as “first editor”. We will miss this collaboration now it has ended, but will feel
rewarded if this book is appreciated by our aphidological colleagues and proves of value to
them and those starting research on aphids for a good number of years to come.
Helmut van Emden

Richard Harrington
19th June 2006

List of Contributors
Caroline S. Awmack, Department of Entomology, University of Wisconsin-Madison,

Madison, WI 53706, USA (Chapter 6)
Jeffrey S. Bale, School of Biosciences, University of Birmingham, Edgbaston, Birmingham,
B15 2TT, UK (Chapter 11)
Sebastiano Barbagallo, Dipartimento di Scienze e Tecnologie Fitosanitarie, University of
Catania, Italy (Chapters 29, 30)
Roger L. Blackman, Department of Entomology, Natural History Museum, London, SW7
5BD, UK (Chapter 1)
Jacques Brodeur, Département de Phytologie, Université de Laval, Sainte-Foy, Québec,
G1K 7P4, Canada (Chapter 8)
John D. Burd, Plant Science and Water Conservation Laboratory, USDA-ARS, Stillwater,
OK 74075, USA (Chapter 27)
Giuseppe Cocuzza, Dipartimento di Scienze e Tecnologie Fitosanitarie, University of
Catania, Italy (Chapters 29, 30)
Rosemary H. Collier, Warwick HRI, The University of Warwick, Wellesbourne, Warwick,
CV35 9EF (Chapter 21)
Piero Cravedi, Istituto di Entomologia e Patologia Vegetale, University ‘Cattolica Sacro
Cuore’, Piacenza, Italy (Chapters 29, 30)
Jean-Philippe Deguine, Centre de Coopération Internationale en Recherche Agronomique
pour le Développement, Avenue Agropolis, 34398 Montpellier Cedex 5, France (Chapter
23)
Gregor Devine, Plant and Invertebrate Ecology Division, Rothamsted Research, Harpenden,
Herts, AL5 2JQ, UK (Chapter 10)
Alan L. Devonshire, Biological Chemistry Division, Rothamsted Research, Harpenden,
Alan M. Dewar, Entomology Research Group, Broom’s Barn Research Station, Higham,
Bury St. Edmunds, Suffolk, IP28 6NP, UK Correspondance address: Dewar Crop
Protection Ltd, Drumlanrig, Great Saxham, Bury St Edmunds, Suffolk, IP29 5JR, UK
(Chapter 15)
Anthony F. G. Dixon, School of Biological Sciences, University of East Anglia, Norwich,
NR4 7TJ, UK (Chapters 3, 12)
Angela E. Douglas, Department of Biology, University of York, Heslington, York, YO10
5YW, UK (Chapter 5)
xxv
xxvi List of Contributors
Victor F. Eastop, Department of Entomology, Natural History Museum, London, SW7 5BD,
UK (Chapter 1)
Helmut F. van Emden, School of Biological Sciences, University of Reading, Whiteknights,
Reading, Berks, RG6 6AJ, UK (Chapters 5, 17, 20)
Stan Finch, Warwick HRI, The University of Warwick, Wellesbourne, Warwick, CV35 9EF
(Chapter 21)
Stephen P. Foster, Plant and Invertebrate Ecology Division, Rothamsted Research,
Harpenden, Herts, AL5 2JQ, UK (Chapter 10)
Bernd Freier, Institute for Integrated Plant Protection, BBA, 14532 Kleinmachnow,
Germany (Chapter 25)
Robert T. Glinwood, Department of Ecology, Swedish University of Agricultural Sciences,
750 07 Uppsala, Sweden (Chapter 9)
Geoff M. Gurr, School of Rural Management, Charles Sturt University, PO Box 883,
Orange, New South Wales 2800, Australia (Chapter 16)
Jim Hardie, Division of Biology, Imperial College London, Silwood Park campus, Ascot,
Berks, SL5 7PY, UK (Chapter 4)
Richard Harrington, Plant and Invertebrate Ecology Division, Rothamsted Research,
Harpenden, Herts, AL5 2JQ, UK (Chapter 19)
Maurice Hullé, INRA, UMR BiO3P, 35653 Le Rheu, France (Chapter 19)
Michael E. Irwin, Department of Natural Resources and Environmental Sciences,
University of Illinois, Urbana, IL 61801, USA (Chapter 7)
Rufus Isaacs, Department of Entomology, Michigan State University, East Lansing, MI
48824, USA (Chapter 22)
Vojtech Jarosík, Department of Ecology, Charles University, 12844 Prague 2, Czech
Republic and Institute of Botany, Czech Academy of Sciences, 25243 Pruhonice, Czech
Republic (Chapter 12)
Gail E. Kampmeier, Section of Ecological Entomology, Illinois Natural History Survey,
Urbana, IL 61801, USA (Chapter 7)
Nikos I. Katis, Aristotle University of Thessaloniki, Department of Agriculture, Laboratory
of Plant Pathology, 541 24, Thessaloniki, Greece (Chapter 14)
Pavel Kindlmann, Faculty of Biological Sciences, University of South Bohemia and
Institute of Landscape Ecology, Czech Academy of Sciences, 37005 Ceské Budgjovice,
Czech Republic (Chapter 12)
A. Michael Klüken, Institute of Plant Protection and Plant Diseases, University of
Hannover, 30419 Hannover, Germany (Chapter 25)
Jonathan D. Knight, Centre for Environmental Policy, Imperial College London, Silwood
Park campus, Ascot, Berks, SL5 7PY, UK (Chapter 31)
Shinkichi Komazaki, Grape and Persimmon Research Station, National Institute of Fruit
Tree Science, NARO, Hiroshima, Japan (Chapters 29, 30)
Simon R. Leather, Division of Biology, Imperial College London, Silwood Park campus,
Ascot, Berks, SL5 7PY, UK (Chapter 6)
François Leclant (the late), Ecole Nationale Supérieure Agronomique, 34060 Montpellier
Cedex 5, France (Chapter 23)
Hugh D. Loxdale, Plant and Invertebrate Ecology Division, Rothamsted Research, Harpenden,
Gugs Lushai, Plant and Invertebrate Ecology Division, Rothamsted Research, Harpenden,
Manfred Mackauer, Department of Biological Sciences, Simon Fraser University, Burnaby,
B.C., V5A 1S6, Canada (Chapter 8)
Gerald J. Michels, Jr., Texas Agricultural Experiment Station, Bushland, TX 79012, USA
(Chapter 27)
List of Contributors xxvii
Xinzhi Ni, USDA-ARS Crop Genetics and Breeding Research Unit, Coastal Plain
Experiment Station, Tifton, GA 31793, USA (Chapter 13)
Judith K. Pell, Plant and Invertebrate Ecology Division, Rothamsted Research, Harpenden,
Herts, AL5 2JQ, UK (Chapters 8, 18)
Jan Pettersson, Department of Ecology, The Swedish University of Agricultural Sciences,
750 07 Uppsala, Sweden (Chapter 4)
John A. Pickett, Biological Chemistry Division, Rothamsted Research, Harpenden, Herts,
AL5 2JQ, UK (Chapter 9)
Manuel Plantegenest, INRA, UMR BiO3P, 35653 Le Rheu, France (Chapter 19)
Hans-Michael Poehling, Institute of Plant Protection and Plant Diseases, University of
Hannover, 30419 Hannover, Germany (Chapter 25)
Katherine L. Ponder, School of Biosciences, University of Birmingham, Edgbaston,
Birmingham, B15 2TT, UK (Chapter 11)
Glen Powell, Division of Biology, Imperial College London, Wye campus, Ashford, Kent,
TN25 5AH, UK (Chapter 14)
Wilf Powell, Plant and Invertebrate Ecology Division, Rothamsted Research, Harpenden,
Jeremy Pritchard, School of Biosciences, University of Birmingham, Edgbaston,
Birmingham, B15 2TT, UK (Chapter 11)
Sharron S. Quisenberry, College of Agriculture and Life Sciences, Virginia Tech,
Blacksburg, VA 24061, USA (Chapter 13)
Edward B. Radcliffe, Department of Entomology, University of Minnesota, St. Paul, MN
David W. Ragsdale, Department of Entomology, University of Minnesota, St. Paul, MN
Katherine A. Robinson, National Centre for Advanced Bio-Protection Technologies,
Lincoln University, Canterbury 7647, New Zealand (Chapter 16)
Mark Stevens, Broom’s Barn Research Station, Higham, Bury St. Edmunds, Suffolk, IP28
6NP, UK (Chapter 14)
Robert A. Surányi, McLaughlin Gormley King Company, 8810 Tenth Ave. N, Minneapolis,
MN 55427, USA (Chapter 26)
G. Mark Tatchell, Department of Biological Sciences, University of Warwick, Gibbet Hill
Road, Coventry, CV4 7AL, UK (Chapter 24)
Deborah J. Thackray, Centre for Legumes in Mediterranean Agriculture, MO80, Faculty
of Natural and Agricultural Sciences, University of Western Australia, 35 Stirling
Highway, Crawley, WA 6009, Australia and Plant Pathology, Department of Agriculture
and Food Western Australia, Locked Bag 4, Bentley Delivery Centre, WA 6983, Australia
(Chapter 31)
W. Fred Tjallingii, Laboratory of Entomology, Wageningen University, PO Box 8031,
Wageningen, The Netherlands (Chapter 4)
John A. Tsitsipis, University of Thessaly, Department of Agriculture, Crop Production and
Rural Environment, Laboratory of Entomology and Agricultural Zoology, 384 46, Nea
Ionia Magnissias, Greece (Chapter 14)
Jason M. Tylianakis, School of Biological Sciences, University of Canterbury, Private Bag
4800, Christchurch 8020, New Zealand (Chapter 16)
Maurice Vaissayre, Centre de Coopération Internationale en Recherche Agronomique pour
le Développement, Avenue Agropolis, 34398 Montpellier Cedex 5, France (Chapter 23)
Wolfgang Völkl, Department of Animal Ecology, University of Bayreuth, 95440 Bayreuth,
Germany (Chapter 8)
Susan E. Webb, Entomology and Nematology Department, University of Florida, Gainesville,
FL 32611, USA (Chapter 28)
Wolfgang W. Weisser, Institute of Ecology, Friedrich-Schiller-University, 07743 Jena,
Germany (Chapter 7)
xxviii List of Contributors
Iain S. Williams, Science Directorate, DEFRA, Nobel House, Smith Square, London, SW1P
3JH, UK (Chapter 3)
J. A. Trefor Woodford, Scottish Crop Research Institute, Invergowrie, Dundee, DD2 5DA,
UK (Chapter 22)
Steve D. Wratten, National Centre for Advanced Bio-Protection Technologies, Lincoln
University, Canterbury 7647, New Zealand (Chapter 16)
1 Taxonomic Issues
Roger L. Blackman and Victor F. Eastop

