Acarología

Manjit
Singh Dhooria
Fundamentals
of Applied
Acarology
Fundamentals of Applied Acarology
Manjit Singh Dhooria
Fundamentals of Applied
Acarology
Manjit Singh Dhooria
Department of Entomology
Punjab Agricultural University
Ludhiana, Punjab, India
ISBN 978-981-10-1592-2 ISBN 978-981-10-1594-6 (eBook)

DOI 10.1007/978-981-10-1594-6
Library of Congress Control Number: 2016953350
© Springer Science+Business Media Singapore 2016

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The registered company address is: 152 Beach Road, #22-06/08 Gateway East, Singapore 189721,
Singapore
My Wife: Rajinder Dhooria
My Sons:
1. Ramanjot Dhooria: Reena, Sarah and Geet
2. Taranjot Dhooria: Saveena and Mannat
Preface
Acarology, the study of mites and ticks, is a subdiscipline of Zoology and is many
times considered in the field of Entomology (the study of insects). Mites and ticks
are distributed throughout the world and inhabit almost every ecosystem (both ter-
restrial and aquatic). More than 55,000 species of mites and ticks are already
described, and the number of undescribed species may reach about 20-fold as mite
fauna in most of the world regions are still poorly explored. Mites and ticks directly
affect humans as pests of different crops, fruit plants, vegetable crops and field
crops; as parasites of human beings, veterinary animals, poultry and pets; as pests
of stored grains and other products, mushrooms and cheese; and as parasites of
honeybees. Mite infestations are responsible for economic losses worth billions of
dollars in terms of reduced crop yields and lowered quality of produce. Some spe-
cies of mites serve as vectors of various plant diseases. Ticks cause losses through
blood feeding and by transmitting Lyme disease, babesiosis, Rocky Mountain fever,
tularaemia and many other diseases among man and animals. House-dust mite aller-
gies and tick bite allergies are also common in many parts of the world.
Our knowledge of biology, ecology and the importance of mites has largely
expanded. At the same time, the actual and potential economic importance of these
creatures continues to grow worldwide, and their success in colonizing the new
regions makes them an ongoing quarantine threat in many parts of the world. There
is a great necessity of an up-to-date compilation of basic and applied knowledge on
mites and their proper management that is otherwise found scattered in a variety of
languages and literature throughout the world.
The present book, Fundamentals of Applied Acarology, is written keeping in
view nonavailability of any standard text dealing in different aspects of acarology at
one place. Separate chapters in this book are devoted to the importance of acarol-
ogy, historical account, acarine technology, morphology and anatomy of Acari; and
feeding, development and reproduction providing knowledge on diapause and water
balance. The book includes a recent classification of subclass Acari, and illustra-
tions are provided for the identification of 107 important families of Acari. The role
of mites in breaking down soil organic matter and affecting the quality of water
bodies is discussed in detail. Molecular developments in relation to mites and ticks
are also discussed. The role of mites and ticks in quarantines of plants and animals,
forensic/criminal investigations and the importance of accidental acarophagy are
also discussed in detail. Transmission of plant diseases by mite vectors is
vii
viii Preface
thoroughly discussed. Safe usage of pesticides based on their mode of action

(IRAC’s groups), development of acaricide resistance and measures to mitigate it
are also discussed.
Mite pests of fruit trees, vegetable plants, floricultural plants and field crops;
mite problems in greenhouses/polyhouses; mite problems encountered under
organic cultivation of plants and their management through minimum usage of pes-
ticides are emphasized in different chapters. The role of different predaceous mites
in controlling plant pests like thrips, aphids and scale insects is elaborately dis-
cussed. Biological control of phytophagous mites is discussed in detail.
Different animal parasitic mites and ticks are discussed from veterinary and
medical point of view.
At the end of each chapter, many important references for further reading and
electronic references (ER) in the form of YouTube links and other weblinks are
given to understand fully how these tiny creatures look like and behave, feed and
reproduce, the nature of damage they cause to plants and animals and measures to
mitigate them. At the end, a glossary of important taxonomic and integrated pest
management terms is provided which makes understanding the contents in a more
suitable way creating great lust for knowing more and more about these creatures.
I am grateful to the late Dr G.P. ChannaBasavanna and Dr S.K. Gupta, both
distinguished acarologists from India, who encouraged me and lent their moral
support for writing such a book on applied aspects. I also sincerely acknowledge
Dr Uri Gerson of Israel and Dr K. Ramaraju of India, eminent acarologists, for their
useful suggestions.
The knowledge contained in the book may prove very useful for ‘General and
Applied Acarology’ course for graduate and postgraduate levels, teachers and
researchers in entomology, pest control advisors, professional entomologists, pesti-
cide industry managers, policy planners and others having interest in mites and
ticks.
Ludhiana, India M.S. Dhooria

About the Book
Acarology, the study of mites and ticks, is a subdiscipline of Zoology and is many
times considered in the field of Entomology (the study of insects). Mites and ticks
are distributed throughout the world and inhabit almost every ecosystem (both ter-
restrial and aquatic) including grassland soils. More than 55,000 species of mites
and ticks are already described, but the number of undescribed species may reach
up to 20-fold as mite fauna in most of the world regions are still poorly explored.
Mites and ticks directly affect humans as pests of different crops, fruit plants, veg-
etable crops and field crops; as parasites of human beings, veterinary animals, poul-
try and pets; as pests of stored grains and other products, mushrooms and cheese;
and as parasites of honeybees. Mite infestations are responsible for economic losses
worth billions of dollars in terms of reduced crop yields and lowered quality of
produce. Many species of mites serve as vectors of various plant diseases; some
species of ticks cause losses through blood feeding and by transmitting Lyme dis-
ease, babesiosis, Rocky Mountain fever, tularaemia and many other diseases among
man and animals. House-dust mite allergies and tick bite allergies are also common
in many parts of the world.
The present book, Fundamentals of Applied Acarology, is written keeping in
view nonavailability of any standard text dealing in different aspects of acarology at
one place. Separate chapters in this book are devoted to the importance of acarol-
ogy; historical account; acarine technology; morphology and anatomy of Acari; and
feeding, development and reproduction providing knowledge on diapause and water
balance. Molecular developments in relation to mites and ticks are also discussed.
The role of mites and ticks in quarantines of plants and animals, forensic/crimi-
nal investigations, and the importance of accidental acarophagy are discussed in
detail in different chapters. Safe usage of pesticides based on their mode of action
(IRAC’s groups), development of acaricide resistance and measures to mitigate it
are also discussed.
Mite pests of fruit trees, vegetable plants, floricultural plants and field crops;
mite problems in greenhouses/polyhouses; mite problems encountered under
organic cultivation of plants and their management through minimum usage of pes-
ticides are emphasized in different chapters. The role of different predaceous mites
in controlling plant pests like thrips, aphids and scale insects is elaborately dis-
cussed. Biological control of phytophagous mites is discussed in detail.
ix
x About the Book
Different animal parasitic mites and ticks are discussed from veterinary and
medical point of view. Many YouTube and other weblinks explain in an interesting
way emphasizing the importance of salient information on these creatures and mea-
sures to combat them.
At the end of each chapter, many important references for further reading and
electronic references (ER) in the form of YouTube links and other weblinks are
given to understand fully how these tiny creatures look like and behave, feed and
reproduce, the nature of damage they cause to plants and animals and measures to
mitigate them. Weblinks will stimulate interest in the readers for more information
about different mites and ticks.
The knowledge contained in the book may prove as best material for ‘General
and Applied Acarology’ course for graduate and postgraduate levels, teachers and
researchers in entomology, pest control advisors, professional entomologists, pesti-
cide industry managers, policy planners and others having interest in mites and
ticks.
562-I Block, B.R.S. Nagar, M.S. Dhooria

Ludhiana-141012, Punjab, India
E-mail: msdhooria@yahoo.com
Contents
1 Acarology and Its Importance................................................................ 1

Further Reading......................................................................................... 7
2 Historical Account of Acarology............................................................. 9
2.1 Developments Prior to 1850 AD................................................... 10
2.2 Progress During 1851–1950 AD................................................... 10
2.3 Developments Since 1951............................................................ 11
2.4 OSU Acarology Development Programme................................... 12
2.5 Acarological Societies.................................................................. 12
2.6 Journals in Acarology................................................................... 16
2.7 Important Books Published........................................................... 17
2.8 Conclusions................................................................................... 18
Further Reading......................................................................................... 18
3 Acarine Technology.................................................................................. 21
3.1 Collection...................................................................................... 22
3.1.1 Plant-Associated Mites................................................. 22
3.1.2 Water Mites................................................................... 23
3.1.3 Soil Mites...................................................................... 24
3.1.4 Parasitic Mites.............................................................. 25
3.1.5 Collection of Ticks........................................................ 27
3.2 Preservation/Killing in a Medium................................................. 28
3.3 Preparation (Slide Mounting)....................................................... 28
3.3.1 Preparation for Water Mites.......................................... 29
3.3.2 Plant-Associated Mites................................................. 29
3.3.3 Preparation of Ticks for Study...................................... 30
3.3.4 Mounting Ticks on Glass Slides................................... 31
3.3.5 Staining......................................................................... 31
3.3.6 Slide Preparation Technique......................................... 31
3.4 Preparation for Scanning Electron Microscopy Studies (SEM)... 32
3.5 Preparation for Molecular Analysis Studies................................. 33
3.6 Photomicrography of Mites.......................................................... 33
xi
xii Contents
3.7 Packing and Shipping Mites......................................................... 33

3.7.1 Specimens in Vials........................................................ 33
3.7.2 Shipping Microscope Slides......................................... 34
3.7.3 Shipping Specimens...................................................... 34
3.8 Working with Live Mites.............................................................. 34
3.9 Rearing Techniques....................................................................... 35
3.10 Sampling Methods........................................................................ 35
3.10.1 Direct Counting............................................................ 35
3.10.2 Imprint Counting.......................................................... 36
3.10.3 Mite-Brushing Machine................................................ 36
3.10.4 Beating Foliage............................................................. 36
3.10.5 Washing Technique....................................................... 36
3.11 Acaricide Testing Techniques....................................................... 36
3.11.1 Slide-Dip Method......................................................... 37
3.11.2 Leaf-Dip Method.......................................................... 37
3.11.3 Residual Leaf Disc Assays............................................ 37
3.11.4 Ovolarvicide Tests........................................................ 37
3.12 Conclusions................................................................................... 38
Further Reading......................................................................................... 38
4 Morphology and Anatomy of Acari........................................................ 41
4.1 External Morphology.................................................................... 42
4.1.1 Integument.................................................................... 42
4.1.2 Differences Between Hard Ticks
and Soft Ticks............................................................... 44
4.1.3 Body Divisions............................................................. 45
4.1.4 Sensory Receptors........................................................ 51
4.2 Internal Anatomy.......................................................................... 55
4.2.1 Digestive System.......................................................... 55
4.2.2 Central Nervous System............................................... 56
4.2.3 Circulatory System....................................................... 56
4.2.4 Excretion....................................................................... 56
4.2.5 Respiratory System....................................................... 57
4.2.6 Reproductive System.................................................... 57
4.2.7 Musculature.................................................................. 57
4.3 Conclusions................................................................................... 58
Further Reading......................................................................................... 58
5 Classification of Subclass Acari.............................................................. 63
5.1 Superorder Parasitiformes............................................................ 64
5.1.1 Order Opilioacarida...................................................... 64
5.1.2 Order Holothyrida......................................................... 65
5.1.3 Order Mesostigmata...................................................... 65
5.1.4 Order Ixodida................................................................ 66
Contents xiii
5.2 Superorder Acariformes................................................................ 67

5.2.1 Order Trombidiformes.................................................. 68
5.2.2 Order Sarcoptiformes.................................................... 68
5.3 Conclusions................................................................................... 70
Further Reading......................................................................................... 70
6 Important Acarine Families.................................................................... 73
6.1 Superorder Parasitiformes............................................................ 74
6.1.1 Order Holothyrida......................................................... 74
6.1.2 Order Opilioacarida...................................................... 74
6.1.3 Order Ixodida................................................................ 75
6.1.4 Order Mesostigmata...................................................... 80
6.2 Superorder Acariformes................................................................ 96
6.2.1 Order Trombidiformes.................................................. 96
6.3 Conclusions................................................................................... 156
Further Reading......................................................................................... 157
7 Feeding, Development and Reproduction.............................................. 161
7.1 Development................................................................................. 162
7.1.1 Eggs.............................................................................. 162
7.1.2 Prelarva......................................................................... 162
7.1.3 Larvae........................................................................... 163
7.1.4 Nymphs......................................................................... 163
7.1.5 Protonymph................................................................... 163
7.1.6 Deutonymphs................................................................ 164
7.1.7 Tritonymphs.................................................................. 164
7.1.8 Adults............................................................................ 164
7.1.9 Life cycle...................................................................... 164
7.2 Feeding......................................................................................... 166
7.2.1 Ticks.............................................................................. 166
7.2.2 Phytophagous Mites...................................................... 167
7.3 Reproduction................................................................................. 168
7.3.1 Mites............................................................................. 168
7.3.2 Ticks.............................................................................. 169
7.4 Tick Ecology................................................................................. 169
7.5 Water Balance............................................................................... 170
7.6 Diapause in Acari.......................................................................... 171
7.6.1 Mites............................................................................. 171
7.6.2 Ticks.............................................................................. 171
7.7 Conclusions................................................................................... 172
Further Reading......................................................................................... 172
8 Molecular Biology and Acarology.......................................................... 175
8.1 Phylogenetic Relationship............................................................ 176
8.2 Species Diversity.......................................................................... 177
8.3 Molecular Markers........................................................................ 178
xiv Contents
8.4 Dispersal of Populations and Species........................................... 180

8.5 Development of Host Races and Species Link............................. 180
8.6 Mite Associates............................................................................. 181
8.7 Molecular Identification of House Dust and Storage Mites......... 181
8.8 Genome Sequencing of Two-Spotted Spider Mite....................... 182
8.9 Tick Genomics.............................................................................. 182
8.10 Conclusions................................................................................... 183
Further Reading......................................................................................... 184
9 Water Mites.............................................................................................. 187
9.1 Hydrachnidia Diversity................................................................. 188
9.2 Feeding Habits.............................................................................. 188
9.3 Habitat........................................................................................... 189
9.4 Effect of Water Pollution.............................................................. 189
9.5 Potential as Indicators of Environmental Safety........................... 190
9.6 Websites on Water Mites............................................................... 190
9.6.1 Water Mites of North America...................................... 190
9.6.2 Hydracarina.Org........................................................... 191
9.6.3 Key to Families and Subfamilies of Water Mites
(Hydracarina)................................................................ 191
9.6.4 Acari1: European Water Mites Research...................... 191
9.7 Life Cycle..................................................................................... 191
9.8 Role of Water Mites in Food Chain.............................................. 192
9.9 Biological Control of Insect Pests................................................ 193
9.9.1 Parasites........................................................................ 193
9.9.2 Parasites of Mosquitoes................................................ 193
9.9.3 Predators....................................................................... 194
9.10 Predators of Water Mites.............................................................. 194
9.11 Conclusions................................................................................... 194
Further Reading......................................................................................... 195
10 Soil Mites................................................................................................... 197
10.1 Oribatid Diversity......................................................................... 198
10.2 Distribution................................................................................... 199
10.3 Life History and Development..................................................... 200
10.4 Feeding Habits.............................................................................. 200
10.5 Population Dynamics.................................................................... 201
10.6 Nematophagous Mites.................................................................. 201
10.7 Exposure to Predators................................................................... 202
10.8 Effect of Drought/Burning............................................................ 202
10.9 Intermediate Hosts of Tapeworms................................................ 202
10.10 Effect of Pesticides and Industrial Wastes.................................... 202
10.11 Effect of Fertilizers....................................................................... 203
10.12 Economic Importance of Oribatid Mites...................................... 203
10.13 Conclusions................................................................................... 204
Further Reading......................................................................................... 205
Contents xv
11 Quarantine Acarology............................................................................. 207

11.1 Acari Dispersal............................................................................. 208
11.2 Quarantine Pests........................................................................... 209
11.3 Invasive Mites and Ticks............................................................... 209
11.4 Phytophagous Mites...................................................................... 210
11.5 Bee Mites...................................................................................... 211
11.6 Animal Parasitic Mites and Ticks................................................. 211
11.7 Quarantine Organizations............................................................. 212
11.8 Internet Resources for Information on Pest.................................. 213
11.9 Other Internet Resources Providing Information on Mites.......... 214
11.10 Pest Information/Fact Sheets/Field Guides.................................. 214
11.11 Conclusions................................................................................... 215
Further Reading......................................................................................... 216
12 Mite Pests of Horticultural Crops.......................................................... 219
12.1 Fruits............................................................................................. 220
12.1.1 Temperate Fruits........................................................... 220
12.1.2 Tropical/Subtropical Fruits........................................... 232
12.2 Vegetables..................................................................................... 248
12.2.1 Family Alliaceae........................................................... 249
12.2.2 Family Convolvulaceae................................................ 250
12.2.3 Family Cruciferaceae.................................................... 250
12.2.4 Family Cucurbitaceae................................................... 250
12.2.5 Family Leguminosae..................................................... 253
12.2.6 Family Solanaceae........................................................ 254
12.2.7 Family Malvaceae......................................................... 259
12.2.8 Family Umbelliferae..................................................... 260
12.2.9 Other Vegetables........................................................... 261
12.3 Mite Pests of Floricultural Plants................................................. 263
12.3.1 Bulb Mites (Rhizoglyphus spp.).................................... 264
12.3.2 Cyclamen Mites (Steneotarsonemus pallidus)............. 264
12.3.3 False Spider Mites (Brevipalpus spp.).......................... 264
12.3.4 Lewis Mite (Eotetranychus lewisi)............................... 265
12.3.5 Two-Spotted Spider Mite (Tetranychus urticae).......... 265
12.3.6 Broad Mite (Polyphagotarsonemus latus).................... 268
12.3.7 Erinose Mite (Aceria hibisci)........................................ 269
12.3.8 Aloe Gall Mite (Eriophyes aloinis).............................. 269
12.3.9 Southern Red Mite (Oligonychus ilicis)....................... 269
12.3.10 Jasmine Mite (Aceria jasmini)...................................... 270
12.3.11 Phyllocoptes fructiphilus.............................................. 270
12.4 Conclusions................................................................................... 271
Further Reading......................................................................................... 271
xvi Contents
13 Mite Pests of Field Crops......................................................................... 275

13.1 Cereals.......................................................................................... 276
13.1.1 Barley (Hordeum vulgare; Poaceae)............................. 276
13.1.2 Maize (Zea mays; Poaceae).......................................... 277
13.1.3 Rice (Oryza sativa; Poaceae)........................................ 278
13.1.4 Sorghum (Sorghum bicolor; Gramineae)...................... 278
13.1.5 Wheat (Triticum aestivum; Gramineae)........................ 279
13.1.6 Oats (Avena sativa; Poaceae)....................................... 281
13.2 Pulses............................................................................................ 282
13.2.1 Pigeon pea (Cajanus cajan; Fabaceae)......................... 282
13.2.2 Cowpea (Vicia spp.)...................................................... 284
13.2.3 Black gram (Vigna mungo)........................................... 285
13.2.4 Green gram (Vigna radiata).......................................... 285
13.2.5 Winter Pulses................................................................ 286
13.3 Fibre Crops................................................................................... 286
13.3.1 Cotton (Gossypium spp.; Malvaceae)........................... 287
13.3.2 Jute (Corchorus capsularis; Malvaceae)...................... 288
13.3.3 Sunn Hemp (Crotalaria juncea; Fabaceae).................. 289
13.4 Oilseed Crops................................................................................ 289
13.4.1 Soybeans (Glycine max; Fabaceae)............................. 289
13.4.2 Canola (Brassica napus; Brassicaceae)........................ 290
13.4.3 Groundnut (Arachis hypogaea; Leguminosae)............. 291
13.4.4 Castor Bean (Ricinus communis; Euphorbiaceae)........ 292
13.4.5 Sesame (Sesamum indicum; Pedaliaceae).................... 292
13.4.6 Safflower (Carthamus tinctorius; Asteraceae).............. 294
13.4.7 Sunflower (Helianthus annuus; Asteraceae)................. 294
13.4.8 Jatropha (Jatropha curcas; Euphorbiaceae).................. 294
13.5 Plantation Crops............................................................................ 294
13.5.1 Sugarcane (Saccharum officinarum; Poaceae)............. 294
13.5.2 Cassava (Manihot esculenta; Euphorbiaceae).............. 297
13.5.3 Tea (Camellia sinensis; Theaceae)............................... 298
13.5.4 Cardamom (Elettaria sp.; Zingiberaceae).................... 300
13.5.5 Areca Nut (Areca catechu; Arecaceae)......................... 300
13.5.6 Rubber (Hevea brasiliensis; Euphorbiaceae)............... 301
13.5.7 Cashew (Anacardium occidentale; Anacardiaceae)...... 301
13.6 Fodder Crops................................................................................ 301
13.6.1 Alfalfa or Lucerne (Medicago sativa; Fabaceae).......... 301
13.6.2 Rye Grass (Lolium sp.; Poaceae).................................. 302
13.6.3 Vicia (Vicia spp.; Fabaceae).......................................... 302
13.6.4 Timothy (Phleum pratense; Poaceae)........................... 303
13.7 Conclusions................................................................................... 303
Further Reading......................................................................................... 304
Contents xvii
1 4 Mite Pests of Greenhouse Crops............................................................. 307

14.1 Mite Pests in Greenhouses............................................................ 308
14.1.1 Spider Mites.................................................................. 308
14.1.2 False Spider Mites........................................................ 309
14.1.3 Tarsonemid Mites......................................................... 310
14.1.4 Rust Mites..................................................................... 310
14.1.5 Bulb Mites.................................................................... 311
14.2 Biological Control of Mite Pests.................................................. 311
14.2.1 Through Predatory Phytoseiid Mites............................ 311
14.2.2 Through Predatory Insects............................................ 312
14.3 Management of Mite Pests in Greenhouses.................................. 313
14.4 Conclusions................................................................................... 314
Further Reading......................................................................................... 314
1 5 Management of Mites Under Organic Farming.................................... 317
15.1 Management of Mites in Greenhouses......................................... 318
15.2 Management of Mites on Organic Crops...................................... 319
15.3 Cultural Control............................................................................ 320
15.4 Water Management....................................................................... 321
15.5 Good Farm Sanitation................................................................... 321
15.6 Other Cultural Practices................................................................ 321
15.7 Monitoring for Mite Pests and Their Natural Enemies................ 322
15.8 Biological Control........................................................................ 322
15.9 Microbiol Control......................................................................... 323
15.10 Chemical Control.......................................................................... 323
15.11 Important Websites....................................................................... 324
15.12 Conclusions................................................................................... 325
Further Reading......................................................................................... 326
1 6 Mite Transmission of Plant Diseases...................................................... 327
16.1 Plant Viral Diseases Vectored by Eriophyid Mites....................... 328
16.1.1 Wheat Streak Mosaic Virus (WSMV).......................... 329
16.1.2 Wheat Spot Mosaic Virus (WSpMV)........................... 330
16.1.3 Ryegrass Mosaic Virus (RgMV)................................... 331
16.1.4 Agropyron Mosaic Virus (AMV).................................. 331
16.1.5 Fig Mosaic Virus (FMV).............................................. 332
16.1.6 Peach Mosaic................................................................ 332
16.1.7 Cherry Mottle Leaf Virus (CMLV)............................... 332
16.1.8 Currant Reversion Disease............................................ 333
16.1.9 Pigeon Pea Sterility Mosaic Disease (PPSMV)............ 333
16.1.10 Rose Rosette Disease (RRD)........................................ 334
16.2 Plant Viral Diseases Vectored by False Spider Mites................... 334
16.2.1 Citrus Leprosis.............................................................. 335
16.2.2 Passion Fruit Green Spot Virus (PGSV)....................... 335
16.2.3 Coffee Ring Spot Virus (CoRSV)................................. 336
16.2.4 Orchid Fleck Virus (OFV)............................................ 336
xviii Contents
16.3 Plant Viral Diseases Vectored by Spider Mites............................. 336

16.4 Plant Fungal Diseases Vectored by Eriophyid Mites.................... 337
16.4.1 Mango Malformation Disease (MMD)......................... 337
16.5 Management of Mite-Vectored Plant Viral Diseases.................... 337
16.6 Conclusions................................................................................... 338
Further Reading......................................................................................... 338
17 Mite Pests of Mushrooms........................................................................ 341
17.1 Pest Mites...................................................................................... 342
17.1.1 Tyrophagus spp............................................................. 342
17.1.2 Tarsonemus myceliophagus.......................................... 343
17.1.3 Red Pepper Mites (Pygmephorus spp.)........................ 343
17.1.4 Other Mite Pests of Mushrooms................................... 344
17.2 Predaceous Mites.......................................................................... 344
17.3 Management of Mushroom Mite Pests......................................... 344
17.4 Chemical Control of Mites........................................................... 345
17.5 Conclusions................................................................................... 345
Further Reading......................................................................................... 346
18 Mite Problems of Stored Foods............................................................... 349
18.1 Stored Product Mites.................................................................... 350
18.1.1 Pest Mites...................................................................... 350
18.1.2 Predaceous Mites.......................................................... 355
18.1.3 Parasitic Mite................................................................ 356
18.1.4 Scavengers and Saprobes.............................................. 356
18.2 Temperature/Humidity Relations.................................................. 356
18.3 Detection of Mites........................................................................ 357
18.4 Allergy Due to Mites in Storages................................................. 357
18.5 Management of Mites in Storages................................................ 358
18.5.1 Stored Grains................................................................ 358
18.5.2 Bulb Mites.................................................................... 359
18.5.3 Stored Packages in Groceries....................................... 360
18.6 Biological Control........................................................................ 361
18.7 Conclusions................................................................................... 361
Further Reading......................................................................................... 361
1 9 Mites Predaceous on Pests of Agriculture.............................................. 363
19.1 Insects........................................................................................... 364
19.1.1 Order Diptera................................................................ 364
19.1.2 Order Orthoptera........................................................... 365
19.1.3 Order Hemiptera........................................................... 366
19.1.4 Order Thysanoptera...................................................... 369
19.1.5 Order Trichoptera.......................................................... 370
19.1.6 Order Thysanura........................................................... 370
19.1.7 Order Collembola......................................................... 370
19.1.8 Order Blattoidea............................................................ 371
Contents xix
19.1.9 Order Coleoptera.......................................................... 371

19.1.10 Order Lepidoptera......................................................... 373
19.1.11 Order Hymenoptera...................................................... 374
19.2 On Snails and Slugs...................................................................... 374
19.3 On Symphylans............................................................................. 375
19.4 On Nematodes.............................................................................. 376
19.5 Mite Associations on Earthworms................................................ 377
19.6 Conclusions................................................................................... 377
Further Reading......................................................................................... 378
2 0 Biological Control of Phytophagous Mites............................................ 381
20.1 Biological Control of Phytophagous Mites.................................. 382
20.1.1 Predaceous Mites.......................................................... 382
20.1.2 Insects Predaceous on Mites......................................... 386
20.1.3 Microbial Control of Phytophagous Mites................... 388
20.1.4 Biological Control of Weeds by Phytophagous Mites.. 390
20.2 Conclusions................................................................................... 394
Further Reading......................................................................................... 394
21 Pesticides................................................................................................... 399
21.1 Mite Outbreaks............................................................................. 400
21.2 Resurgence.................................................................................... 401
21.3 Development of Specific Acaricidal Compounds......................... 401
21.4 Classification of Acaricides.......................................................... 402
21.4.1 Inorganic Acaricides..................................................... 402
21.5 Synthetic Organic Acaricides........................................................ 403
21.5.1 Acaricides Based on Their Chemistry.......................... 403
21.5.2 Acaricide Classification Based on Mode of Action...... 403
21.6 Development of Specific Acaricides............................................. 405
21.6.1 New Synthetic Acaricides............................................. 406
21.6.2 Natural Acaricides and Other Alternative Solutions..... 407
21.7 Role of Acaricides in IPM of Spider Mites.................................. 407
21.8 Acaricide Resistance..................................................................... 408
21.9 Management of Acaricide Resistance........................................... 409
21.10 Conclusions................................................................................... 409
Further Reading......................................................................................... 410
22 Parasitic Mites on Honeybees................................................................. 413
22.1 Parasitic Mites.............................................................................. 414
22.1.1 Tracheal Mite (Acarapis woodi; Tarsonemidae)........... 414
22.1.2 Tropilaelaps clareae..................................................... 417
22.1.3 Varroa jacobsoni........................................................... 418
22.1.4 Varroa destructor.......................................................... 421
22.1.5 Other Mites Found on Bees.......................................... 423
22.2 Conclusions................................................................................... 423
Further Reading......................................................................................... 423
xx Contents
23 Medical and Veterinary Acarology......................................................... 425