Department of Entomology, Natural History Museum, London, SW7 5BD, UK
Introduction All these insects are phytophagous, and most

of them are sap-sucking. Historically, the
There are about 4700 species of Aphididae Sternorrhyncha have been grouped with the
in the world (Remaudière and Remaudière, Auchenorrhycha (leafhoppers, cicadas, etc.)
1997). Of these, about 450 species have been as Homoptera, but molecular work has pro-
recorded from crop plants (Blackman and vided strong support for the long-standing
Eastop, 2000), but only about 100 have success- view, based on morphological and embryo-
fully exploited the agricultural environment logical evidence, that Sternorrhyncha and
to the extent that they are of significant eco- Auchenorrhyncha do not share a common
nomic importance (Table 1.1). The agricul- ancestor, so this grouping is not phylogen-
turally important species are mostly in the etically sound. The general consensus now
subfamily Aphidinae, not only because this is that the primary division of the Hemiptera
is the largest subfamily, but also because it is into Sternorrhyncha and a sister group
contains a very high proportion of the aphids comprising Auchenorrhyncha plus Heterop-
that feed on herbaceous plants (Blackman and tera (von Dohlen and Moran, 1995).
Eastop, 2006). Some quite large aphid sub- One major feature, their well-known
families – the Calaphidinae and Lachninae, cyclical parthenogenesis, sets aphids apart
for example – are associated almost exclu- from other Hemiptera, and has influenced
sively with woody plants, as are most of the every aspect of their biology. The system of
smaller ones. alternating one bisexual generation with a
The Aphididae is one of three families succession of parthenogenetic, all-female
of Aphidoidea, the other two being the Adel- generations evolved in the ancestral line of
gidae, or conifer woolly aphids, and the all Aphidoidea, probably as far back as the
Phylloxeridae, which are also nearly all asso- Triassic. At first, the parthenogenetic females
ciated with trees but include the notorious must have laid eggs like their sexual coun-
Viteus (= Daktulosphaira) vitifoliae (grape terparts, as do the parthenogenetic females
phylloxera) (Table 1.1). In the context of the of present-day adelgids and phylloxerids.
order Hemiptera as a whole, the superfamily Then, in the line leading to all modern Aphi-
Aphidoidea is placed in the sub-order Sterno- didae, the parthenogenetic generations devel-
rrhyncha, along with Coccoidea (scale insects oped the further refinement of viviparity,
and mealybugs), Aleyrodoidea (whiteflies) which effectively ‘telescoped’ generations into
and Psylloidea (psyllids or jumping plant lice). one another, thereby greatly reducing the
©CAB International 2007. Aphids as Crop Pests
(eds H. van Emden and R. Harrington) 1
2
Table 1.1. Biological features and distribution of agriculturally important species among the major groups of aphids.
No. of
agriculturally
Characteristic Geographical Host important Representative
Taxon host plants distribution alternation? spp./total spp. genera
Adelgidae Coniferae Holarctic Most 0/49 Adelges, Pineus
Phylloxeridae Fagaceae, Juglandaceae Holarctic (mostly nearctic) Some 1/64 Phylloxera, Viteus
Aphididae:
Eriosomatinae Ulmaceae, Salicaceae, Holarctic, oriental Most 9/319 Eriosoma, Pemphigus
Anacardiaceae, roots of
gymno- and angiosperms
Hormaphidinae Hamamelidaceae, Styracaceae, Mostly oriental Many or 5/176 Astegopteryx, Cerataphis,
palms, bamboos most Ceratovacuna
Anoeciinae Cornus, grass roots Holarctic Most 2/24 Anoecia
Calaphidinae Deciduous trees Holarctic None 6/331 Chromaphis, Myzocallis,
Therioaphis
Saltusaphidinae Sedges Holarctic None 0/68 Iziphya, Subsaltusaphis,
Thripsaphis
Chaitophorinae Salicaceae, Acer, grasses Holarctic None 2/164 Atheroides, Sipha
Greenideinae Dicotyledonous trees Eastern palaearctic, None 2/151 Anomalosiphon,
oriental Greenidea
Pterocommatinae Salicaceae Holarctic None 0/51 Pterocomma
Aphidinae:
Aphidini Some 20/738 Aphis, Rhopalosiphum,
R.L. Blackman and V.F. Eastop