23.1 Mites............................................................................................. 426
23.1.1 Chicken Mite (Dermanyssus gallinae;
Dermanyssidae)............................................................ 426
23.1.2 Northern Fowl Mite (Ornithonyssus sylviarum;
Macronyssidae)............................................................. 427
23.1.3 Tropical Rat Mite (Ornithonyssus bacoti;
Macronyssidae)............................................................. 427
23.1.4 House Mouse Mite (Liponyssoides sanguineus;
Dermanyssidae)............................................................ 427
23.1.5 Grain Mites (Glycyphagus domesticus; Acaridae)....... 428
23.1.6 Straw Itch Mites (Pyemotes ventricosus;
Pyemotidae).................................................................. 428
23.1.7 Chiggers (Eutrombicula alfredugesi;
Trombiculidae).............................................................. 429
23.1.8 Burrowing Mites........................................................... 429
23.1.9 Non-Burrowing Mites................................................... 432
23.1.10 Pneumonyssus caninum; Halarachnidae....................... 433
23.1.11 Snake Mite (Ophionyssus natricis;
Macronyssidae)............................................................. 433
23.1.12 House Dust Mites......................................................... 434
23.1.13 Other Important Mites.................................................. 435
23.2 Ticks.............................................................................................. 435
23.3 Conclusions................................................................................... 437
Further Reading......................................................................................... 438
24 Forensic Acarology................................................................................... 441
24.1 Decomposition of Cadaver........................................................... 442
24.2 Decomposition Stages of Corpse.................................................. 443
24.3 Arthropods and Decomposing Body............................................ 444
24.4 Megnin and Forensic Acarology................................................... 444
24.5 Mite Diversity Found on Carcases................................................ 444
24.6 Role of Freshwater Mites.............................................................. 446
24.7 Conclusions................................................................................... 446
Further Reading......................................................................................... 447
25 Accidental Acarophagy............................................................................ 449
25.1 Plant-Associated Mites................................................................. 450
25.2 Stored Food................................................................................... 451
25.2.1 Stored Wheat Flour....................................................... 451
25.2.2 Prophylactic Measures.................................................. 452
25.3 Dried Fruits................................................................................... 453
25.4 Mushrooms................................................................................... 454
25.5 Soil Mites...................................................................................... 454
Contents xxi
25.6 Cheese........................................................................................... 455

25.7 Meat (Including Cured Fish)........................................................ 456
25.8 Conclusions................................................................................... 456
Further Reading......................................................................................... 457
26 Glossary.................................................................................................... 461
Further Reading......................................................................................... 470
About the Author
Dr M.S. Dhooria was born in 1946 and did his B.Sc. in Agriculture in 1967, and
M.Sc. in Entomology in 1969 from Punjab Agricultural University, Ludhiana
(Punjab), India. In 1980, did Ph. D in Entomology from Indian Agricultural
Research Institute (IARI), New Delhi (India). He was awarded Junior Research
Fellowship in M.Sc., and Senior Research Fellowship in Ph. D studies by the Indian
Council of Agricultural Research (ICAR), New Delhi. He has more than 36 years of
experience in teaching, research and extension in the field of Entomology. He did
research work on: biology, ecology, and management of phytophagous mites.
Taught ‘General and Applied Acarology’ course at post graduate level for many
years. Published more than 125 research publications in different national and inter-
nationally reputed journals. Contributed some book chapters and a booklet on Citrus
Mites as a special publication of ICAR. Attended several national and international
level workshops on different aspects of insects and mites. He is also a life fellow of
Entomological Society of India, New Delhi; Indian Society for the Advancement of
Insect Science, Ludhiana; and Acarological Society of India (ASI), Bangalore.
Remained as Vice President of ASI for two terms. Friendship Forum of India, New
Delhi, awarded him the Certificate of Honour and Gold Medal for his lifetime
achievements in Entomology.
xxiii
Acarology and Its Importance
1
Abstract
Acarology is the study of mites and ticks which are distributed throughout the
world and inhabit almost every ecosystem. Mites are most common arthropod
ectoparasites of vertebrates including humans. They are the largest and diverse
group within the arthropod class Arachnida with over 55,000 described species,
but the number of undescribed species may reach upto 20-fold as mite fauna in
most of the world regions are still poorly explored. Mites and ticks are of great
interest to human beings in a variety of ways and affect the humans as pests of
economic plants including vegetable crops, fruit crops, field and fibre crops, and
different ornamental plants. They also act as parasites of human beings, cattle,
poultry and pets; as pests of stored grains and processed foods; and as parasites
of honeybees. Mite infestations are responsible for economic losses worth bil-
lions of dollars in terms of reduced crop yields, lowered quality of product, and
poor health of mankind. The economic importance of mites and ticks is dis-
cussed to create more interest in these creatures.
Keywords
Acarology • Mites • Ticks
Acarology is a sub-discipline of Zoology and is sometimes included in the field of

Entomology (the study of insects), and Arachnology (the study of spiders and
related organisms). Word acarology is derived from Greek words: akari, means a
type of mite; and logos, meaning knowledge. So acarology is the study of mites and
ticks–the members of the Subclass Acari
Mites are distributed throughout the world and inhabit almost every ecosystem.
They have successfully colonized both terrestrial and aquatic habitats, and are also
very commonly found in the forests and grassland soils, and in temporary accumu-
lations of organic debris. Mites are most common arthropod ectoparasites of verte-
brates, including humans. Follicle mite (Demodex spp.), and scabies mite (Sarcoptes
© Springer Science+Business Media Singapore 2016 1

M.S. Dhooria, Fundamentals of Applied Acarology,
DOI 10.1007/978-981-10-1594-6_1
2 1 Acarology and Its Importance
spp.) are present in association with most people throughout the world (Krantz and
Walter 2009; Evans 1992).
Mites are in fact the first land arthropods, the oldest known so far Late
Devonian. However, because of their small size, the Acari are uncommon in fossil
records. A few are known from the Carboniferous, and more from Tertiary amber
from Canada, Mexico and Europe. Insects and mites inhabited the earth long
before man appeared. They inhabited earth 350 million years ago but man got
inhabited the earth only one million years ago. The first described fossil mite,
Protacarus crani, was found in the Devonian Rhynie Chert of Scotland. Other
fossil mites have been found afterwards, especially bodies included in amber
(Walter and Proctor 1999).
Mites or Acari are the largest and diverse group within the arthropod Class
Arachnida (Subphylum Chelicerata, and Phylum Arthropoda) with over 55,000
described species. Even this number seems misleading since it is estimated that only
5–10 % of all the mite species have been formally described, but the number of
undescribed species may reach upto 20-fold as mite fauna in most of the world
regions are still poorly explored. Acari are the largest and more diverse group in the
class Arachnida. In contrast with other arachnid groups such as spiders and scorpi-
ons, mites are of smaller size 0.1–30 mm in length. Mouthparts of mites and ticks
are specialized for piercing tissues and sucking out the contents (Gupta 2012; Hoy
2011).
Mites and ticks are of great interest to human beings in a variety of ways. They
directly affect the humans as pests of different economic plants including vegetable
crops, fruit trees, field and fibre crops, and different ornamental plants (mostly in
greenhouses); as parasites of human beings, cattle, poultry and pets; as pests of
stored grains and processed food, mushrooms and cured- fish; and as pests of hon-
eybees. Mite infestations are responsible for economic losses worth billions of dol-
lars in terms of reduced crop yields and lowered quality of the produce (Gerson
et al. 2003). The economic importance of mites is briefly discussed here under dif-
ferent heads.
Plant Feeding Mites About 7000 species of plant feeding mites are known world-
wide, and about half of them belong to the Superfamily Eriophyoidea (gall, bud,
rust mites etc.). Remaining half are distributed within the Superfamily Tetranychoidea
(spider mites, flat mites, and peacock mites), and the lesser known bulb mites of the
family Acaridae.
Plant feeding mites play an important role as pests of vegetable crops, fruit trees,
ornamental plants (both under field and protected cultivation), fibre crops, forage
crops, oilseed crops and timber trees. Under many situations, lack of information
about the correct identity of mites, and inadequate knowledge regarding their biol-
ogy and ecology, have hampered our ability to combat these pests. Small size and
their cryptic appearance makes them difficult to detect, and thus infestations are
many times overlooked. Once established in a new area, certain biological charac-
teristics allow their rapid escalation to pest status. High rate of fecundity, short life
1 Acarology and Its Importance 3
cycles, a myriad of their dispersal techniques, and adaptability to diverse ecological

conditions are the main reasons for their success under different agroecosystems
(Hoy 2011). Besides, the great increase in world trade has set the stage for poten-
tially devastating situations that may threaten the world’s agroecosystems. Plant
feeding mites like two-spotted spider mite Tetranychus urticae Koch, citrus mites
Panonychus ulmi (Koch) and Panonychus citri (McGregor); and flat mite Brevipalpus
phoenicis (Geijskes) are amongst the most serious pests under diverse conditions.
Some eriophyid mites and spider mites are the vectors of several viral diseases of
some economic crops and cause economic losses in their production.
Controller of Weeds Weeds compete cultivated plants for water and for nutritive
elements. The skeleton weeds gall mite, Aceria chondrilla (Canestrini) feeds spe-
cifically on troublesome weed, Chondrilla juncea, and reduce the damage from
weed by controlling their further spread in new areas. Some other eriophyid mties
have also been reported specifically feeding on certain other weed species resulting
in reduced use of herbicides and pesticides (Ochoa et al. 1994).
Soil Mites Soil mites are typical representatives of soil microfauna inhabiting soil
pores and other minute soil spaces. They inhabit primarily in upper layers of soil,
and together with the collembolans are the important representatives of aerobiotic
microfauna. In grasslands (meadows, pastures etc.) and forest habitats, their repre-
sentation is dominant among different microfauna. The soil mites are mainly phy-
tophagous, fungivorus and saprophagous forms; besides predaceous and parasitic
mites. Soil mites have exceptional importance in the circulation of substances in soil
and are more resistant to desiccation in contrast to other soil microfauna. Soil or
oribatid mites are also known as beetle mites, and are especially abundant in humus
rich soil. Significance of soil mites in processing the organic matter into humus, for
the life of the soils as an organic component of ecosystem, in which they help to
maintain an equilibrium and increase the fertility of soil. Several soil mites may
play an important role in transmitting different pathogens to man. Chiggers – larvae
of family Trombiculidae, transmit rickettsia responsible for scrub typhus, and are
typical microcavernicolous inhabitants of the soil penetrating the deeper soil layers.
Some oribatid mites are utilized as intermediate hosts by cestodes of sheep and
other ruminants, like hares and rabbits (Baker and Wharton 1952; Evans 1992;
Krantz and Walter 2009; Walter and Proctor 1999).
Predatory Mites Diverse ecological relationships among mites may be utilized in

biological control of phytophagous mites under different situations such as green-
houses/screenhouses etc. Different predaceous mites belonging to family
Phytoseiidae are considered as efficient predators of pest mites resulting in com-
paratively lesser use of pesticides. Commercial insectaries produce these mites for
biological control of pest mites on tomato, ornamental plants (including roses) par-
ticularly under protected cultivation and under organic cultivation. Some tydeid
mites have been reported feeding on hyphae of fungus that cause powdery mildew
disease. Some of the predatory mites have been reported to control successfully
aphids, thrips, and whiteflies. Straw itch mite. Pyemotes tritici (Fossat & Montagne),
have been used as a biocontrol agent against Anagastra kuhniella.which damages
stored grains and flour (Gerson et al. 2003; Gupta 2012; Hoy 2011; Zhang 2003).
Stored Products Mites living originally in small numbers on seeds, plants, decay-
ing vegetation, and in nests of mammals, are gradually introduced to all types of
stored products due to mans’ economic activities (food storage). Mites attack either
stored unprocessed products (grains, vegetables, fruits, meat, bulbs, hay etc.) or
prepared products (flour or flour products, cheese, dried fruits, cured-fish etc.).
Stored mites primarily belong to family Acaridae; and infestation of stored products
by mites pose great problems all over the world. Besides damaging grains, they
affect the quality of grain products. Even the stored bulbs and corms of different
ornamental plants are badly affected by mites. The mites consumed together with
infested food cause intestinal and urinary problems in animals (Gerson et al. 2003;
Hoy 2011).
Acariasis ‘Acariasis’ in animals is caused by a variety of mites. The zoonotic spe-

cies include Sarcoptes scabiei (DeGeer) – cause of sarcoptic mange (scabies) in
humans, and more than 100 other species of mammals and marsupials. Humans can
also be infested by the scabies mites of dogs, pigs, cattle, goats, sheep, hares, buf-
falo, camels, and some wild or zoo animals. Many members of family Psoroptidae
affect only domestic animals and result in conditions known as ‘psoroptic mange’.
Several species of chiggers or harvest mites (family Trombiculidae) are parasites of
different animals or humans. Hundreds of food storage items (including grains and
insect cultures) are faced by certain allergic reactions and pruritis caused by differ-
ent mite infestations. The bites of Dermanyssus gallina (DeGeer), Liponyssoides
sanguineus Hirst, Ornithonyssus bacoti (Hirst), O. sylviarum (Canestrini &
Fanzago), and O. bursa can be painful or pruritic, and may cause irritation and
localized dermatitis (Krantz and Walter 2009; Walter and Proctor 1999).
Mushrooms and Mites Some mites are important pests of cultivated fungi espe-
cially mushrooms. Linopodes spp. (family Eupodidae), and Tyrophagus spp. (fam-
ily Acaridae), Tarsonemus spp. and Pygmephorus spp. occur in large mumbers in
commercial mushroom houses where they often destroy whole crop of mushrooms
particularly when improperly conditioned compost is used and poor sanitary condi-
tions are followed.
Flavour Enhancer for Cheese For centuries French and German cheese makers
have used mites, Acarus siro and Tyrophagus casei, to refine the taste of cheese.
However, it is not properly known regarding the production of typical sour taste,
loved by fans resulting from mite infestation. Quarantine authorities of USA and
other countries, however, do not allow such cheese for import.
1 Acarology and Its Importance 5
House-dust Mites and Allergy Dermatophagoides spp. are the primary source of
allergens in house dust. These mites are commonly known as house-dust mites, and
they produce many proteins that induce allergic responses in sensitive persons.
House-dust allergy may take the form of respiratory distress or skin irritation. Mites
typically inhabit beds, chairs, carpets, mattresses and pillows in houses, and their
shed skins and faeces provide the bulk of the allergens in house-dust extracts
(Colloff 2009).
Beekeeping and Mites Ectoparasitic mites like Varroa destructor Anderson &
Trueman, V. jacobsoni Oudemans, and Tropilaelaps clareae Delfinado & Baker;
and endoparasitic mite Acarapis woodi (Rennie) which infest trachea, are also very
important natural enemies of honeybees. Sometimes these mites cause lot of devas-
tations of bee colonies resulting in reduced pollination of crops, and lowered yields
of honey. Bee mites are very difficult to control because of toxicity of pesticides to
bees and development of resistance to pesticides in parasitic mites (Krantz and
Walter 2009).
Crime Solvers The precise date of ones’ death can be determined by taking sam-
ples of the mites that develop on the victim. The establishment of these mites follow
a well- defined chronology. Certain species of mites invade the human corpse faster
than others. Therefore, the Acari can be used for the estimation of postmortem inter-
vals. Macrocheles spp. are abundant in first stage of decomposition while
Tyroglyphidae and oribatid mites which are consumers of dry human skin appear
later. Moreover, as some Acari have a precise geographical distribution they may
serve to provide valuable supporting data to determine the location of the body and
the scene of the crime at the time of death. Similarly the characteristics of the aca-
rines found on the body of drowned people can be useful to determine the time of
submersion of a corpse. So mites can sometimes provide vital clues about murder
and in solving the cause of death.
Poultry Pests The poultry red mite, Dermanyssus gallinae (DeGeer), is regarded
as the most important ectoparasite of laying hens in organic or conventional egg
production. These mites can cause severe anaemia and associated mortality. Mite
infestations may cause staining of egg shell surface and thus reduce their market
value. Even low mite infestation can irritate hens to the extent that they refuse to use
the hen house or rest on the perches. Serious infestation of poultry mites, D. galli-
nae, may result in great reduction in terms of eggs and meat yield. Red mites may
also act as a vector for numerous pathogens of medical and veterinary importance
(Evans 1992; Krantz and Walter 2009).
Medical and Veterinary Importance Ecto-, and endoparasitic mites attack both
vertebrates and invertebrates. Most endoparasitic mites live in the respiratory tract
of vertebrates. Ectoparasitic mites are the most common arthropod parasites of ver-
tebrates (including human beings). Human follicle mite, Demodex spp. are present
on most people throughout the world. Parasitic mites Sarcoptes scabiei (DeGeer)
causes ‘scabies’ in humans and ‘mange’ in wide range of domestic and wild mam-
mals throughout the world. Mites infest the skin in a variety of mechanisms like
mechanical trauma, contact through their secretions, retained mouthparts, or with
their living/dead bodies (irritant or allergic contact reactions). Storage mites
Glycyphagus destructor Schrank, play an important role in causing ‘atopic derma-
titis’ (atopic eczema) in children (Baker and Wharton 1952).
Ticks Ticks are actually a specialized group of mites, and share many features with
the mites. In general they are larger than most mites, ranging from 0.6 mm to 1 cm
in length or even more in diameter when fully engorged with blood. All ticks are
blood-sucking parasites (ER 2). They are found in most parts of the world but are
generally limited to those habitats (wood, tall grass and shrubby vegetation) which
are most frequented by their hosts, as the ticks climb onto tops of plants and estab-
lish contact with a passing host. Ticks are obligate ectoparasites of cattle, poultry
and human beings. Dermacentor andersoni Stiles, causes paralysis in cattle; D
variabilis (Say), is of nuisance value to tourists and rural people; D. albipictus
(Packard) infests deer and neglected range of horses, and may act as an important
source of nuisance to game animals confined in parks. Otobius megnini (Duges)
causes ‘toxicosis’ of cattle. Rhipicephalus sanguineus (Latreille) commonly parasit-
izes dogs, and is a pest in kennels and homes. Direct feeding of ticks on cattle
causes blood loss as high as one litre per cattle head in one season. A high infesta-
tion may cause anaemia in cattle. Ticks sucking in clusters may cause additional
bleeding from the wounds cut by cluster feeding. The damaged skin permits the
entry of their infectious germs which may cause local or general afflictions of
domestic animals. There may be decrease in milk yield from 25 to 40 % in tick-
infested cows. Ticks also serve as vectors of animal diseases caused by arboviruses,
rickettsiae, bacteria, spirochetes, protozoa, parasitic worms etc. (Krantz and Walter
2009; Walter and Proctor 1999).
Considering great economic importance of mites and ticks (both Acari) in rela-
tion to agriculture, horticulture, animal husbandry and human-beings, the informa-
tion regarding different aspects of studies on them are found in literature published
in different scattered journals, book chapters, and other reports in different lan-
guages and can not be readily used by students or other interested readers. So it was
thought necessary to present the information in different chapters on different
aspects in the form of a book. Most important information gathered from different
sources are included in this book in a way that it may stimulate readers to do more
studies in this area. For knowing more about the subject, further reading including
salient sources of study are given at the end of each chapter. Additionally Electronic
Resources (ER) and weblinks are provided for better understanding mites and ticks.
Further Reading 7
Further Reading
Baker EW, Wharton GW (1952) An introduction to acarology. Macmillan, New York, 464 pp
Colloff MJ (2009) Dust mites. CSIRO Publishing, Collingwood, 496 pp
Evans GO (1992) Principles of acarology. CAB International, Wallingford, 563 pp
Gerson U, Smiley RL, Ochoa R (2003) Mites(Acari) for pest control. Blackwell Scientific, Oxford,
521 pp
Gupta SK (2012) Handbook: injurious and beneficial mites infesting agri-horticultural crops in
India and their management. Nature Books India, New Delhi, 362 pp
Hoy MA (2011) Agricultural acarology: an introduction to integrated mite management. CRC
Press, Boca Raton, 410 pp
Krantz GW, Walter DE (eds) (2009) A manual of acarology, 3rd edn. Texas Tech University Press,
Lubbock, 807 pp
Ochoa R, Aguilar H, Vargas C (1994) Phytophagous mites of Central America: an illustrated guide.
Centro Agronomico Tropical de Investigacion y ensenanza (CATIE), Turrialba, 220 pp
Walter DE, Proctor HC (1999) Mites, ecology, evolution and behaviour. CAB International,
Wallingford, 322 pp
Zhang ZQ (2003) Mites of greenhouses: identification, biology and control. Wallingford, CAB
International, 235 pp
Electronic References (ER)
1. https://www.youtube.com/watch?v=_BPE9-JymTA Arachnids: the different kinds

2. https://www.youtube.com/watch?v=kAQNoEp8P0o&feature=iv&src_vid=_BPE9-
JymTA&annotation_id=annotation_2302628687 Ticks up close
Historical Account of Acarology
2
Abstract
Awareness about the mites and ticks dates back to ancient Egypt (1550 BC) and
was continuously demonstrated throughout the writings of major Greek scholars;
however, the science of acarology originated in the eighteenth-century Europe.
Pliny the Elder (23–79 AD) reported ticks as the foulest and nastiest creatures.
References to mites and ticks have also been found in the early writings of
Hippocrates and Aristophanes. Greek terms Kroton or Kynoraistis for ticks and
A-Kari (without head) had been used by Aristotle in describing about non-ixodid
ticks. During medieval times, mites were commonly referred to as lice, beesties
or little insects. The rediscovery of the term Akari or Acari appears to have
occurred about 1650, but the term acarology began to appear regularly in the
literature. Carl von Linnaeus used the generic word Acarus in the first edition of
the Systema Naturae published in 1738. Historical account of acarology is dis-
cussed into three parts, i.e. prior to 1850 AD, between the period 1851 and 1950
and the developments from 1951 till today. A brief account of important books
published, history of formation of acarological societies at regional and interna-
tional levels and important journals published in acarology are discussed. Interest
in acarology is rapidly developing because of the role of mites and ticks and their
associations on different crops, storages and vectors of diseases and their impact
as parasites/pests on human beings, domestic animals and poultry.
Keywords
Acari • Acarina • History • Linnaeus
Awareness of the mites and ticks dates back to ancient Egypt (1550 BC) and was
continuously demonstrated throughout the writings of the major Greek scholars;
however, the science of acarology originated in the eighteenth-century Europe.
Historical account has been divided into three parts, i.e. prior to 1850 AD, between
1851 and 1950 and the developments from 1951 till today (Prasad 1982).

DOI 10.1007/978-981-10-1594-6_2
10 2 Historical Account of Acarology
2.1 Developments Prior to 1850 AD
The oldest reports are about a tick in the 1550 BC (Egyptian Ebers Papyrus), occur-
rence of ticks in the 850 BC in Ulysses’ dog (Homer) and a record of mites on
cheese in the 350 BC (Aristotle). Subsequently after another 500 years, a mite para-
site of locusts (probably Eutrombidium) has been reported in De Animalibis Histori
Libri. Pliny the Elder (23–79 AD) reported ticks as the foulest and nastiest crea-
tures. References to mites and ticks have been found in the early writings of
Hippocrates and Aristophanes.
Greek terms Kroton or Kynoraistis for ticks and A-Kari (without head) had been
used by Aristotle in describing about non-ixodid ticks. Another possible derivation
of the word Acari may have been from the Greek word akares, which means small
or short. During medieval times, mites were commonly referred to as lice, beesties
or little insects. As in Aristotle’s time and until relatively recently, ticks were con-
sidered as an entity separate from mites and were referred to as ticia (old English)
or Zecken (in Middle High German). The old English word mite, meaning very
small, may well have been derived from the ‘widows’ mite – an early Flemish coin
of very small size (Walter and Proctor 1999).
Rediscovery of the term Akari, or Acari, appears to have occurred about 1650,
but it was not until the early twentieth century that the term acarology began to
appear regularly in the literature (Krantz 1996). Carl von Linnaeus used the generic
word Acarus in the first edition of the Systema Naturae published in 1738. Later,
Linnaeus named the type species, Acarus siro, in the tenth edition of Systema
Naturae published in 1758. In this edition, 30 mite species reported all under the
genus Acarus. In this way Linnaeus did the ground work for the field of systematic
acarology. During the one hundred years that followed Linnaeus, several schemes
of higher classification of mites and ticks were introduced (DeGeer 1778; Leach
1815; Duges 1839). Many newly recognized mites and ticks were reported in this
period. Oudemans’ publication Kritish Historisch Overzicht der Acarologie that
appeared in three volumes (1926, 1929 and 1936–1937) included all acarological
contributions made before 1850. But some early naturalists after Linnaeus many
times mixed up the genera of mites with diverse groups such as crustaceans, the
common sea spider, small beetles and harvestmen.
William Elford Leach (1790–1836) deserves credit for recognizing mites as an
arachnid order Monomerostomata Leach 1815. But 4 years later, Leach (1819)
replaced his ordinal name with a class Acari, although the credit for naming the taxon
is given to Christian Ludwig Nitzsch (1782–1837) who introduced essentially the
same name a year earlier as Acarina Nitzsch, 1818. By this time the ‘mites’ were now
on their way to bring a definable and recognizable group (Krantz and Walter 2009).
2.2 Progress During 1851–1950 AD
The emergence of acarology as a modern science during the late nineteenth and
early twentieth centuries was centred primarily in Europe and North America.
Michael (1884) placed all Acari in two groups based on the basis of respiratory
2.3 Developments Since 1951 11
systems: (i) Acari, Atracheata (including Glycyphagus, Histiostoma, Phytoptus,

Demodex and Sarcoptidae), and (ii) Acari, Tracheata (including Prostigmata,
Oribatidae, Gamasidae, Ixodidae, Tarsonemidae and Myobiidae). By this time
members of mites and ticks were known by the names: Acarina, Acariens, Acaros
and Acarida.
Grandjean (1882–1975) did an excellent work on mites regarding their systemat-
ics, morphology, phylogeny and ontogeny. Historic contributions during the above
period were also made by Kramer (1877), Canestrini (1891), Banks (1904), Reuter
(1909), Sig Thor (1929), Vitzthum (1940–1943) and Tragardh (1946) mostly regard-
ing description of new taxa and classification of present day Acari. Many valuable
contributions of the above scientists towards acarine systematics and classification
proved instrumental in establishing acarology as a discrete discipline.
After World War II, intensive studies on mite-transmitted diseases such as scrub
typhus and tick-borne haemorrhagic fever were reported. Following the introduc-
tion of several organochlorinated and organophosphate pesticides for the control of
different insect and mite pest problems of agricultural and veterinary importance,
the impact of these pesticides in disrupting the natural enemies of mites on agricul-
tural crops was more thoroughly investigated. Several new mites and ticks were
found and their damage was investigated in different locations.
2.3 Developments Since 1951
The eminent scientists, EW Baker and GW Wharton, investigated thoroughly diver-

sity of plant-associated and animal parasitic mites. Both of them published jointly a
monumental and very useful book: An Introduction to Acarology in 1952. The above
text included briefly all the most important contributions towards systematics and
classification of Acari and provided the base of modern day acarology. Many biolo-
gists were attracted to this branch of science after the publication of the above text.
Baker and coworkers published in 1958 another very useful book: Guide to Families
of Mites. The efforts of different scientists led to the development and production of
some more books, establishment of acarological societies in different countries and
production of various electronic publications and websites, which have put acarol-
ogy on very sound footing. In many universities, General and Applied Acarology
course is being taught at both graduate and postgraduate levels, and many shining
students are opting this subject for their thesis/dissertation studies. To stimulate
further interest in the subject of acarology, V. Prasad – an eminent acarologist – pub-
lished a book History of Acarology in 1982 which included historical account of
acarological investigations made in 22 different countries. The Acarological Society
of America (ASA) honoured EW Baker and GW Wharton for their lifetime contri-
butions in the field of acarology, at a specially organized symposium, Acari: Life
history, and Reproductive Patterns, and considered them as the Fathers of Acarology
(Prasad 1982; Lindquist 1984; Lindquist 2006).
2.4 OSU Acarology Development Programme
At Duke University (USA), Dr George W. Wharton established a section ‘The