Many families including Holarctic, a few southern Schizaphis, Toxoptera
Macrosiphini Rosaceae, Poaceae, hemisphere Some 60/2016 Acyrthosiphon,
Asteraceae Brevicoryne,
Macrosiphum, Myzus
Lachninae Trees (dicotyledonous Holarctic None 4/356 Cinara, Lachnus, Trama
and coniferous)
15 other Mostly dicotyledonous Some are oriental and None 0/185
subfamilies trees southern hemisphere
Taxonomic Issues 3
generation time and enabling far more effi- examples, we will discuss 14 taxa that prob-
cient exploitation of periods of rapid plant ably head the list of economically important
growth. aphid species.
Aphids (Aphididae) can be recognized
by a number of shared morphological char-
acteristics that developed before the diver- Interpreting Variation in Aphids
gence into present-day subfamilies: e.g.
siphunculi (secretory organs, but with their In the year 2000, soybean crops in the USA
precise function still strangely enigmatic); and Australia were attacked for the first time
five- or six-segmented antennae composed of by large numbers of an aphid closely resem-
two basal segments and a segmented flagel- bling the well-known polyphagous species,
lum with a terminal process; two-segmented Aphis gossypii (cotton or melon aphid), but
tarsi with the first segment much shorter than obviously with a far greater affinity for soy-
the second; and a cauda, or tail, that is often bean. Taxonomists identified the species as
used for flicking away droplets of honeydew Aphis glycines (soybean aphid), previously
from the anus. These features have been mod- known only from the Far East. It had been
ified, reduced, or secondarily lost in some introduced probably a year or two earlier,
species, but are evident in most aphids that building numbers and spreading until field
are pests of crop plants. entomologists and growers realized that they
Of the present-day subfamilies of Aphi- had something new on their hands. Aphis
didae, one in particular was successful at glycines is biologically quite different from
exploiting the rapid expansion of numbers A. gossypii; it is specific to soybean in sum-
and diversity of herbaceous flowering plants mer and has host alternation, overwintering
in the Tertiary period. This subfamily, the on Frangula spp. Identification immediately
Aphidinae, with 2750+ extant species, is unlocked much crucial information about
predominantly a northern temperate group, this species and its biology, as well as where
with life cycles closely tied to temperate to look for its natural enemies. In this case,
seasonality and the phenologies of temper- the question ‘What is it?’ seems to have a
ate plants. Originally on woody plants, they fairly clear-cut answer; the soybean aphid is
evolved a system of host alternation, migrating a relatively well-known and well-studied
to completely unrelated herbaceous plants for species in Eastern Asia. Although morpho-
the summer months, where their parthenoge- logically resembling A. gossypii and requir-
netic generations could continue to utilize ing specialist identification, it is clearly a
stages of rapid plant growth. However, today, distinct species. There are plenty of other
only about 15% of Aphidinae host-alternate. cases, however, including other close rela-
Some of the other 85% live only on woody tives of A. gossypii, where the question of
plants, but most of them, including some of identity is not so easily answered. Several
the largest and most successful genera, have of these are discussed in the next section of
lost or given up the ancestral woody (pri- this chapter. Sometimes, taxonomic diffi-
mary) host, and now live all-year-round on culties arise as a result of founder effects
herbaceous plants. Host alternation has, in and, even in the case of the soybean aphid,
fact, evolved independently in several other it may be necessary to bear in mind that the
aphid subfamilies (von Dohlen and Moran, introduced population has been through a
2000), but the Aphidinae are the only sub- recent ‘bottleneck’, and can be expected
family to exploit numerous families and to have less allelic diversity than East Asian
genera of flowering plants. populations.
In the rest of this chapter, we will look Taxonomy and identification are a mat-
at aphids from a taxonomist’s viewpoint. ter of interpreting observed variation. The
We will highlight some of the problems of first problem to be overcome is the effect of
interpreting the observed variation within the environment on the phenotype, especially
and between species, and discuss what the if the only available data are morphological.
names we give to aphids really mean. Then, as Most species are distinguished and described
4 R.L. Blackman and V.F. Eastop
originally, often from one small sample only, on the length relationships between body
using morphological criteria, and most iden- parts (allometry), which can play havoc with
tifications are based on keys that use morpho- the morphometric ratios and functions often
logical discriminants. As a group, aphids are used to discriminate between closely related
renowned for the considerable extent to which species (Blackman and Spence, 1994).
the phenotype is influenced by environmen- Thus, it is important to take into acc-
tal factors. Within any aphid species, there ount the possible effects of season, host plant,
are a number of different forms (morphs) with and climate when examining samples of
discrete morphological differences, which field-collected aphids. It is also necessary to
may be triggered by specific environmental bear in mind that a sample might consist of
stimuli such as day length or crowding. It is a single clone, especially if it comes from a
well known, for example, that in most aphids, warm temperate or subtropical region where
the parthenogenetic females can be winged the population has not gone through a sex-
or wingless, the differences being not just in ual phase and is therefore more likely to be
the presence or absence of wings but invol- clonally structured. All the aphids in a
ving every part of the body. Separate identi- clone are, discounting mutations, geneti-
fication keys are therefore needed for each cally identical, so a sample consisting of a
morph, but the structural distinction between single clone will give a misleading idea of
the winged and wingless morphs is not always the range of variation in the species. Some-
as clear as might be expected, as wingless times ‘abnormal’ morphological features
individuals may occur with some tendency may occur, which may be due to mutation,
towards the characters of the winged morph. but are often no more than character states
Likewise, intermediates between other morphs near the extremes of a range of continuous
can occur; for example, between viviparous variation, and part of the natural variation
parthenogenetic females and oviparous sex- of all living organisms. In sexually repro-
ual females. ducing organisms, any particular abnormal
The range of continuous morphological character state is likely to occur in only the
variation is also wider than in many other occasional, rare individual within a popula-
insect groups. Increases or decreases of size tion, and thus will be instantly recognizable
due to nutritional effects, for example, can as an extreme or rare condition. In aphids, it
accumulate over several generations, because is possible to find a whole colony of indi-
the size of the mother can affect the size of viduals all with the same anomalous char-
her offspring. There may be large seasonal acters. Such a colony has sometimes been
differences, with some species producing erroneously described as a new species.
dwarf individuals when food quality is poor
in midsummer. The parthenogenetic female
that hatches from the overwintering egg – The Taxonomy of Pest Aphids –
the fundatrix or stem mother – is morpho- What’s in a Name?
logically different from the later generations
of parthenogenetic females, and sometimes The species that become pests are those that
the particular features of the fundatrix are only are best able to adapt to and exploit man-
gradually lost in subsequent generations. In modified environments. Many of them belong
some aphid groups – although not in most to groups that were probably already specia-
Aphidinae – the generations on the secondary ting rapidly before human intervention, and
host differ greatly in morphology from those on the colonization of new geographical regions
the primary host, and were originally descri- and/or new habitats are potent factors lead-
bed as different species and often placed in ing to further divergence and change. Sev-
different genera. The temperature experi- eral examples of this are discussed in the
enced during development can also have concluding part of this chapter.
profound effects on morphology, involving Although it is clear that pest aphids are
not only general body size and pigmenta- highly dynamic, rapidly evolving systems,
tion, but also having more subtle influences there still seems to be a tendency, among
Taxonomic Issues 5
agricultural entomologists in particular, to new taxa in animal taxonomy. It was devel-