Institute of Acarology’ in 1951. Then Dr GW Wharton and Dr Edward W Baker
together conducted ‘Acarology Summer Programme’ for the young acarologists at
Ohio State University, at Columbus (Ohio), in the same year when Dr GW Wharton
joined as Chairman of Department of Zoology and Entomology. Since then this
course is being regularly conducted annually for training interested acarologists in
the field of agriculture, medical, veterinary and soil acarology. Many present-day
acarologists have got training from the above programme and produced useful find-
ings which have made the subject of acarology as an important discipline (ER1).
A brief history of acarological developments in Hawaii (USA) and Australia is
also reported in literature (ER2 and ER6). An overview of the story of French aca-
rologists (ER4), and history of acarology in Egypt (ER5), is also available. Trends
of publications made in the last 150 years in different aspects of acarology in differ-
ent journals are also reported (ER3). The information pertaining to different acaro-
logical societies, journals devoted to acarology and important books published in
the field of acarology are given below.
2.5 Acarological Societies
1. International Congress of Acarology (ICA)

http://www.acarology.org/ica/
Meeting of the International Congress of Acarology is held after every 4 years,
and publications in book form are released. ICA also compiles a directory of
acarologists of the world. First such meeting was held in Colorado (USA) in
1963, and the 14th International Congress of Acarology was held at Kyoto
(Japan) in 2014. Details about ICA congresses held and proceedings published
thereof are given in Table 2.1.
2. Acarological Society of America (ASA)
https://www.facebook.com/AcarologicalSocietyofAmerica/
https://sites.google.com/site/acarologicalsociety/home
ASA meetings are held in coordination with the Entomological Society of
America (ESA). The society also publishes a directory of member acarologists,
news and meetings. ASA also publishes on irregular basis its newsletter (ER2).
3. Acarological Society of India (ASI)
ASI was founded in 1973. The Indian Journal of Acarology, later renamed as
Journal of Acarology; and Acarology Newsletter published regularly by the
society till 2000 (Vol. 16 and 17), but subsequently thereafter published
irregularly.
4. Systematic and Applied Acarology Society (SAAS)
(http://www.nhm.ac.uk/hosted_sites/acarology/saas/)
SAAS was founded in 1996 and published Systematic and Applied Acarology
Journal. SAAS also published Special Publications – a rapid publication for
2.5 Acarological Societies 13
Table 2.1 International Congress of Acarology

Congress Held at Period President Proceeding published
1st Fort Collins, 2–7 Sept. 1963 G. W. Wharton Proceedings published as:
Colorado, USA Proceedings, in a special
number of Acrologia.
Acarologia, 1964–1966
(h.s.):1–439
2nd Sutton, Bonington 19–25 July 1967 T. E. Hughes Proceedings by
England (UK) Akademiai Kaido,
Budapest, in 1969, 652 pp
3rd Prague (then 31 Aug.–6 Sept, B. Rosicky Proceedings of the 3rd
Czechoslovakia) 1971 International Congress of
Acarology 1971, 837 pp
4th Saalfelden, 12–19 Aug. R. Schuster Proceedings of the 4th
Austria 1974 International Congress of
Acarology 1979,752 pp.
(ISBN 963 05 16950)
5th East Lansing, 6–12 Aug. 1978 E. W. Baker Recent advances in
Michigan (USA) Acarology Vol I 631 pp.
(ISBN 0-12-592201) &
Vol. II 569H. (ISBN
0-12-592201-9)
6th Edinburgh, 5–11 Sept. 1982 G. O. Evans Acarology VI, Vol:
Scotland 645645 pp. (ISBN
1984-085312-603) and,
Vol. II (ISBN
0-85312-604-6)
7th Bangalore, India 3–9 Aug. 1986 G. P. Channa Progress in Acarology
Basavanna 1988, Vol. I: 523 pp.
(ISBN 81-204-0346-0),
Vol. II: 484 pp. (ISBN
81-204-0347-9)
8th Ceske Budejovice 6–11 Aug. 1990 B. Rosicky Modern Acarology,
(Czechoslovakia) 1991 Vol. I: 651 pp.
(ISBN 80-200-0364-9)
and Vol. II: 779 pp.
(80-200-0369-9)
9th Columbus, Ohio 17–22 July 1994 D. E. Johnston Proceedings: Acarology
(USA) IX 1996, 718 pp
10th Canberra, 5–10 July 1998 R. B. Halliday Proceedings: Acarology
Australia Proceeding of the 10th
International Congress
1998, 657 pp. (IISBN
0-643-06658-6)
11th Merida, Yucatan, 8–13 Sept. 2002 Tila M. Perez Acarology XI:
Mexico Proceedings of the
International Congress of
Acarology, 2007, 726 pp.
(ISBN
978-970-32-4451-5)
(continued)
Table 2.1 (continued)

Congress Held at Period President Proceeding published
12th Amsterdam, the 21–16 Aug. Maurice Trends in Acarology:
Netherland 2006 W. Sabelis Proceedings of the 12th
International Congress
2010, 566 pp. and (ISBN
978-90-481-9836-8 and
12th e-ISBN
978-90-481-9837-5)
13th Recife, Aug. 2010 G. J. Demores Acarology XIII:
Pernambuco and Heather Proceedings of the 13th
Proctor International Congress
(2011) ISBN: 978-1-
86977-804-0 (online
edition)
14th Kyoto, Japan 14–18 July 2014 H. Amano Proceedings 14th
International Congress of
Acarology (2016)
Flechtmann (2011)
short papers and monographic works – which is now merged with the journal in
2012. It also publishes Acarology Bulletin, a newsletter of the society. It also
issues books of special interest to members and maintains an online acarologi-
cal e-reprint library for acarologists around the world. Anyone interested in the
study of mites and ticks is welcome to join SAAS.
5. African Acarology Association (AAA)
http://www.nhm.ac.uk/hosted-sites/acarology/saas/Hosted/aaa/index.htm
AAA was founded in 1998 in South Africa. It holds symposia in acarology
(regarding different topics) regularly after 3 years duration. The first sympo-
sium was held in 1998, and the fifth symposium was held at Livingstone,
Zambia, in 2011. Abstracts of the third, fourth and fifth symposium are avail-
able on the website of AAA.
6. The Acarological Society of Japan (ASJ)
(http://en.acarology-japan.org/)
ASJ was founded in 1973 and replaced the earlier Japanese Association of
Acarology which was established in 1973. ASJ publishes semi-annually a
research journal, Journal of Acarological Society of Japan. ASJ also holds
annual meetings and conducts symposiums regularly. The 14th International
Congress of Acarology (ICA) was held at Kyoto, Japan, in 2014.
7. European Association of Acarologists (EURAAC)
(http://euraac.webs.upv.es/)
EURAAC publishes an irregular newsletter and organizes symposia in Europe
at regular intervals (ER4). Proceedings of different symposia have been pub-
lished in book form as given in Table 2.2.
8. The Egyptian Society of Acarology (ESA)
(http://www.esaeg.org/)
ESA was founded in 2005 and is publishing annually a journal, Acarines (ER5).
2.5 Acarological Societies 15
Table 2.2 EURAAC symposiums

Symposium Place held Dates Editors Proceedings published as
1st Graz (Austria) 1988 Schuster, R and The Acari: Reproduction,
Murphy P. W. Development and
life-history strategies,
pp. 564. Chapman and
Hall, London, 1991
2nd Krynica 1992 Kropczynska, D; The Acari: Physiological
(Poland) Boczek, J; and and Ecological aspects of
Tomczyk, A. Acari-Host relationships,
Pp. 698. Dabor Warsaw,
1995
3rd Amsterdam 1996 Bruin J, Van der Geest Ecology and Evolution
(Netherland) LPS and Sabelis MW of the Acari. Pp 677,
Kluwer Acad.
Publishers, Dordrecht
1999
4th Sienna (Italy) 2000 Bernin, F; Nannelli, Acari Polygeny and
R.; Nuzzaci, G.; and Evolution (Adaptations
de Lillo, E. in Mites & Ticks). Pp.
472, Kluwer Acad.
Publishers Dordrecht,
2002
5th Berlin 2004 Weigmann G; Alberti, Acarine Biodiversity in
(Germany) G; Wohltmann, A; and the Natural and Human
Ragusa, S. Sphere. Pp. 765.
Phytophaga XIV, 2004
(2005)
6th Montpellier 2008 Bertrand, M; Kreikter, Integrative Ecology. Pp.
(France) S.; Mc Coy, K. D.; 492, Euraac
Migeon, A.; Navajos,
M.; M. S.; and Vial, L.
7th Vienna (Austria) 2012 K. D. Migeon, A; Acari in a changing
Navajos, M: Tyl world. Acarologia 53
(2),2013
9. The Acarological Society of Iran (ASI)

(http://www.acarology.ir/)
ASI was founded in 2008 and started publishing semi-annually – Persian
Journal of Acarology – from January 2012.
10. Sociedad Latinoamericana de Acarologia (SLA)
(www.slaacarologia.com)
SLA was established for South American acarologists in 1994.
11. Societe Internationale Des Acarologues De Langue Francaise (SIALF)
(www.nhm.ac.uk/hosted-sites/acarology/saas/hosted/sialf/index.htm)
SIALF aims to promote research in acarology by exchanges of information
among French-speaking members to support the growth of acarology. The soci-
ety also organizes ‘International Courses of Acarology’ regularly.
2.6 Journals in Acarology
1. International Journal of Acarology (IJA). [ISSN: 0164-7954 (print); and ISSN:

1945-3892 (online)]
(http://www.tandfonline.com/loi/taca20#. VPLsJHyUfp8)
IJA was published biannually from 1975 till 2010. Taylor and Francis, from the
United Kingdom, is publishing this journal from January 2011. Since its incep-
tion, this journal under the dynamic editorship of Dr V. Prasad – an eminent
acarologist – honoured erstwhile and present distinguished acarologists through
different activities published in IJA.
2. Acarologia (ISSN: 0044-586-x (print); 2107-7207 (online))
http://www1.montpellier.inra.fr/CBGP/acarologia/latest.php
Marc Andre and Francois Grandjean founded this journal in 1959. This is the
first and longest publishing acarology journal published quarterly and is devoted
exclusively to acarology. From 2010, the online version is free through open
access. A paper version is also available but through subscription.
3. ACARINA
http://insects.ummz.lsa.umich.edu/acarina/
Russian Journal of Acarology (ISSN: 2221-51515 (online); and 0132-8077
(print).
Published by KMK Scientific Press © on behalf of the Zoological Museum of
Moscow University, since 1993. The journal is published in English; free PDFs
are available.
4. Acarology Bulletin (ISSN: 1361-8091)
http://www.nhm.ac.uk/hosted_sites/acarology/saas/ab.html
http://biotaxa.org/saa/
Acarology Bulletin is a newsletter published quarterly by ‘Systematic and
Applied Acarology Society’. Contents of online issues are available from 1996
to 2008; thereafter, it merged with its sister journal and is publishing online since
1912 (DOI: http://dx.doi.org/10.11158/saasp.23.1.
5. Experimental and Applied Acarology (ISSN 0-168-8162)
http://www.springer.com/life+sciences/entomology/journal/10493
The journal started publishing in 1985 and is publishing regularly. About 114
issues are available electronically (1997–2011).
6. Indian Journal of Acarology (ISSN: 970-1400)
(www.acarology.in/htm/ijoa.htm)
Published semi-annually from 1976 to 2000 and is publishing thereafter
irregularly.
7. Persian Journal of Acarology
(http://www.acarology.ir/online/20issue.htm)
A quarterly journal published by the Acarological Society of Iran since 2012 and
is publishing regularly.
8. ACARINES
http://www.esaeg.org/acarines.htm
Published biannually by the Egyptian Society of Acarology, since 2007.
2.7 Important Books Published 17
2.7 Important Books Published
Baker EW, Wharton GW. 1952. An introduction to Acarology, MacMillan,

New York, 465 pp
Chillar BS, Gulati R, Bhatnagar P. 2007. Agricultural Acarology. Daya Publishing
House, Delhi (India), 353 pp
Evans GO, Sheals JG. Macfarlane D.1961. Terrestrial Acari of the British Isles,
Vol.1, Adland and Son. Barthobnew Press, Durking, England
Evans GO. 1992. Principles of Acarology. CAB International, Wallingford,
UK.563 pp
Gerson U, Smiley RL. 1990. Acarine Biocontrol Agents: An Illustrated Key and
Manual. Chapman and Hall, London, 174 pp
Gerson U, Smiley RL, Ochoa R. 2003. Mites (Acari) for pest control. Blackwell
Science Ltd, Cambridge, 539 pp
Helle W, Sabelis MW. 1985. Spider mites: Their Biology, Natural Enemies and
Control. World Crop Pests, Vols I A and I B, Elsevier, Amsterdam
Harvey MS. 1998. The Australian Water Mites: A Guide to Families and Genera
CSIRO Publishing, Collingwood, Victoria
Jeppson LR, Keifer HH., Baker EW 1975. Mite pests of economic plants. University
of California Press, Berkeley, 614 pp
Krantz GW, Walter DE., eds. 2009. A Manual of Acarology. Third Edition, Texas
Tech. University Press, Lubbock, Texas, USA
Lindquist EE; Sabelis MW, Brun J., eds. 1996. Eriophyoid mites: Their biology,
natural enemies and control. World Crop Pests, Vol. 6. Amsterdam, Elsevier
Prasad V, ed. 1982. History of Acarology. Indira Acarology Publishing, Michigan,
USA
Walter DE, Proctor HC. 1999. Mites-Ecology, Evolution and Behaviour University
of New South Wales Press and CAB International, Sydney and Wallingford,
322 pp
Zhang Z-Q 2003. Mites of Greenhouses: Identification, Biology and Control. CABI
Publishing, CAB International, Wallingford, UK, 244 pp
In addition to the information given above, there may be several regional acaro-
logical associations and many other journals publishing important acarological
investigations regularly throughout the world. Many books, monographs and tech-
nical bulletins/reports on different fundamental and applied aspects of acarology are
being published by agricultural, medical and veterinary universities of different
countries (Zakhvatkin 1952; Zhang 2011, 2014). So interest in acarology is picking
up gradually as the role of mites and ticks is being realized because of their associa-
tions on different crops, storages, vectors of diseases and their impact as parasites/
pests on human beings, domestic animals, zoo animals, poultry, etc. Recently many
molecular studies on understanding higher levels of taxonomy and even at species
level have been made. The results obtained from these studies are proving very use-
ful tools even where morphological studies are not providing true information of
relationship at different levels (Walter and Proctor 2010). Studies on genetic bar
coding of different mites and tick species have also been reported.
2.8 Conclusions
Survey of literature reveals awareness about mites and ticks in ancient Egypt (1550
BC), but the science of acarology originated in the eighteenth century alone.
References to the term Akari or Acari began appearing regularly in the literature,
and Linnaeus used the generic word Acarus for mites in his first edition of Systema
Naturae published in 1758. Later in tenth edition of Systema Naturae published in
1758, Linnaeus included 30 species of mites all under genus Acarus. Thereafter,
references to mites and ticks regularly appeared and these organisms were reported
under several schemes of higher classification. But the science of acarology (scien-
tific study of mites and ticks) started emerging as a modern science during the late
nineteenth and early twentieth centuries; however, during this period the mites and
ticks were known by the names: Acarina, Acariens, Acaros and Acarida. Historic
significant contributions that were made by Kramer (1877), Canestrini (1891),
Banks (1904) and Vitzthum (1940–1943) proved instrumental in establishing aca-
rology as a modern science. In 1952 a very useful book An Introduction to Acarology
by Baker EW and Wharton GW appeared, and this stimulated interest in the science
of acarology globally. Thereafter, many useful publications started appearing on
taxonomy, ecology and other applied aspects from different regions of the world.
This led to publication of the journal Acarologia in 1957; this journal published
information exclusively on mites and ticks. Interest in acarology further resulted in
formation of different societies at international, regional and country levels. Some
more journals exclusively publishing information on mites and ticks and some good
books were written by eminent scientists in their field of specialization, which fur-
ther helped in the development of this discipline. At the international level, the
‘International Congress of Acarology’ was founded which organizes regularly after
4 years interval the meetings of the congress in different countries. The first con-
gress was held in the USA, and the 14th congress was held in 2014 in Japan.
Proceedings of the congress are being published regularly in book form. At regional
and country levels, many associations were formed which are conducting sympo-
siums at different intervals.
Further Reading
ChannaBasavanna GP, Nangia N (1988) Acarology in developing countries: glimpses and pros-
pects. In: ChannaBasavanna GP, Viraktamath CA (eds) Progress in acarology, vol II. Oxford
and IBH, New Delhi, pp 469–473
Evans GO, Till WM (1979) Mesostigmatid mites of Britain and Ireland (Chelicerata: Acari:
Parasitiformes): an introduction to their morphology and classification. Trans Zool Soc London
35:139–270
Flechtmann CHW (2011) Summary of the international congresses of acarology. Zoosymposia
6:9–13
Halliday RB (2001) 250 years of Australian acarology. In: Halliday RB, Walter DE, Norton RA,
Coloff MJ (eds) Acarology: proceedings of the 10th international congress of acarology.
CSIRO Publishing, Melbourne, pp 3–16
Hughes TE (1959) Mites or the Acari. University of London, London, 400 pp
Further Reading 19
Johnston DE (1968) An Atlas of Acari 1. The families of Parasitiformes and Opilioacariformes.

Acarology Laboratory, Ohio State University, Columbus, 110 pp
Krantz GW, Walter DE (2009) A manual of acarology, 3rd edn. Texas-Tech University Press,
Lubbock, 809 pp
Lindquist EE (1984) Current theories on the evolution of major groups of Acari and on their rela-
tionships with other groups of Arachnida with consequent implications for their classification.
In: Griffiths DA, Bowmann CF (eds) Acarology VI, vol 1. Ellis Harwood, Chichester, pp 28–62
Lindquist EE (2006) Poising for a new century. In: Halliday RB (ed) Proceedings of 10th interna-
tional congress of acarology. CSIRO Publishing, Melbourne, pp 17–34
Prasad V (1982) History of acarology. Indira Publishing House, Michigan, 472 pp
Vitzthum HG (1943) Acarina. In: Bronnes Klassen und Ordnengen des Tierreikhes, 5 Abt.4, Buch
5, Lief 1-7, 1-1011, Leipzig
Walter DE, Proctor C (2010) Mites as modern models: acarology in the 21st century. Acarologia
50:131–141
Zakhvatkin AA (1952) Subdivision of mites (Acarina) into orders and position of these in the
system of the Chelicerata. Paraz Sbornik 14:5–46
Zhang ZQ (2011) Animal diversity: an introduction to higher level classification and taxonomic
richness. Zootaxa 3148:7–12
Zhang ZQ (2014) Acarological publications in the last 150 years: historical trends and an assess-
ment of current journals. Syst Appl Acarol 11:250–252
Electronic Reference (ER)
1. https://acarology.osu.edu/history. Acarology History

2. https://scholarspace.manoa.hawaii.edu/bitstream/10125/8124/1/3Swift.pdf. History of Acarology
in Hawaii
3. http://www.biotaxa.org/saa/article/viewFile/saa.19.2.16/9061. Acarological publications in the
last 150 years: historical trends and an assessment of current journals
4. http://www1.montpellier.inra.fr/CBGP/Montpellier2008/AcarologuesFrancais.pdf. A quick over-
view of the Story of French Acarologists
5. http://www.esaeg.org/volumes/Vol1no1History.htm. History of Acarology in Egypt
6. http://www.researchgate.net/publication/242137696. 250 years of Australian Acarology
Acarine Technology
3
Abstract
Ticks and mites – members of the Subclass Acari – are a constant threat to eco-
nomically important crops, stored grains and other stored products, livestock,
wildlife and humans. Because of their small size, it is very difficult to study their
intricate structures on both the dorsal and ventral surface of their body which are
otherwise useful in identifying them. Different techniques used in collection of
plant-associated mites, water mites, soil mites, stored product mites, bee para-
sitic mites, and parasitic mites and ticks associated in different ways on both
invertebrates and vertebrates are discussed. Techniques employed in their preser-
vation in different killing mediums are given in good details. Slide-mounting
techniques for water mites, ticks and plant-associated mites for temporary as
well as for permanent preparation and formulation of different mounting medi-
ums are mentioned for their efficient studies. Preparation of specimens for scan-
ning electron microscopy studies (SEM) and molecular analysis are
given. LT–SEM technology has been mentioned for producing magnified clear
images needed for showing details of intact mites and for studying their interac-
tions on plant and insect hosts. Techniques for packing and shipping mites for
technical assistance, and techniques for rearing and sampling, and for testing
different pesticides against them are properly considered.
Keywords
Collection • Preservation • Mounting • Acaricides
The members of the Subclass Acari are a constant threat to economically important
crops, stored grains, livestock, wildlife and humans. Because of their small size, it
is very difficult to study their intricate structures on both the dorsal and ventral sur-
face of mite body which are otherwise useful in identifying them. Different tech-
niques utilized for their collection, preparation, preservation, culturing and

DOI 10.1007/978-981-10-1594-6_3
22 3 Acarine Technology
biological studies are being reported hereunder which may help in their easy
understanding.
3.1 Collection
For different taxonomic and faunistic studies, a good collection is needed. A good
collector knows from where to collect the mites, or for which type of damage symp-
toms on the host to be looked for, and when or how to collect the mites. Before
collection of the mites, it is important to know the habits and habitats of the different
mites. Different technologies used to collect the mites under different situations are
given below briefly.
3.1.1 Plant-Associated Mites
Since most of the plant-associated mites are negatively phototropic, so mites are
normally found on the undersurface of the leaves either in colonies (like members
of the families Tetranychidae, Tenuipalpidae or Eriophyidae) or in solitary forms
(like members of the family Tarsonemidae and majority of predatory mites). The
mite species which do not live in colonies are found either on the leaf lamina, along
midribs or veins or in angles formed by major veins. A good number of mites (mem-
bers of the family Phytoseiidae and Tydeidae) are found on twigs, stems or under
barks. The eriophyid or worm-like tiny mites are found on various malformed or on
deformed plant parts in buds, flowers and axils of leaves. Eriophyid mites are also
commonly found within the galls which are usually of open type contrary to the
insect-caused galls which are of closed types. The following different methods are
used for collection of different plant-associated mites.
Handpicking Infested leaves or other infested plant parts are examined under a
10× hand lens in the field, and the mites found can be picked up by a fine hairbrush
(00 series) moistened with 70 % alcohol or water. Better results are obtained if the
infested leaves or other plant parts are brought in the laboratory in polythene bags
after tightly closing the mouth of the bag with a rubber band. In the laboratory, the
mites are examined on plant parts under a microscope. During field trips, the plant
material may be stored in the thermal box with ice to reduce movements of mites,
their predation by natural enemies and desiccation. Mites can be examined or
counted under a microscope in the laboratory or picked with a fine brush for preser-
vation in the suitable preservative (usually 70–80 % alcohol) or mounted directly on
a glass slide for studies (Evans 1992; Krantz and Walter 2009; Gupta et al. 1972).
Sweeping On low herbaceous plants, grasslands, etc., sweeping can be done with
a fine net several times. A large number of mites are found on the inner surface of
the net wall which can be picked with a fine brush.
3.1 Collection 23
Beating Different plant parts are beaten with a wooden rod on a white enamel tray.
Beating will dislodge the mites on the tray which can be collected with a fine brush.
Samples of the mite-infested foliage may also be beaten on a 1–2 cm mesh screen
covering a large tray, thus avoiding leaves and other residues to fall in the tray.
Dislodged mites can be picked with a fine brush and put in a preservative.
Aspirators An aspirator is made of a large vial capped by a rubber bung with two
tubes, one tube leading to a sucking source (a rubber bulb or the mouth of collector)
and the other for pointing to the mites. A gauge may be fixed to the end of the suck-
ing tube inside the vial to prevent the mites being sucked back outsurfaces. Aspirators
are quite useful for collection of plant mites directly from its habitat to the preserva-
tive (70 % alcohol) eliminating the necessity of handling the specimens (ER7).
Aspirators are used for quick-moving mites of relatively large size on leaf and
ground surfaces (Singer 1964).
Washing in Hot Water Mites can also be collected by washing infested plant parts
in hot water. When infested plant parts are shaken in water in a container, mites fall
from the plants. Mites in the water may be separated by pouring the water through
a series of sieves of different mesh sizes to separate mites of different sizes.
Rust and Bud Mites These worm-like mites can be collected by pouring a thin
syrup of sorbitol (made up of 25 % solution of isopropyl alcohol in which a few
crystals of iodine have been added) over the leaf or other plant part in a small con-
tainer, and such leaves can be examined later under a stereoscopic microscope in the
laboratory. The mites are individually transferred by a needle to the slides or vials.
Mite-galled plant parts are allowed to dry out completely in paper envelopes. Mites
can even be recovered many years later, if necessary, by warming the dried galls in
a small amount of sorbitol syrup in a cavity slide (Krantz and Walter 2009).
Vacuum Technique A hand-operated vacuum apparatus can be used for collecting

a large number of mites from the surface of ground vegetation.
3.1.2 Water Mites
In intertidal and slow subtidal zones, collection of mites by the hand at low tide is
recommended. Colonies of mussels, polychaetes, barnacles, bryozoans and hydro-
zoans are scrapped off. Algal tufts, holdfasts and fronds, shells or pebbles covered
with unicellular algae and fine and course deposits from a beach surface as well as
from deeper layers are gathered. From 50 to 500 cm3 of the substratum may yield
hundreds of water mites. Sublittoral collection can be made with trawl, dredge and
grab. Halacarid mites inhabiting the flocculent ooze are collected with epibenthic
sleds.
Aquatic Net Aquatic net consists of a strong net with a wide opening (25–40 cm
diameter) and fine mesh size (250 μm). The net is methodically and vigorously
treaded down the plants while dragging the net along to scoop up the debris, silt and
organisms that become temporarily dislodged as the water is forced up through the
disturbed layer of vegetation. When a mixture of fine salt, plant fragments and
organisms begin to fill the bottom of the net, it is transferred to a plastic bag contain-
ing a small amount of water (Krantz and Walter 2009). This process is repeated till
the bag is approximately half-full of the material. The bag is then nearly filled with
clean water and the contents stirred thoroughly to allow heavy inorganic material
such as gravel and sand to settle to the bottom. The contents of the bag are then
examined under a hand lens or dissecting microscope. Alternatively, the contents of
the net may then be shaken onto a black (or white) tray and mites can be sorted and
picked using a hairbrush (ER6). If there is a lot of debris in the net, the contents may
be washed into the hot water and separated using different methods.
3.1.3 Soil Mites
Modified Tullgren Apparatus Berlese in 1905 used a funnel technique for collec-
tion of mites inhabiting the soil, leaf litter and other organic detritus. The system
consisted of a metal funnel equipped with a hot water jacket; the soil and organic
material were supported on a sieve with a wider opening. The material was slowly
dried out by heat from the water jacket and the mites and other soil fauna dropped
through the sieve into a collecting tube below. In the collection tube, 70–80 % alco-
hol is kept; however, 1–3 % glycerine is put so as to keep the specimens moist even
if alcohol dries. Later Tullgren in 1918 replaced the water jacket with an electric
bulb suspended over the funnel as a source of heat (Colloff 2009; Edwards and
Fletcher 1971; Norton and Kethley 1988; Steykstal and Murphy 1986). For quanti-
tative work, several Tullgren funnels are mounted together in batteries, and in this
way a large number of samples are handled in a shorter time. Simple Berlese–
Tullgren funnels are usually adequate, but for expedition purposes collapsible fun-
nels made of plastic sheeting can also be used (ER8).
Simple Floatation Technique The collected soil sample is stirred in a magnesium

sulphate solution of high specific gravity and a stream of air bubbles is put into the
suspension from below. After settling, the liberated organisms and organic matter
float to the surface and are collected on a filter paper or through a sieve. Floatation
technique is better for collection of mites, their eggs and even inactive stages, but it
is time consuming and unsuitable for the material containing higher proportion of
organic matter. Moreover, washing and sieving process also tend to damage the deli-
cate species of mites (Gupta et al. 1972).
3.1 Collection 25
Grease Film Technique Extraction of soil mites on a grease film is done from a
mixture of the soil sample and water. Since most soil dwelling mites possess a water
repellent cuticle, they tend to stick to a suitable grease in a soil water mixture.
3.1.4 Parasitic Mites
3.1.4.1 Bee-Associated Mites (Barlow 2009; Bush 2009)