regard pest species identifications provided oped mainly by vertebrate taxonomists, who
by taxonomists as names set in stone, and to define subspecies as geographically local-
react with alarm to the suggestion that cer- ized populations that are morphologically
tain common and well-known pest species distinguishable. In aphid taxonomy, consid-
may, in fact, be more accurately described erable use has been made of the subspecies
as species complexes. The name has great category. Remaudière and Remaudière (1997)
importance, as it is the key to what we know list 141 accepted subspecies names in the
about a species and its way of life, and to subfamily Aphidinae alone. Unfortunately,
how we might expect it to behave. The name the subspecies designation has been used with
Myzus persicae, for example, identifies a set an almost complete lack of consistency. In
of populations that are closely related and some cases, a single sample has been described
share numerous attributes. But this must as a subspecies, for no other reason than that
not be allowed to mask the heterogeneity that it shows some deviation from the known
is also present, and which may in particular range of variation of a species. This could be
include populations that have diverged due simply to it comprising a single clone
genetically to such an extent that they have with certain anomalous features, or a colony
evolved past the stage of being host races that developed under unusual microclimatic
and achieved a degree of permanency, so conditions, or part of a continuous geograph-
that they can be regarded as incipient or sib- ical cline of variation from which intermedi-
ling species (or subspecies) with particular ate populations have not been sampled.
attributes of their own. Aphid taxonomists studying groups of
The recognition of such divergent pop- very closely related taxa (‘species com-
ulations or taxa can add to our understand- plexes’) have also used subspecies in a com-
ing of the ecology and evolution of a pest pletely different way to define populations
species complex, and increase the possibil- that are morphologically very similar but
ity of devising effective control measures. It that have been shown by field observation
is also important that they are named in and/or experimental studies to differ in
some way, because without any consistent their life cycle or host-plant relationships
method of referring to them, new informa- (Müller, 1986). Such an approach recognizes
tion is inaccessible, lost in the mass of liter- that speciation processes in aphids may be
ature about the species as a whole. Yet, both very different from those of vertebrates,
the recognition and the naming of such cat- with changes in life cycle and/or host rela-
egories are fraught with difficulty. Discrimi- tions acting as the primary isolating mecha-
nation of recently derived taxa may require nism and trigger for speciation, rather than
sophisticated techniques, which are likely spatial isolation (Guldemond and Mackenzie,
to be time-consuming and/or expensive. It is 1994). Crucial evidence that such speciation
also hard to decide what to call them. Insect processes may operate in aphids has been
taxonomists are usually reluctant to provide provided recently by Hawthorne and Via
formal names and descriptions and to give (2001). Thus, in rapidly speciating groups
full species status to members of species com- of aphids, one may expect to find incipient
plexes that cannot be distinguished readily species that are at an early stage of divergence
by their morphology alone, even where con- where they can be distinguished more easily
sistent differences can be demonstrated in by biological rather than by morphological
biology and/or host relationships. This is properties.
understandable, given that they probably Every speciation process is a unique event,
will be asked to recognize and identify dead or series of events, which may include tran-
specimens of their ‘new’ species! sitional phases involving a wide variety of
The subspecies category is the only intra- reproductive relationships between the incip-
specific category recognized by the Zoologi- ient species. It is impractical to try to define
cal Code of Nomenclature, and therefore the such transitional phases except in the most
only one that can be used formally to designate flexible terms. Blackman (1995) suggested
that the term ‘semispecies’ could convey this recognizable entities which show certain
idea of ‘species in the making’, without imply- consistent distinguishing properties, indi-
ing anything about the nature of the specia- cating an interruption or significant restric-
tion process or the degree of reproductive tion of gene flow and making it reasonable
isolation. Semispecies have been used already to conclude that speciation is in progress, but
for many years by both plant and animal has not reached the stage of irreversibility.
taxonomists with experience of the problems A working definition of an aphid sub-
of describing species complexes, including species. A group of populations that is rec-
some of the best-studied ones such as those ognizably part of an existing species yet
of the fruit fly, Drosophila. They have not, maintains a consistent suite of properties
however, been used for aphids, or most distinguishing it from other populations
other insect groups of agricultural impor- within that species. The species should be
tance, nor have they been accommodated sufficiently well known for it to be reason-
into the Zoological Code of Nomenclature. ably certain that the observed variation is
On further reflection, it seems unrealistic discontinuous, and the consistency of this
to expect the wide acceptance and use in discontinuity should be demonstrated by
aphid taxonomy or agricultural entomology samples from more than one time and place.
of the term semispecies and, even if it were The likely cause of the discontinuity should
accepted, that would still not resolve the be identifiable, for example a difference in
problem of nomenclature. host-plant relationships, life cycle proper-
Rakauskas (2004) has recently revived ties, or geographic location, and should not
Müller’s (1986) proposal for a broader use be of such a kind as to make it irreversible
of the subspecies category to validate its (e.g. permanent parthenogenesis). The des-
application to aphid species complexes, cription of a new subspecies should give a
and – of particular relevance to the present clear and accurate account of its morphologi-
chapter – to meet the practical need of pro- cal and biological properties in comparison
viding names that identify intraspecific cat- with those of other populations within the
egories in groups that include pest species. species, and include the best possible
We support this idea and accommodate it in morphological discriminants. One clone
our discussion of the taxonomy of some of should be designated as the type, with slide-
the major pest aphids that occupies the mounted specimens deposited in a national
remainder of this chapter. collection and other specimens of the same
Blackman (1995) drew attention to the clone deep-frozen and/or preserved in abso-
extensive misuse of the term ‘species concept’ lute ethanol for future DNA studies.
in the literature on evolution and speciation,
which has made it difficult for taxonomists
and evolutionists working in different
fields and on different groups of organisms The 14 Aphid Species of Most
to agree about the nature of species. The Agricultural Importance
same difficulties have also impeded the use (See also colour plates in this volume.)
of the subspecies category. To emphasize the
distinction between a concept and a working There is a very large literature about all the
definition in the case of subspecies, we offer major pest aphid species. Much of the recent
below (i) a description of the ‘subspecies work is readily accessible on the Internet,
concept’ that is applicable to all organisms and summary accounts are provided by
irrespective of their modes of speciation, Blackman and Eastop (2000). The treatments
and (ii) a working definition for the subspe- that follow are therefore largely concerned
cies category in aphids. with the taxonomic issues raised by these
The subspecies as a concept. Subspecies 14 species, and the two factors that have the
can be conceived as a species in the making; greatest influence on intraspecific variation
populations or groups of populations that in aphids, the life cycle and the host plant.
have diverged to the extent that they are These factors are considered further in the
Taxonomic Issues 7
next chapter from a population geneticist’s sexual phase on Vicia. Recent molecular
viewpoint. work has shown that populations coloniz-
ing peas, lucerne, and red clover in France
are genetically divergent and also differ in
Acyrthosiphon pisum (pea aphid) their symbionts (Simon et al., 2003).
There seem to be several other specia-
Acyrthosiphon pisum is a rather large, green tion events currently in progress in Europe,
or pink aphid with long, slender appendages, with populations also in the process of
forming colonies on young growth and devel- diverging on Lotus and Sarothamnus. A
oping pods of many leguminous plants. Its form that lives on Ononis in Europe is mor-
host plants are mostly Fabaceae of the the phologically recognizable and is currently
tribes Genistae (Cytisus, Genista, Sarothamnus, regarded as a subspecies (A. pisum ononis),
Spartium), Trifoliae (Medicago, Melilotus, but could be a distinct species.
Ononis, Trifolium, Trigonella), Fabeae (Lathy- Populations introduced to other parts
rus, Lens, Pisum, Vicia), and Hedysareae of the world must at least originally have
(Hippocrepis, Onobrychis), and it also colo- been genetically depauperate, and hence
nizes a few members of other tribes, e.g. Lotus their biology and host-plant relations may
(Loteae) and Glycine (Phaseolae). Many have diverged significantly from the species
legumes, including some of economic impor- in Europe. The genotype(s) introduced to
tance (e.g. Phaseolus), and almost all other North America initially lacked the dominant
plants are not colonized, although under dry red (pink) allele that is frequent in European
conditions it is sometimes found on Capsella populations, and were particularly well adap-
bursa-pastoris. It is a vector of more than 30 ted to lucerne. They were named as a new
virus diseases, including non-persistent viruses species (Johnson, 1900), and this name
of beans, peas, beet, clover, cucurbits, Nar- (destructor) has been proposed as a sub-
cissus, and Brassicaceae, and the persistent specific name not only for the North Ameri-
viruses, Pea enation mosaic virus (PEMV) and can populations (Hille Ris Lambers, 1947),
Bean leaf roll virus (BLRV). but also for the green, pea-adapted form in
Originally a palaearctic species, A. pisum Europe from which it was assumed to have
now has an almost worldwide distribution been derived (Müller, 1985; but see below).
(see CIE Distribution Map 23, last revised Variation within and between North American
1982). In cold temperate regions, it is populations has been studied with respect
holocyclic, producing oviparae and males on to resistance in lucerne (Frazer, 1972) and
various leguminous hosts. As in other members pea (Cartier, 1963) varieties, life-history traits
of the genus Acyrthosiphon, there is no true (MacKay et al., 1993), photoperiodic respon-
host alternation. The ancestral primary host ses (Smith and MacKay, 1990) and migra-
was presumably a member of the Rosaceae, as tory tendencies (Lamb and MacKay, 1979).
in related genera, but this was probably lost Harper et al. (1978) provided a bibliography
long ago. Clones may produce either apterous of all the earlier work on A. pisum.
or alate males, or both. At warmer latitudes, In ground-breaking work that has illu-
it overwinters without a sexual phase. minated the evolutionary processes that may
In Europe and Central Asia, A. pisum be going on in crop-colonizing aphids, Via
seems to be a complex of races and subspe- (1991, 1999) demonstrated that populations
cies with different host ranges and prefer- of A. pisum on lucerne and red clover in
ences (Müller, 1980, 1985). Populations north-eastern USA perform significantly
attacking peas (Pisum sativum) in Europe better on their respective host plants, and are
consist entirely of green genotypes and reproductively isolated from one another as a
have some morphological differences from result of inherited differences in host selec-
those colonizing other leguminous crops such tion by alatae, which lead to assortative
as lucerne (Medicago sativa), which may be mating. Hawthorne and Via (2001) have
green or pink. They produce alate sexuparae shown recently that there is close genetic
and males in autumn, and go through a linkage at several loci between the two key
traits involved in host specialization, i.e. Aphis craccivora (cowpea aphid)

host selection by alatae, and subsequent
performance (measured by fecundity) of This is a small, dark brown aphid with a
populations on each host. This could lead shiny black dorsal shield. It occurs most com-
to extremely rapid divergent selection of pea monly on legumes, but is much more poly-
aphids on lucerne and clover. phagous than A. pisum, with a wide range of
The implications of this work are far- hosts not only in the Fabaceae (e.g. Arachis,
reaching, although much is still unclear. Colutea, Glycine, Medicago, Melilotus, Trifoli-
The origins of the populations studied by um, Vicia), but also in many other plant
Via’s group are unknown, but subsequent families (altogether it attacks about 50 crops
work in Europe by Simon et al. (2003) sug- in 19 different plant families). It is a vector of
gests that the observed differences may be about 30 plant virus diseases, including non-
long-standing and the result of separate persistent viruses of beans, cardamom (Elett-
introductions from the palaearctic of two aria cardamomum), groundnuts, peas, beets,
genotypes with different host associations, cucurbits and crucifers, and the persistent
as seems to have happened with some other Subterranean clover stunt virus, Peanut mottle
pest aphids in North America (see under virus, and the complex of viruses causing
Therioaphis trifolii, Schizaphis graminum, groundnut rosette disease. Jones (1967) com-
and Myzus persicae). More detailed genetic pared two clones from East and West Africa
comparisons of European and North Ameri- differing in ability to transmit strains of
can populations are needed to confirm this. groundnut rosette and found that there were
However, regardless of their origins, Via’s also differences in ability to colonize various
A. pisum populations seem to have a genetic host plants. Apart from this, there has been
system that can lead to rapid divergence little work of any significance on intraspe-
and incipient speciation in crop environ- cific variation in Aphis craccivora.
ments. Add in the probability that the same Aphis craccivora now occurs in most
processes of selection and reinforcement of parts of the world (CIE Distribution Map 99,
correlations between key traits are occur- revised 1983), but its origins are clearly in
ring in large populations throughout the Europe, as the most polyphagous member
regions where these crops are grown and of a group of closely related species (subge-
there seems to be a very potent mechanism nus Pergandeida), most of which are spe-
for evolutionary change. cific to particular species of Fabaceae. None
If the host-adapted forms of A. pisum of this group of Aphis has host alternation.
demonstrated by Via’s and Simon’s groups Aphis craccivora has a sexual phase on var-
are indeed ‘incipient species’, can they be ious Fabaceae in Central Europe (Germany),
classed as subspecies? It would obviously and sexual morphs have also been reported
be undesirable to give names to locally from India and Argentina, but through most
divergent populations, even though the first of the world, reproduction seems to be exc-
stage in the speciation process can occur lusively parthenogenetic. It is particularly
only at the level of the local population. To common as a pest in warmer climates, and
conform with the working definition of it seems likely that the pest populations may
aphid subspecies above, it would be neces- have originated from the warmer part of its
sary to demonstrate that the observed differ- original distribution area in Southern Europe
ences were consistent in both time and or the Middle East.
space, and verify the existence of monophy-
letic host-adapted lineages. It is already
apparent that the name used originally for Aphis fabae (black bean aphid)
the North American lucerne-feeding form
(destructor) was applied wrongly to the Aphis fabae is perhaps the most familiar
pea-feeding populations in Europe, although aphid in Europe, due to its predilection for
it still may be valid for a lucerne-adapted Phaseolus and Vicia, although it is probably
clade if this is common to both continents. just as important as a pest and virus vector
Taxonomic Issues 9
of sugarbeet. Adult apterae in new colonies on colonize certain other plants, but is not
young plant growth are matt black in life, most found on Vicia or Solanum.
of the pigmentation of the body being internal, 4. Aphis euonymi, a brown aphid that
so that it is no longer present in cleared spec- stays on Euonymus all year round.
imens prepared for the microscope. Individ- 5. Aphis fabae mordvilkoi, again almost
uals in older colonies and feeding on leaves indistinguishable from A. fabae sensu stricto,
tend to develop white wax markings. but with sexual generations on Viburnum
Aphis fabae is a very polyphagous spe- opulus or Philadelphus coronarius, and mig-
cies, but the actual host range of the aphid rating for the summer mainly to various
that colonizes beans and sugarbeet is unclear, secondary hosts, but not colonizing Vicia,
because it is a member of a bewildering com- Solanum, or Cirsium, and often occurring
plex of species, at least some of which also on Arctium spp. and Tropaeolum majus.
have wide host ranges. Many people have
tried to sort out this complex. Stroyan (1984: However, the host ranges of all members of
119–122) reviewed the taxonomy and host the group seem to overlap and some plants,
relations of the group as then understood. such as Rumex obtusifolius, seem to be accep-
There has been significant work since then ted by them all (Thieme, 1987, 1988).
(e.g. Müller and Steiner, 1986; Thieme, 1987, To complicate the story still further,
1988; Thieme and Dixon, 1996; Raymond the primary host relationships of the mem-
et al., 2001), but a lot of questions still remain. bers of the group are also not at all clear-cut.
To summarize the situation in Northern Aphis fabae sensu stricto and A. solanella
Europe as succinctly as possible, there are are able to live on Viburnum and Philadel-
five closely related taxa, four of which go phus, the primary hosts of A. f. cirsiiacan-
through their sexual phase on the spindle thoidis and A. f. mordvilkoi, and may in
tree, Euonymus europaeus: autumn produce males on these plants, but
not oviparae (Iglisch, 1968). Hybridization
1. Aphis fabae sensu stricto (A. fabae ssp. between any members of the group is there-
fabae), which has Euonymus as its only pri- fore theoretically possible, and can be accom-
mary host and migrates for the summer to a plished rather easily in the laboratory, even
wide range of plants, including Vicia faba, between slightly less closely related forms
sugarbeet, Chenopodium album, and pop- such as A. fabae and the brown species, A.
pies, but not Solanum nigrum. euonymi (Müller, 1982). However, it is now
2. Aphis solanella, which also has Euony- clear that such results are misleading, and
mus as its only primary host and migrates to that strong prezygotic isolating mechanisms
a wide range of plants including S. nigrum, are likely to be operating to prevent hybrid-
the leaves of which it crumples and curls ization and promote assortative mating in
characteristically, but will not colonize beans, the field. Aphis fabae sensu stricto and A.
sugarbeet, Chenopodium album, or poppies. solanella occurring together on Euonymus
Aphis solanella, which has shorter hairs show differences in the diurnal patterns of
than A. fabae and a greater tolerance of high pheromone release by oviparae and male
temperatures, has usually been classed as a responsiveness, and in choice experiments,
subspecies of A. fabae, but Thieme and Dixon males preferred the sex pheromones of con-
(2004) suggested that it should have full specific females (Thieme and Dixon, 1996).
species status. It is a pest in its own right, Prezygotic isolating mechanisms have also
especially on Solanaceae. been demonstrated in the laboratory between
3. Aphis fabae cirsiiacanthoidis, which A. fabae sensu stricto and A. f. mordvilkoi,
normally uses Euonymus as a primary host although these subspecies do not share the
but can also go through its sexual phase on same primary host in nature (Raymond et al.,
Viburnum opulus. This form is morphologi- 2001).
cally hardly distinguishable from A. fabae In spite of this evidence of substan-
sensu stricto, but has Cirsium arvense as tial reproductive isolation between the mem-
its most characteristic summer host. It can bers of the A. fabae complex, analysis of
mitochondrial DNA and of a plasmid from cocoa, aubergine, peppers, potato, okra, and
their symbionts showed differences within, many ornamental plants including chry-
but not between, species (Raymond et al., santhemums and Hibiscus. Populations on
2001). This is similar to the situation in A. cotton and cucurbits can be particularly large
pisum already discussed (where host/habi- and damaging. More than 50 plant viruses are
tat selection rather than mate selection is transmitted, including non-persistent viruses
the isolating factor), and suggests that such of beans and peas, crucifers, celery, cowpea,
isolating mechanisms may evolve very rap- cucurbits, Dahlia, lettuce, onion, pawpaw,
idly, possibly by the process known as rein- peppers, soybean, strawberry, sweet potato,
forcement (Coyne and Orr, 1989; Mackenzie tobacco, and tulips, and the persistent Cotton
and Guldemond, 1994). anthocyanosis virus, Lily symptomless virus,
On the evidence of its primary host and PEMV, and lily rosette disease.
closest relatives, A. fabae must be of Euro- In all the warmer parts of the world, A.
pean origin. Aphis fabae sensu stricto gossypii reproduces continuously by parthe-
occurs in Europe, Western Asia, Africa, and nogenesis. It is particularly abundant and
South America. It is a vector of more than widely distributed in the tropics, including
30 plant viruses, including non-persistent many Pacific islands. During prolonged dry
viruses of beans and peas, beets, crucifers, seasons in hot countries, small colonies
cucurbits, Dahlia, potato, tobacco, tomato, may survive on a great variety of plants on
and tulip, and the persistent Beet yellow net which they are seldom seen during the grow-
virus (BYNV) and Potato leaf roll virus ing season, including Poaceae. Deguine and
(PLRV). In warmer regions – the Mediterra- Leclant (1997) provided a comprehensive
nean and the Middle East, the Indian sub- account with an extensive bibliography.
continent, and hotter parts of Africa and Certain morphological features – short
South America – it is replaced by A. hairs on legs and antennae, and a cauda that
solanella, reproducing parthenogenetically is usually paler than the siphunculi and
throughout the year on its secondary host bears rather few hairs – make it easy to apply
plants, particularly Solanaceae, Asteraceae, the name A. gossypii to aphids collected on
and Polygonaceae. The Tropaeolum-feeding crops or other non-indigenous plants any-
subspecies A. f. mordvilkoi has a more where in the world. This is, however, a con-
northerly, holarctic distribution, and uses siderable oversimplification of the taxonomic
Viburnum trilobum as its main primary host problem, as becomes evident, for example,
in Canada (Barber and Robinson, 1980; as when one compares accounts of A. gossypii
A. barbarae). in Europe and East Asia.
In Europe (Stroyan, 1984; Heie, 1986), A.
gossypii is classed as a subspecies in the Aphis
Aphis gossypii (cotton or melon aphid) frangulae complex, a group of closely related
and morphologically almost indistinguish-
If the multiplicity of populations that are able indigenous European species that use
lumped under the name A. gossypii are buckthorn (Frangula alnus) as their primary
really all one species, then it is indeed a host. Aphis gossypii is regarded (that is, more
remarkable one, with greater diversity in or less defined) as the only member of the
terms of host relationships, life cycle, and group that does not have a sexual phase on
geographical range than any other aphid. buckthorn, overwintering parthenogenetically
Small aphids that vary greatly in colour from in Northern Europe in protected situations
pale yellow dwarfs at high temperatures, such as glasshouses. One might conclude from
through dirty yellow-green to dark bluish- this scenario that this worldwide pest origi-
green or almost black at lower temperatures, nated in Europe as a permanently partheno-
occur on plants in numerous families, genetic, highly polyphagous and adaptable
including nearly a hundred species of crop offshoot of the A. frangulae complex, and
plants, throughout the world. Crops attac- spread from there to all parts of the world as
ked include cotton, cucurbits, citrus, coffee, the classic ‘general purpose genotype’.
Taxonomic Issues 11
However, such a conclusion is difficult Particular host associations have been noted