Ether (or Alcohol) Roll Brush or shake worker bees sampled from near the middle
of the hive into the wide-mouthed mason jar, and place the lid on the jar of the cap-
tured bees. Spray a short burst (about one second) of ether or alcohol into the closed
container. Gently roll the jar from side to side to coat all of the bees with the ether
or alcohol.
Powdered-Sugar ‘Shake’ Brush the worker bees into the wide-mouthed mason
jar with a two-piece lid. Remove the central portion of the lid and replace it with
approximately 9 mesh screen (3 mm × 3 mm mesh). Replace the modified lid and
add a tablespoon of powdered sugar through the mesh screen. Roll the jar from side
to side to distribute the sugar over all of the bees. Pour the sugar and dislodged mites
through the screen onto the cheese cloth. The powdered sugar makes it difficult for
the mites to adhere to their host, causing them to fall off the bees.
Sticky Board The screened sticky board is placed between the hive floor and brood
frames. The screen separates the mites from the bees and prevents the bees from
reaching the sticky paper and becoming entrapped. As mites are dislodged during
the grooming process, they will fall through the screen cover and adhere to the
sticky white paper on the bottom board. The sticky board should be placed in the
hive for 24 h and removed as necessary to examine the mites.
Drone/Brood Sampling The varroa mites refer to feed on drone brood. So for tak-
ing out drone pupae, a capping scratcher is used to remove the caps and pupae from
the brood comb. The cappings and pupae are lifted from the comb, and the pupae
are examined for mites.
Sticky Board with Acaricide By using the sticky board (as given above) with an
acaricide provides as a good method for monitoring mite population levels within
the hive. The use of sticky board with acaricides like Apistan® (formulated as
impregnated plastic strips) is recommended above all other techniques as it is effec-
tive even when varroa mites are low or clumped within the hive.
Dissection Technique Since the endoparasitic mite Acarapis woodi infests the tra-
chea of bees, dissection of bees to expose the trachea is a must. Sometimes, even
dyes are used to differentiate the position of parasitic mites in the trachea (ER1).
3.1.4.2 Mites Associated with Other Insects

Many members of the morphologically unusual heterostigmatine family
Podapolipidae are ectoparasitic beneath the wings/elytra or in intersegmental mem-
branous areas of orthopterans, hymenopterans and cockroaches. But in locusts,
Locustacarus spp. are endoparasitic in tracheae and air sacs of locusts.
Otopheidomenid species, Katydiseius nadchatrami and Orthopteroseius sinicus, are
endoparasitic in the tracheal system of their katydid and grasshopper hosts. Moth
mites, Otopheidomenis spp., are found in the wing bases and the thoraco-abdominal
cleft of their sphingid and noctuid moth hosts. Moth ear mites, Dicrocheles spp.,
invade the tympanal cavities where they feed and lay eggs.
3.1.4.3 Mites from Other Invertebrates

Ereynetid mites, Riccardoella limacum, are found on the surface and in the mantle
cavity of slugs and snails and can easily be found under a microscope. Some mites,
Ewingia spp. (family Acaridae), attach to the grills or other body regions of coeno-
bitid hermit crabs.
3.1.4.4 Mites from Vertebrates

Endoparasites Some endoparasitic mites are associated with the respiratory sys-
tems of their hosts. Members of family Rhinonyssidae are largely confined to the
nasal passages of birds, lizards and snakes but some species even invade the lungs
and air sacs. Representatives of astigmatine family Pneumocoptidae are commonly
found in lungs of rodents. Dermanyssid mites and mites belonging to family
Entonyssidae are exclusively parasitic in the lungs and trachea of reptiles. Nasal
passages of seals and walruses, apes, squirrels and dogs have been commonly found
infested by various endoparasitic mites. Burrowing mites, Sarcoptes scabiei, are
found beneath the skin of their hosts. Burrowing mite, Knemidocoptes mutans, is
found on the epidermal scales of legs of chickens.
Ectoparasites Ectoparasitic mites are collected from their habitats (nests, runaway
litter, mammals, birds, etc.). The nest and litter material can be easily found through
Berlese–Tullgren funnel separation technique. From rodents, mites can also be col-
lected through combing and brushing of the body on a white paper or enamel tray.
Parasitic mites can also be collected by dragging a flannel cloth over suspected
infestation sites (flagging). Collection of mites from living hosts can be done even
by anaesthetizing the host with an ultramuscular injection of anaesthetizers. Even
through tracheobronchial swab irrigation techniques, living mites of monkeys can
be found. Sometimes the mites from dead rodents and birds are removed simply by
immersing the infested animal in the water detergent mixture which is shaken vigor-
ously. Chiggers can be found on smaller animals simply by examination of ears,
3.1 Collection 27
areas on the muzzle or chin, axillary and inguinal areas and areas on the midline of
the belly. Many non-burrowing, burrowing and hair follicle mites can be found sim-
ply through examination of skin scrapings under a microscope (Steykstal and
Murphy 1986).
3.1.4.5 Collection from Stored Products

From the stored grains or from other storages, sieving and floatation methods can be
used for the detection of mites. In stored grains PC™ trap and BT traps are quite
effective in determining the presence of mites, and these traps may provide an ear-
lier indication of their presence than traditional methods. BT traps have been
designed specifically for detection of storage mites in various storage and produc-
tion premises. The addition of the lure (oilseed rape) significantly improves the trap
catch of Acarus siro, Tyrophagus longior and Lepidoglyphus destructor in both the
BT traps and PC™ traps (Thind 2005).
3.1.5 Collection of Ticks
3.1.5.1 Direct Examination

Ticks being larger in size than mites can be detected on animals by direct examina-
tion even without the use of a hand lens or microscope. Engorged ticks (those filled
with blood) are particularly easy to see as they look somewhat like a castor bean in
shape and colour.
3.1.5.2 Combing Technique

Combing hairs or feathers exposes the skin which can then be directly examined.
Ticks are generally found attached on areas around the head, in the ears, on the
shoulders and other parts of the body. On cattle, ticks can be commonly found in
highly vascularized areas of the skin (i.e. tail, udder, etc.). Unattached ticks can be
recovered from the host by simply combing. The ticks can also be collected directly
from the hosts by using tweezers or fingers or through Tick Key. Grasp the tick as
close to the skin as possible and pull firmly away until it detaches (Krantz and
Walter 2009).
3.1.5.3 Dipping Technique

Heavily infested animals can be dipped, dusted or shampooed with an acaricide and
then combed thoroughly to rid the ticks from the body of the animal.
3.1.5.4 Dragging Technique

By dragging a white cloth (2.5 m × 2.5 m) stapled to a dowel to which a cord is
attached with a second dowel or board at the end to weigh the cloth down. Questing
ticks grab onto the cloth as it is dragged over the grass and brush. The drag is then
inspected for ticks at regular intervals. Tick drags are effective when the vegetation
is dry and the temperature exceeds 5 °C (Evans 1992).
3.1.5.5 Carbon Dioxide (CO2) Trapping

For many species of ticks, carbon dioxide is found very useful as the ticks are known
to sense the carbon dioxide and move towards its source. Dry ice is placed in venti-
lated containers, or compressed gas is used as a source of carbon dioxide (Krantz
and Walter 2009).
3.1.5.6 Host Trapping

This seems to be an effective method for detecting ticks. In areas with dense tick
populations, simply walking through the habitat of animals is an effective surveil-
lance method.
3.2 Preservation/Killing in a Medium
Terrestrial mites can be killed/fixed and satisfactorily preserved in 70–80 % ethyl

alcohol after adding 5 % of glycerol to prevent the specimens from drying out com-
pletely because of evaporation of the alcohol during long periods of storage.
Isopropyl alcohol can also be used for preservation of mites. Sometimes Oudemans’
fluid (ethyl alcohol, 87 parts; glycerol, 5 parts; and glacial acetic acid, 8 parts) is
employed for long-term preservation of the specimens. In this medium, the mites
are killed and their appendages are extended, and their orientation during mounting
can more easily be done. However, progressive hardening of tissues occurs in alco-
hol – preserved material which may render specimens unusable for phase contrast
or interference purposes. Koenike’s fluid can be used in general for a permanent or
semi-permanent storage of water mites as the tissues and appendages remain in a
flexible state, thus alleviating the problems of breakage during mounting/dissection.
The composition of Koenike’s fluid is glacial acetic acid (10 parts), glycerine (50
parts) and distilled water (40 parts). Some workers use lactic acid (65 %) for preser-
vation of soil mites (Steykstal and Murphy 1986).
Mite specimens should be stored in small vials. Proper labels containing infor-
mation about the host, locality, date of collection and place should be inserted in the
vial. But formalin should never be used for mite preservation since it hardens the
specimens and renders their clearing very difficult. Good seals on vials are a neces-
sity especially in warmer areas, and refrigeration can be used to reduce fluid loss.
Sometimes isopropyl alcohol and methylated spirits are also used for killing and
preservation of mites for shorter periods.
3.3 Preparation (Slide Mounting)
Preparation of mites before mounting on the glass involves the removal of their
internal organs in order that other structures of taxonomic value can be readily
examined. During the preparation procedure, proper clearing of the mites should be
done, but it should be kept in view that clearing procedures should not cause any
distortion of the specimens. The main objective of clearing is to macerate and
3.3 Preparation (Slide Mounting) 29
dissolve internal tissues especially muscles. To soften the cuticle for dissection and
maceration purposes, lactic acid (60–95 %) is commonly used. But lactic acid
should not be used for the oldest alcohol-preserved specimens. For such purposes,
Nesbitt’s solution or 5–10 % potassium hydroxide (KOH) should be used. Nesbitt’s
solution is composed of chloral hydrate (40 g), concentrated hydrochloric acid
(2.5 ml) and distilled water (25 ml). The time required for keeping in clearing agents
depends on the temperature used, length of the time specimen is preserved, size and
sclerotization of the specimen (Singer 1967).
3.3.1 Preparation for Water Mites
Puncture dorsally the mite body after taking out from the Koenike’s fluid and then
place the specimen in about 8 % solution of potassium hydroxide (KOH) for 2 hours.
Thereafter remove the specimens from KOH and place it in a mixture of glycerine
jelly and water for proper dissection purposes. Then mount the specimen in glycer-
ine jelly.
3.3.2 Plant-Associated Mites
The following techniques for temporary and permanent mounting plant mites are
used commonly.
3.3.2.1 Temporary Preparation

The specimens are placed in a small quantity of 60 % lactic acid on the glass slide
and orientated and then apply the coverslip. The preparation is then gently heated
until clear. The duration of the clearing process will depend on the degree of sclero-
tization of the specimen. Specimens engorged in blood should be pricked with a fine
needle to facilitate their clearing. Living or preserved specimens can be placed in
cold or warm lactic acid (50–100 %) depending upon the degree of sclerotization of
specimen. Weakly sclerotized mites of family Tetranychidae and astigmatine mites
should be treated with 50–60 % lactic acid, while heavily sclerotized mites such as
some Mesostigmata may be treated in lactic acid (80–100 %). Lactic acid is quite
suitable for almost all terrestrial mites except the Eriophyoidea. If body contents are
not being cleared with lactic acid or lactophenol, then they may be treated with
2–5 % KOH, xylene or hydrogen peroxide. But KOH treatment should be followed
by washing in acid alcohol and the use of xylene will require different grades of
xylene concentrations. Temporary mounts sealed with glyceel or euparal may be
transformed into semi-permanent preparations and kept for 2–3 years (Evans and
Browning 1955).
3.3.2.2 Permanent Preparation