or impossible to reconcile with accounts of previously in A. gossypii, for example, in
what is purportedly the same species in European glasshouses, where aphids from
Japan and China, where the parthenogenetic chrysanthemums will not colonize cucum-
generations seem equally polyphagous, but ber, and vice versa (Guldemond et al., 1994).
there is an annual sexual phase. Overwin- These glasshouse populations on both chry-
tering as eggs occurs in East Asia on a vari- santhemums and cucurbits are normally par-
ety of unrelated plants, including Frangula thenogenetic, but can produce sexual morphs
spp., but also Hibiscus syriacus, Celastrus under certain conditions (Guldemond et al.,
orbiculatus, and Rubia cordifolia (Inaizumi, 1994 and Fuller et al., 1999, respectively).
1980; Zhang and Zhong, 1990). It is possible The work of Vanlerberghe-Masutti and Chavi-
that some of these populations have diverged gny is the first to show that forms of A.
as a result of differential selection among gossypii with particular host associations
these primary hosts; populations overwin- may be distributed over a wide area. One
tering on R. cordifolia in Japan, for exam- may hypothesize that two forms have become
ple, seem to be isolated from those on other widely distributed, possibly with different
primary hosts, and are possibly a separate geographic origins, both with rare sexual
taxon (Inaizumi, 1981). Earlier, Kring (1959) reproduction, one found particularly on
had demonstrated that populations in Con- Cucurbitaceae and Malvaceae (and causing a
necticut, USA, also have a sexual phase, major pest problem on cotton), and the other
using H. syriacus and also Catalpa bigno- specializing on Asteraceae. These may corre-
nioides as primary hosts. spond to the two forms, one cucurbit-feeding
So, did the worldwide pest called A. and the other chrysanthemum-feeding, iden-
gossypii originate in Europe, East Asia, or tified in European glasshouses. Further evi-
North America? North America is unlikely dence of the existence of a distinct, widely
to be the ancestral homeland, because there distributed Asteraceae-feeding form of A.
are no indigenous North American Aphis spe- gossypii has come recently from a multi-
cies of the group of species closely related to variate morphometric study (Margarito-
A. gossypii that use Frangula as primary poulos et al., 2006). However, there were no
hosts. In East Asia, there are indigenous consistent host-associated RAPD bands in
species related to A. gossypii that have a Japanese A. gossypii populations studied by
sexual phase on Frangula, such as the soybean Komazaki and Osakabe (1998), where there is
aphid A. glycines, but none of these seem regular genetic recombination. The problems
quite so similar in morphology to A. gossy- of identity and origin of A. gossypii and the
pii as the European A. frangulae group. The taxonomic status of host-associated forms,
answer to this enigma can only come from thus still remain, and require investigation
some extensive work encompassing the of the much more complex genetics of pop-
entire geographical and host-plant range of ulations in places where A. gossypii and its
A. gossypii and including comparisons with relatives have an annual sexual phase.
related species in Europe and East Asia.
Vanlerberghe-Masutti and Chavigny
(1998) provided some interesting pointers Aphis spiraecola (green citrus aphid
for further work, with a study of random or spiraea aphid)
amplified polymorphic DNA (RAPD) of 18
A. gossypii populations from Southern This is a small yellow or greenish-yellow
Europe, La Réunion, and Laos. They found aphid with black siphunculi and cauda,
certain RAPD bands that were fixed in popu- found in dense, ant-attended colonies, curl-
lations collected from cucurbits and absent ing and distorting leaves near the stem api-
from those collected on non-cucurbit hosts, ces of a wide range of plants, particularly
and the 18 populations therefore clustered those of shrubby habit. Its numerous hosts
into two groups according to host plant, are in more than 20 plant families, espe-
irrespective of their geographical origins. cially Caprifoliaceae, Asteraceae, Rosaceae,
Rubiaceae, Rutaceae, and Apiaceae. Proba- Compared with the problems raised by
bly, its most important crop host is Citrus. A. fabae and A. gossypii, the taxonomic sta-
Although not particularly efficient at trans- tus, origins, and identity of A. spiraecola
mitting viruses, very large populations occur seemed to be fairly clear. However, Komazaki
in spring in some regions – the Middle East, et al. (1979) found that A. spiraecola in
for example – and can make it an important Japan was using Citrus unshiu as well as
vector of Citrus tristeza virus. It also transmits Spiraea thunbergii as a primary host, and
Cucumber mosaic virus (CMV), Plum pox experimental work demonstrated that there
virus, an isolate of Alfalfa mosaic virus from were genetically inherited differences in
Viburnum, Water melon mosaic virus 2, hatching time (Komazaki, 1983, 1986) and
and Zucchini yellow mosaic virus (ZYMV). egg diapause (Komazaki, 1998), correlated
Now almost worldwide, there seems with esterase differences (Komazaki, 1991),
little doubt that A. spiraecola is indigenous that seem to indicate a degree of genetic iso-
to East Asia. It has been in North America since lation and divergence between the popula-
at least 1907, and had reached Australia by tions on the two primary hosts. Presumably,
1926, New Zealand by 1931, Argentina by the ancestral primary host was the rosaceous
1939, the Mediterranean region by about plant (Spiraea), and Citrus unshiu was acqui-
1939, and Africa by 1961. Populations in red more recently as a primary host. This
most parts of the world are permanently seems to be an example of incipient specia-
parthenogenetic on secondary hosts, but in tion, as in Via’s populations of A. pisum, but
East Asia and North America, A. spiraecola again it is not yet clear whether it is more
has a sexual phase on Spiraea. than a local phenomenon. There are no
There is an extensive literature on A. records of sexual generations on Citrus out-
spiraecola, particularly in relation to its eco- side Japan, and it could be that they are
nomic importance on Citrus (it was referred unable to survive on other Citrus species. In
to as A. citricola in the literature from 1975– Japan, alatae migrating from spring popula-
1988 because of a misidentification). The tions that had developed from overwinte-
most comprehensive accounts of A. spira- ring eggs on Citrus unshiu seemed to be the
ecola as a Citrus pest are by Barbagallo (1966) most important source of infestations of Cit-
in Italy and Miller (1929) in Florida. On other rus groves at a time when the main migra-
plants, especially Rosaceae, A. spiraecola is tion from Spiraea had already taken place
often confused with Aphis pomi (green apple (Komazaki, 1983).
aphid). For example, Cottier (1953) wrote
an account of A. spiraecola in New Zealand
under the name A. pomi, and Singh and Diuraphis noxia (Russian wheat aphid)
Rhomberg (1984) studied allozyme varia-
tion in populations nominally of A. pomi This small, narrow-bodied, yellow-green
on apples in North America and found two aphid was little known outside southern
forms, one of which was almost certainly A. Russia until the late 1970s. It then took only a
spiraecola. Aphis pomi has a longer last little over 10 years to colonize the main
rostral segment than A. spiraecola, more hairs wheat- and barley-growing areas of East Asia,
on the cauda, and usually has lateral tuber- South Africa, and both North and South
cles on abdominal segments 2–4 (see also America (see IIE Distribution Map 521, 1991).
Halbert and Voegtlin, 1992). There also is It is still expanding its range northward in
possible confusion of identity with Aphis Europe (Thieme et al., 2001). Hughes and
eugeniae in East and South-east Asia and Maywald (1990) assessed the suitability of the
Australia that can occur on the same hosts; Australian environment for D. noxia, although
A. eugeniae can be recognized by the it still (as at 2006) has not reached Australia.
peg-like hairs on the hind tibia, and by the Diuraphis noxia feeds only on Poaceae,
presence of a median sense peg between concentrating particularly on wheat and
the pair of hairs on the first segment of the barley. It does best on late-sown crops on
hind tarsus. poor soils. It transmits Barley yellow dwarf
Taxonomic Issues 13
virus, but its feeding also has a rapidly toxic in Ukraine, and Tanigoshi et al. (1995) descri-
effect on the plant, the leaves of which bed biological control measures. Chen and
become rolled into tubes and desiccated, Hopper (1997) studied its population dyna-
and infested ears become bent. In cold tem- mics and the impact of natural enemies in
perate regions of Europe and Asia, it has a southern France.
sexual phase without host alternation on
wheat and barley. In North America, it has
been assumed generally to have no sexual Lipaphis pseudobrassicae (mustard
phase. Kiriac et al. (1990) found some ovipa- aphid, also known as the false
rae in Idaho and Oregon, and oviparae have cabbage aphid)
appeared in glasshouse cultures (Puterka
et al., 1992) but no males have been found. Lipaphis pseudobrassicae is a cosmopolitan
Puterka et al. (1993) found some genetic pest of cruciferous crops. Apterae are small to
variation in North American D. noxia using medium-sized, yellowish, grey, or olive green,
RAPD-PCR, although this was between, with a waxy bloom that, in humid condi-
rather than within, populations, suggesting tions, becomes a dense coat of white wax. It
that it was generated only by rare mutatio- can occur in large colonies on the under-
nal or recombinational events. More genetic sides of leaves or in inflorescences of many
variation was found in collections from the species and genera of Brassicaceae, includ-
Middle East, Moldavia, Ukraine, and ing Barbarea, Brassica, Capsella, Erysimum,
Kirghizia. Their data suggest a single source Iberis, Lepidium, Matthiola, Nasturtium,
of spread of D. noxia, perhaps via Turkey to Raphanus, Rorippa, Sinapis, Sisymbrium,
France, South Africa, and North America. and Thlaspi. Often, the leaves are curled and
An interesting addition to this story is that turn yellow. It is a vector of about 10 non-
Mimeur (1942) collected D. noxia in North persisent viruses, including Turnip mosaic
Africa in 1938, described it as a new species virus and Cauliflower mosaic virus. It occurs
(Cavahyalopterus graminearum), and produ- throughout the world (CIE Distribution Map
ced oviparae in culture, but no males. Thus, 203, 1965), but particularly is a pest in
the absence of males may be a long-standing warmer climates, reproducing throughout
feature of the population, which has since the year by continuous parthenogenesis.
become widely distributed. The origin and identity of L. pseudo-
There are no problems with the iden- brassicae were long in doubt. In North
tity of D. noxia, although in Europe there is America, it was at first confused with Brevi-
a very similar species, Diuraphis muehlei, coryne brassicae, until Davis (1914) recog-
which feeds specifically on Phleum pratense, nized it as distinct and named it Aphis
turning the leaves yellow. This species has pseudobrassicae. Because of its weakly cla-
a shorter antennal terminal process than vate siphunculi, it was subsequently trans-
D. noxia; the ratio of the terminal process to ferred by Takahashi (1923) to the genus
the base of the last segment in apterae is Rhopalosiphum, and it was referred to in
1.05–1.65 (muehlei), as opposed to 1.55–2.6 the economic literature as Rhopalosiphum
(noxia), and in alatae 1.2–1.9 (muehlei), as pseudobrassicae (Davis) until 1964. Börner
opposed to 1.8–2.7 (noxia). and Schilder (1932) recognized that pseudo-
There is an extensive older Russian lit- brassicae should be placed in Lipaphis, a
erature on D. noxia, one of the most com- genus erected by Mordvilko (1928) for a
prehensive studies being that of Grossheim palaearctic crucifer-feeding aphid, erysimi.
(1914). The rapid spread and great economic Lipaphis erysimi is a holocyclic species
importance of this aphid have resulted in with a 2n = 10 karyotype (Gut, 1976; Black-
more recent extensive studies; see Poprawski man and Eastop, 2000) that occurs com-
et al. (1992) for a bibliography, and general monly on wild crucifers in Northern and
accounts by Pike and Allison (1991) and Central Europe, but is not usually found on
Hughes (1996). Berest (1980) studied the Brassica crops (Müller, 1986; Heie, 1992).
parasite and predator complex of D. noxia Hille Ris Lambers (1948) could not find
characters to discriminate pseudobrassicae Macrosiphum euphorbiae (potato aphid)

from erysimi, but nevertheless stopped short
of making it a synonym. Others regarded Macrosiphum euphorbiae is one of the few
it as a subspecies of erysimi (e.g. Eastop, cosmopolitan aphid pests of field crops that
1958a; Müller, 1986). Despite these uncer- are undoubtedly of North American origin.
tainties, the name erysimi was used for the The earliest European record is from potato
widely distributed crucifer pest from 1975– at Wye, Kent, England in 1917, after which
2000. it soon became common in Britain and
Although most Lipaphis populations spread to continental Europe (Eastop, 1958b).
throughout the world are continuously par- It is a medium-sized to large, spindle-
thenogenetic, a holocycle does occur on shaped aphid, usually green but sometimes
cruciferous crops (Brassica rapa, Raphanus pink or magenta, the adult apterae often rather
sativus) in western Honshu, Japan (Kawada shiny in contrast to the immature stages,
and Murai, 1979). These aphids have 2n = 8, which have a light dusting of greyish-white
and thus differ in karyotype from holo- wax. In northeastern USA, it has a sexual
cyclic populations of L. erysimi in Northern phase on Rosa, using both wild and culti-
Europe. Chen and Zhang (1985) also repor- vated species as primary hosts (Shands
ted 2n = 8 for Lipaphis in China. In West et al., 1972). In Europe, and probably else-
Bengal, Lipaphis populations are economi- where, M. euphorbiae is mainly anholocyc-
cally important on field crops of mustard, lic, although sexual morphs are produced
Brassica nigra, and here the common karyo- occasionally and the holocycle may some-
type is also 2n = 8 (Kar and Khuda-Bukhsh, times occur (Möller, 1970). The pink form
1991). Sexual morphs have been reported has become much more common in Europe
from northern India, but populations there in the past 10 years. On secondary hosts, M.
are probably mostly anholocyclic. Most euphorbiae is highly polyphagous, feeding
permanently parthenogenetic Lipaphis on more than 200 plant species in more
populations throughout the world have a than 20 different plant families. It is a vec-
9-chromosome karyotype, probably derived tor of more than 40 non-persistent and five
from the 8-chromosome form by dissociation persistent viruses including BYNV, PEMV,
of one autosome to produce a small, unpaired BLRV, Sweet potato leaf-speckling virus,
element. In multivariate morphometric analy- ZYMV, and PLRV. It is an important pest of
sis, samples with 8 and 9 chromosomes group potato, but as a vector of PLRV under field
together and are separated from samples of conditions, it seems to be relatively unim-
European L. erysimi with 10 chromosomes portant in comparison with Myzus persicae
(V.F. Eastop and R.L. Blackman, unpub- (Robert, 1971; Woodford et al., 1995),
lished results), so it was concluded that L. although direct feeding by large numbers
pseudobrassicae should be re-introduced early in the season can cause ‘false top roll’.
for the worldwide crucifer pest, and that it There is a very large literature, but surpris-
probably originated in Eastern Asia (Black- ingly little is known about intraspecific
man and Eastop, 2000). Finding simple mor- variation and specific aphid–host interac-
phological discriminants for the two species tions of M. euphorbiae. In western North
is not easy. Lipaphis pseudobrassicae has America there are several little-known, and
relatively longer antennae and relatively even some undescribed, species closely rela-
shorter siphunculi, and the function ‘(length ted to and almost indistinguishable from
of antennal segment III + length of proces- M. euphorbiae (MacDougall, 1926 and V.F.
sus terminalis) + length of siphunculus’ Eastop, unpublished results). In Europe,
discriminates most specimens. The value of there is a group of closely related species with
this function is more than 2.4 in 90% of more specific host associations (see Watson,
apterae of L. pseudobrassicae, and less than 1982 and Heie, 1994); of these, M. euphor-
2.4 in 90% of apterae of L. erysimi. The biae is most easily confused with Macrosi-
equivalent discriminating value for alatae phum tinctum (= Macrosiphum epilobiellum),
is 3.4. which feeds only on Epilobium spp. It can
Taxonomic Issues 15
be hybridized in the laboratory with Macro- on most of which the populations are highly
siphum stellariae, which is usually found dispersed and individuals are found feed-
on Stellaria holostea, and can colonize other ing singly on the older leaves. The great
plants, but not potato (Möller, 1971). economic importance of M. persicae is due
Meier (1961) provided a general account to its efficiency as a virus vector. It has been
of M. euphorbiae in Europe, Barlow (1962) shown to be able to transmit considerably
studied its development on potato, and more than 100 plant viruses, including the
MacGillivray and Anderson (1964) studied persistent viruses BLRV, Beet western yel-
the factors controlling sexual morph pro- lows virus, Beet mild yellowing virus, BYNV,
duction in eastern Canada. Parasitoids and PEMV, PLRV, Tobacco vein distorting virus,
hyperparasitoids were studied in North Tobacco yellow net virus, and Tobacco yel-
America by Shands et al. (1965) and Sulli- low vein virus. The relationship with PLRV
van and van den Bosch (1971). has received particular attention (e.g. Ponsen,
1972; Eskanderi et al., 1979). Myzus persicae
is also a very efficient vector of numerous
Myzus persicae (peach–potato aphid) non-persistent viruses; e.g. CMV and Bean
yellow mosaic virus to lupins in Western
Myzus persicae is an exceptional species in Australia (Bwye et al., 1997).
many respects; cosmopolitan, extremely poly- As its principal primary host is thought
phagous, highly efficient as a virus vector, to originate from China, one would pre-
and with a great range of genetically-based sume this to be the original homeland of
variability in properties such as colour, life M. persicae. This presumption is, however,
cycle, host-plant relationships, and meth- not without its problems. First, one might
ods of resisting insecticides. Adult apterous expect to find its closest relatives in China.
parthenogenetic females of M. persicae are Yet the species that seem most closely
small to medium-sized, pale greenish-yellow, related to M. persicae, including what many
various shades of green, pink, red, or almost would regard as its sibling species, Myzus
black (apart from the genetically-determined certus, and others with which it readily
colour variation, any one genotype will be hybridizes in the laboratory such as Myzus
more deeply pigmented in cold conditions). myosotidis, are all European. It is difficult
Alatae have a shiny black dorsal abdominal to see how this situation arose. There are
patch, as in other members of the genus no clues from biology, as all other species in
Myzus, and immature alatae are often red or the Myzus subgenus Nectarosiphon except
pink, even of genotypes where the apterae M. persicae have lost their ancestral pri-
are green. Immature males are always some mary host and live all year round on their
shade of yellow or yellow-green. herbaceous host plants. A second problem
The sexual phase of M. persicae occurs concerns the relationship of M. persicae
predominantly on Prunus persica (includ- with PLRV, which seems to be intimate and
ing v. nectarina), except in parts of north- therefore long-standing, but this is in
eastern USA and eastern Canada, where conflict with their respective origins. It
Prunus nigra is the main primary host (Shands is possible, however, that potato leaf
et al., 1969). Host alternation occurs in the roll occurs in some unrecognizable or
temperate regions of all continents, wher- symptomless form in an Asian member of
ever peaches are available and the autumn the Solanaceae.
temperatures are low enough to allow pro- As might be expected of such an adapt-
duction of the sexual morphs (Blackman, able and fast-evolving genome, biology and
1974). Spring populations on peach become host relationships are likely to be changing
very dense, severely curling the leaves. In faster than morphology, causing problems
contrast to its extreme primary host speci- of identification and identity. Specimens of
ficity, the secondary hosts are in more than M. certus on slides are difficult enough to
40 different plant families. They include distinguish from those of M. persicae, althou-
very many economically important plants, gh this species is clearly very different in its
biology and host relations (living all year on Apterae on tobacco are predominantly pink/
Caryophyllaceae and Violaceae, and having red in colour, and have acquired resistance
its sexual phase on these plants, with apterous to insecticides far more slowly than those
males). Two other taxa, Myzus dianthicola on other crops (Takada, 1979; Semtner et al.,
and Myzus antirrhinii, are even more like 1990). Blackman (1987) demonstrated using
M. persicae, and individual slide-mounted multiple discriminant analysis that samples
specimens cannot be distinguished reliably from tobacco in many parts of the world
from that species. These two are both per- could be differentiated from M. persicae on
manently parthenogenetic as far as is other crops, indicating that populations on
known, and their karyotypes are structurally tobacco worldwide constituted a monophy-
heterozygous (Blackman, 1980). Myzus dian- letic lineage for which he proposed the name
thicola is found only on Dianthus, usually Myzus nicotianae. Most of the samples ana-
in glasshouses, where its consistently deep lysed by Blackman (1987) were from regions
yellow-green colour and the leaf chlorosis where populations are permanently parthe-
that it causes distinguish it respectively nogenetic, but Margaritopoulos et al. (2000)
from M. certus and M. persicae. Myzus found that holocyclic populations of tobacco
antirrhinii may be almost as polyphagous as aphids in Greece could also be discrimina-
M. persicae, but has certain characteristic ted from those collected from other crops,
hosts such as Antirrhinum and Buddleja and and from peach away from tobacco-growing
a more consistent mid-green to dark green col- regions. It has been suggested (Clements
our, and there are also differences in et al., 2000a) that these morphological differ-
allozymes and at rDNA and microsatellite ences could be due to phenotypic plasticity.
loci (Fenton et al., 1998; Terradot et al., However, it is clear that the differences are
1999). Both M. dianthicola and M. antirrhinii genetically-based, as all the samples analysed
are found in Europe and North America, the were clones reared under controlled condi-
former also being found in New Zealand tions on the same host plant. The clones origi-
and the latter in Australia. Although probably nating from peach in tobacco-growing regions
of recent origin, they seem to be isolated had never even seen a tobacco plant.
from other members of the group by their Genetic isolation between tobacco-
obligate parthenogenesis, and are therefore adapted and non-tobacco-adapted forms can-
best treated as discrete taxa (Blackman and not be complete, as the E4 and FE4 genes
Brown, 1991). The karyotype of M. antirrhinii amplified in insecticide-resistant aphids
is remarkably variable, and is of cytogenetic are identical in the two forms (Field et al.,
interest because fusions and dissociations of 1994). However, these genes apparently have
chromosomes have occurred in the absence taken many years to cross into tobacco aphids.
of genetic recombination (Hales et al., 2000). For example, holocyclic populations of M.
Although complicating the practical persicae on peach in Southern Europe have
identification of M. persicae, none of the been resistant to organophosphates since
forms discussed above have questionable about 1962, yet such resistance in tobacco
taxonomic status. However, for many years aphids was first reported in holocyclic pop-
it has been recognized that populations of ulations in northern Greece in the mid-
M. persicae on tobacco (the ‘tobacco aphid’) 1980s. This may be where introgression of
are distinct from populations on other these genes into tobacco-adapted genotypes
plants (de Jong, 1929; Brain, 1942; Müller, occurred, selection then strongly favouring
1958; Takada, 1986). The aphid attacking their spread to other populations. Absence
commercial varieties of Nicotiana tabacum of complete reproductive isolation between
forms large, dense colonies at the growing the two forms, perhaps in conjunction with
points and on the youngest leaves, and a very recent origin of the tobacco-adapted
seems able to avoid or tolerate the exudates form, may explain the failure to find consis-
of the glandular trichomes, which are not tent diagnostic genetic markers (Fenton
only sticky but contain repellent or toxic et al., 1998; Margaritopoulos et al., 1998;
chemicals (Georgieva, 1998; Wang et al., 2001). Clements et al., 2000a,b), or the divergence
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Kieron whirled and caught Freka's blade on his own. The courtiers
drew back, giving them room to fight. No one made a move to
interfere. It was known that Valkyrs had sacked the city of Neg, and
according to the warrior code the two warlords must be allowed to
fight to the death if they wished.
Kieron made no attack. Instead he retreated before the
expressionless Freka.
"Did you know, Freka," asked Kieron softly, "that Geller of the
Marshes is dead? He was your father in a way, wasn't he?"
Freka made no reply, and for a moment the only sound in the hushed
chamber was the ring of blades.
Suddenly Kieron lunged. His sword pierced Freka from breast to back.
The Valkyr stepped back and pulled his blade clear. The crowd
gasped, for Freka the Unknown did not fall....
"Are you really unkillable?" breathed Kieron. "I wonder!"
Again he lunged under the mechanical guard of the Kalgan. Again his
blade sank deep. Freka backed away for a moment, still alert and
unwounded.
Kieron shouted derisively at the star-kings: "Great warriors! Do you
see? You have followed the leadership of an android! A homunculus
spawned by the warlock Geller!"
A gasping roar went up in the chamber. A sound of superstitious
horror and growing anger.
Kieron parried a thrust and brought his blade down on Freka's sword
arm. Hard. A sword clattered to the flagstones—still gripped by a
slowly relaxing hand. There was no blood. The android still moved in,
eyes expressionless, his one hand reaching for his enemy. Kieron
struck again. A clean cut opened from shoulder to belly, slicing the
artificial tendons and leaving the android helpless but still erect.
Kieron raised and lowered his blade in glittering arcs. Freka ... or the
thing that had been Freka ... collapsed in a grotesque heap. Still it
moved. Kieron passed his point again and again through the
quivering mass until at long last it was still. Somewhere a woman
fainted.
A thick silence fell over the assemblage. All eyes turned to Ivane. She
stood staring at the remnants of the thing that had been ... almost ...
a man. Her hand fluttered at her throat.
Alys' voice cut through the heavy stillness. "Arrest that woman for the
murder of my brother Toran!"
But the crowd of courtiers was thinking of other things. Jaded and
cynical, they had seen with their own eyes that Ivane was a familiar
of the dreaded Great Destroyer. Someone cried: "Witch! Burn her!"
The mass of courtiers and warriors swept forward, screaming for the
kill. Kieron leaped for the dais, his sword still bared.
"I'll kill the first one who sets foot on the Great Throne!" he cried.
But Ivane had heard the crowd sounds. The black mantle slipped
from her shoulders, and she stood stripped to the waist, like a marble
goddess—her eyes recapturing some of their icy hauteur. Then,
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driven it deep into her breast.
Kieron caught her as she fell, feeling the warm blood staining his
hands. He eased her down on the foot of the Great Throne and laid
his ear to her breast.
There was no pulse. Ivane was dead.
Before the assembled Court, the Warlord of Valkyr knelt before his
Empress. The star-kings had gone, and the Valkyrs were the last
outworld warriors remaining in the Imperial City. Now, they too,
would take their leave.
The Empress sat on the Great Throne, mantled in sable. Somehow,
the huge throne and the vast vaulted chamber seemed to make her
look small and frail.
"Your Imperial Majesty," said Kieron, "have we your leave to go?"
Alys' eyes were bright with tears. She leaned forward so that none
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He looked up at her once, pain in his eyes, but he did not speak.
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"Permission is granted, My Lord of Valkyr. You ... you may return to
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Kieron!"
Kieron raised her jewelled hands to his lips and kissed them.... Then
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Helmut F. van Emden

Emeritus Professor of Horticulture, University of Reading, UK

Plant and Invertebrate Ecology Division, Rothamsted Research, UK

CABI Head Office CABI North American Office

© CAB International 2007. All rights reserved. No part of this

A catalogue record for this book is available from the

ISBN-13: 978 0 85199 819 0

Typeset by AMA DataSet Ltd, UK.

who died while this book was in preparation

He told me he particularly enjoyed writing his chapter,

List of Contributors xxv

2. Population Genetic Issues: The Unfolding Story Using

Application of molecular markers in aphidology 32

3. Life Cycles and Polymorphism 69

4. Host-plant Selection and Feeding 87

5. Nutrition and Symbiosis 115

6. Growth and Development 135

7. Aphid Movement: Process and Consequences 153

The nature of aphid movement 154

8. Predators, Parasitoids and Pathogens 187

9. Chemical Ecology 235

Interactions between aphids 238

10. Insecticide Resistance 261

11. Coping with Stress 287

12. Population Dynamics 311

Biological background 312

13. Feeding Injury 331

14. Transmission of Plant Viruses 353

Cultural practices 372

15. Chemical Control 391

16. Cultural Control 423

17. Host-plant Resistance 447

18. Biological Control 469

Aphids on field crops and amenity plants 486

19. Monitoring and Forecasting 515

21. IPM Case Studies: Brassicas 549

Monitoring and forecasting 552

22. IPM Case Studies: Berry Crops 561

23. IPM Case Studies: Cotton 573

24. IPM Case Studies: Leafy Salad Crops 587

Executive summary 593

25. IPM Case Studies: Grain 597

26. IPM Case Studies: Seed Potato 613

27. IPM Case Studies: Sorghum 627

28. IPM Case Studies: Cucurbits 639

29. IPM Case Studies: Deciduous Fruit Trees 651

31. Decision Support Systems 677

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