For satisfactory mounting of plant-feeding mites particularly tetranychids, Hoyer’s
mounting media is commonly used. Hoyer’s medium consists of distilled water
25 g, gum arabic 15 g, chloral hydrate 100 g and glycerol 10 g.
Living mites or specimens of mites preserved in alcohol can be placed directly
into the Hoyer’s medium. Gum chloral mites are not completely permanent. If nec-
essary, the mounted mites can be removed for remounting by soaking the slides in
warm water. Polyvinyl alcohol has also been used as a permanent mountant but
shrinkage of specimen during drying has been reported. Faure’s medium, contain-
ing gum arabic (15 g), distilled water (25 ml), glycerine (10 ml) and chloral hydrate
(25 g), has also been found to be a satisfactory general mountant for mites. The
above medium is prepared by dissolving gum arabic in water first, and then glycer-
ine and chloral hydrate are added in the order as given above. The solution should
be made without heating. The specimen is transferred directly from the preservative
into the mounting medium in which it eventually gets cleared. However, heavily
sclerotized species are first cleared in 60 % lactic acid before mounting. A liberal
quantity of the medium should be applied to the slide to allow for shrinkage occur-
ring during the hardening process. Since these mediums are hygroscopic, when the
slide gets dried, the edge of the coverslip should be sealed with euparal or nail pol-
ish or paint which is waterproof in nature. A well-made ringed slide in Hoyer’s
medium may last for decades, if stored under proper conditions. Slides should be
kept horizontally in storage boxes (Evans and Browning 1955).
In case of water mites, specimens can be mounted directly from the water or
alcohol into the Hoyer’s medium or polyvinyl alcohol. But specimens cleared in
potassium hydroxide or lactic acid must first be rinsed in the water. Sometimes dis-
section of the material is needed to expose the structures necessary for
identification.
3.3.3 Preparation of Ticks for Study
Larval ticks may be cleared and mounted on slides as given under mites. Nymphal
and adult ticks are usually examined out of alcohol with the stereoscopic micro-
scope. But it is essential to remove all superficial liquid from the body so that orna-
mentation on body can be properly observed. So the ticks should be allowed to
become too dry. Moreover, there is a necessity of observing different body structures
from different directions. So a tick for examination can be picked up with fine for-
ceps and dabbed on wad of dry paper tissue to remove all surface liquid. Ticks should
not be examined under strong and hot illumination for too long. Regarding observa-
tion on ticks, cold lights supplied from fibre optic illuminators are more useful.
3.3 Preparation (Slide Mounting) 31
3.3.4 Mounting Ticks on Glass Slides
Several slide-mounting techniques for ticks have been used with varying degrees of
success. The classic Canada Balsam technique is quite efficient but time consuming.
Water-soluble media are inadequate in the humid/tropic conditions, even when the
slides are ringed with a waterproofing material such as glyptal. Crystallization of
the media, probably due to absorption of water, is the main problem with slide-
mounting techniques. Larvae and nymphs of ticks starved for 15 days (for larvae)
and 24 h (for nymphs) either by killing in hot water or 70 % ethanol (approximately
70 °C) and then preserved in 70 % ethanol. Fixed specimens are subsequently trans-
ported to lactic acid for one hour at 50 ° C and then the specimens are kept for about
24 h in alcoholic solution saturated with phenol. The specimens after clarification
are washed in 100 % ethanol. Washed specimens are transferred to a solution of pure
creosote for approximately 24 h and then mounted in Canada Balsam (Evans 1992).
3.3.5 Staining
In whole mount prepared by lactic acid or lactophenol, structures of taxonomic

importance are usually discerned and staining of the material is rarely needed.
However, certain structures in weakly sclerotized mites may be difficult to differen-
tiate without the use of phase-contrast microscope. In such cases, staining can be
advantageous. Small amount of ‘lignin pink’ can be added in the clearing medium.
Living specimens and specimens preserved in alcohol could be transferred directly
to this clearing staining medium. Fuchsin stain, chlorazol black and toluidine stain
can also be added in the same way. In case of eriophyid mites, staining with iodine
is also quite useful.
3.3.6 Slide Preparation Technique
First place a drop of the medium in the centre of the glass slide. Drop size will
depend on the size of coverslip used, thickness of mite specimen and viscosity of
the medium. Mites can be mounted directly from the water, alcohol or plants to the
Hoyer’s medium or polyvinyl alcohol. But specimens cleared in potassium hydrox-
ide or lactic acid must first be rinsed in water.
Sometimes dissection is required to expose structures necessary for identifica-
tion. Fine watchmaker’s forceps or insect minuten pins mounted on wooden match
sticks are the most useful tools for dissection purposes. Large mites usually require
dissection, and in some groups mites are always dissected before mounting. Large
mites can also be mounted in depression slides or surrounded by protecting plastic
rings. If oil immersion lens is required for observation, then the slides should be of
lesser thickness. Coverslips used should be circular (10–18 mm in diameter) except
for very large mites and thin (0001–1). The small (10–12 mm), thin (000–00) cov-
erslips can be very expensive and provide comparatively less protection for the
specimen. Smaller area need be searched to find the mite specimen, and the oil
immersion lens is easily used (Krantz and Walter 1909).
If the purpose of the slide is to provide taxonomic information, then the rule is
one mite per slide. However, for routine purposes many specimens can be mounted
on a slide to save time and money. A small stainless steel pin with a flattened tip and
mounted in a wooden handle serves as a good tool for transferring mites. Always
place the mites in the centre of the drop of the medium and gently push it to the
surface of the slide. Allow any air bubbles trapped in the medium to rise to the sur-
face and burst. Gently lower the coverslip from one end of the drop to the other and
avoid trapping any air bubbles and then adjust its orientation by gently putting pres-
sure on the coverslip. In case of spider mites, females are orientated dorsoventrally
or in profile; males preferably are mounted in profile to see the aedeagus. Pick up
the mounted slide, invert it and circle the position of the specimen with a fine-tipped
indelible marker. Then revert the slide to its upright position and label it with an
indelible marker. Heat the slides in an oven or slide warmer at 45–50 °C until the
medium around the rim is dry to the touch. When the slide is properly dried, replace
the temporary labelling with a permanent label with all the necessary data regarding
the date collected, host and collector on one side of the coverslip. On the other side,
taxonomic information about the specimen is given. The coverslip of dried slide
should be ringed with a ringing material like glyptal, insulating varnish or nail pol-
ish through a small brush or a bottle applicator. Store the properly labelled slides
horizontally with the coverslip side facing up in slide boxes or flat trays (Travis
1968).
3.4 reparation for Scanning Electron Microscopy Studies

P
(SEM)
Initial killing of mites should follow the guidelines reviewed above. Additionally,
the mites can be boiled alive in a test tube of lightly soaped water and boiling chips
to extend limbs; clean them of some of debris, and fix tissues. For better sclerotized
mites, less treatment is required before SEM. For example, most oribatid mites like
brachypyline and ptychoid mites can be dehydrated in 100 % ethanol, transferred to
a small watch glass of acetone and dried under a fume hood in a few hours with little
or no resulting deformation. Specimens are then transferred to a SEM stub covered
with double-sided sticky tape using a fine brush and sputter coated with gold, gold–
palladium, platinum or carbon. The best approach for soft-bodied mites and espe-
cially for mites feeding on plant tissues appears to be low temperature electron
microscopy (ER4). This technique has been used to produce very good images of
Pyemotidae, Eriophyoidea, Tydeidae and Acaridae and many other soft-bodied
mites (Saito and Osakbe 1992).
3.7 Packing and Shipping Mites 33
3.5 Preparation for Molecular Analysis Studies
DNA can often be recovered from specimens stored in 70 % ethanol, at least over
the short term, but killing and storage of mites in 100 % ethanol produces better
results. Even more better results are obtained if flash freezing live mites in small
plastic tubes or in a buffer solution at 70 ° C or lower. Standard Chelex–DNA tech-
nique works well with mites of moderate size using 1–5 specimens ground up with
a micro-mortar and pestle. A variety of kits are commercially available that digest
DNA from specimens and allow retrieval and sliding of the exoskeletons for use as
voucher specimens (Ochoa 2000; Hillis et al. 1996).
3.6 Photomicrography of Mites
For detailed information about mite’s correct identity, scientists at the Systematic
Entomology Laboratory, USDA, have developed low-temperature scanning elec-
tron microscopy (LT-SEM) technology which is also known as cryo-SEM. Through
this technique, highly magnified clear images are produced that show details of
intact mites and their interactions on plant and insect hosts. Cryogenic SEM is an
effective tool for viewing mites because the high resolution makes it suitable for
viewing small biological samples such as mites. Scanning electron microscopy
allows for clear images of small details that cannot be viewed using traditional
microscopy techniques such as light microscopes. Cryogenic SEM has the ability to
dehydrate and preserve a samples’ original shape (Sircom 2002). Because the cuti-
cle layer enclosing a mite is delicate, it is susceptible to deformation and complete
collapse. By cryogenically preparing the mite, dehydrating the mites while main-
taining their original structure is made possible (ER4).
To prepare or fix a mite specimen for viewing under LT-SEM, liquid nitrogen (at
−195 °C) is used; this cryofixation instantly freezes the mite in its natural state on
the host and prevents it from moving or becoming distorted. With this technique
mites can be magnified for more than 50,000 times; hence, many structural details
of mites can be found at different angles and directions. Through LT-SEM, ultra-
structural features like pores on the body surface can be found which proves very
useful for correct identity of mites. Such details provide vital information regarding
how mites move and hold onto their hosts (Achor et al. 2001; Ochoa 2000).
3.7 Packing and Shipping Mites
3.7.1 Specimens in Vials
Fill each vial with liquid preservative and stopper it tightly by putting a piece of
wire in the vial and the stopper. Cork stoppers should not be used as there are
chances of leakage of the material through them. Screw-topped vial should be firmly
closed and sealed with an adhesive tape or Parafilm around the lower edge of the cap
and part of the vial. Paraffin wax sealing should not be done as it usually breaks and
will not prevent leakage. Wrap each vial with cotton, tissue, paper towelling or simi-
lar other material, and do not allow any glass part of vials to touch each other to
avoid its breakage during trans-shipment. Several vials can be wrapped together or
held tightly with tape or rubber bands as a unit. They may be placed in small card-
board box with enough packing to ensure that they are not shaken around. While
shipping more than one box or packet of vials, tie or wrap them together as a unit
before they are placed in the larger carton (Evans and Browning 1955).
3.7.2 Shipping Microscope Slides
First of all, ensure that slides being shipped are thoroughly dried and cured. Slides
may be shipped in specially designed slide holders which are available commonly
from the biological supplies companies. The slides may also be shipped in standard
storage boxes keeping enough soft tissue around each slide and between the slides
and box lid so that the cover glass does not come into contact with anything. If no
slide holders are available, then a few slides, each wrapped with tissue, may be tied
together at each end with tape and rubber bands, wrapped in strong paper and placed
in a carton with packing material. The carton should be made of strong corrugated
board or other stiff material. The packing material can be in the form of shaved
wood, crumpled newspapers or starch peanuts. But clear plastic sheet material with
a regular pattern of bubbles (bubble wrap or blister pack) can also be used for pack-
ing since it is very light in weight and has excellent shock-deadening properties.
3.7.3 Shipping Specimens
In no case live specimens are to be submitted to experts for identification as the law
of many countries prohibits the importation and movement of the live pests, patho-
gens, vectors and infested articles that might harbour these mites. Living specimens
can only be shipped after complying all conditions required as per quarantine regu-
lations of different countries. It is recommended that all packages be marked
‘FRAGILE’ and a complete return address be written on the outside of each con-
tainer. Valuable specimens like ‘Type Species’ should be sent via registered mail as
it allows misdirected packages to be tracked much more readily. Place a label such
as Dead Insects for Scientific Study and No Commercial Value on the outside of the
package (Steykstal and Murphy 1986; Bergh and French 2000).
3.8 Working with Live Mites
Mites have a high surface/volume ratio, desiccate easily and must be maintained at
a relative humidity similar to the one they experience under normal living. For soil
and leaf litter mites, atmospheric humidity more than 95 % is required. On the
3.10 Sampling Methods 35
contrary, mites adapted to drier habitats often need quite lower humidities. Wet
strips of paper towel supplied through the funnel can be placed in vials containing
live mites. However, for good results, 20–40 % bottom surface of collection vial is
filled with a mixture of powdered activated charcoal and plaster of Paris. Different
types of microscopes are employed in rearing of plant mites. Mites are also cultured
primarily in various types of glass or plexiglass cells or dishes.
3.9 Rearing Techniques
A number of methods for rearing of free-living mites have been developed. Mites
are cultured in various types of glass or plexiglass cells. Leaf disc method is com-
monly used for rearing mites under laboratory conditions. A proper sized leaf is
placed on wet cotton and mites are released for rearing or for developmental studies
at controlled conditions. Cotton swabs need to be moistened periodically to avoid
drying of leaf discs. Sometimes moistened filter paper is also kept instead of cotton.
Rearing of plant mites in mass culture requires healthy vigorous host plants main-
tained under optimal growing conditions. Tashiro leaf cage technique is also used
for laboratory studies. But all these and many other techniques used for rearing (or
development) studies require keeping the leaf discs turgid through periodic provi-
sion of water on the material on which leaf discs are kept. Many other techniques
are also available in literature regarding rearing of plant-associated and animal para-
sitic mites and can be consulted for particular studies (Abou-Setta and Childers
1987; Argov et al. 2002; Helle and Overmeer 1985; Krantz and Walter 2009; Tashiro
1967).
3.10 Sampling Methods
Several techniques have been devised by different workers for sampling of field
populations of spider mites. Each of these techniques has its own advantages and
disadvantages. The following important techniques are mentioned briefly (Gupta
et al. 1972; Chant and Muir 1955; Leigh et al. 1984).
3.10.1 Direct Counting
Leaf samples are collected in the field and then inspected under a microscope to
note mite counts per unit size. This is a time-consuming procedure and is not very
efficient to count the walking mites as they may be counted more than once. But
cold storage of leaves prior to counting may reduce counting errors. In the field, if
the population per leaf is low, then direct counting by a hand lens is quite useful.
3.10.2 Imprint Counting
Infested leaves are pressed between a fold of sheet of mimeograph or any other
paper of suitable absorbance. On pressing with roller, the mites are crushed and
leave an imprint which with experience can be distinguished from imprints of dif-
ferent mite species. This is a semi-permanent record of the mite infestation, and
counting is much easier. Even imprints of different stages of coloured mites can be
differentiated in some mites.
3.10.3 Mite-Brushing Machine
Infested leaves are passed between contra-rotating brushes which dislodge the mites
onto a revolving counting disc (grid). The disc is smeared with sticky coating of
Vaseline or other similar coating material. Even the mites on discs can be easily
photographed, and the observations can also be recorded later after keeping the
discs in cold storage conditions (ER2). This machine distributes the mites uniformly
over an adhesive-coated disc kept on revolving plate. Effectiveness of brushing
machine depends on the smoothness of leaves. Less proportions of mites are brushed
off when the leaves have pronounced ribs and a dense field of hairs. Another disad-
vantage is that tender or large leaves are crumpled or shredded as they pass between
the opposing roller brushes resulting in a meshy sample (ER3 and ER5).
3.10.4 Beating Foliage
Simple beating of infested foliage with pencil or glass rod over a white sheet of
paper which may be divided into different regions can be used. Mites that fall on the
paper can be counted through a hand lens. However, clover mites directly drop from
a plant when it is disturbed.
3.10.5 Washing Technique
The mites are washed from the infested leaves by stirring in a solution of lower
concentration of alcohol. Even mite rinse machine has also been used by some
workers to separate mites from the silk and soil particles on the leaves and also from
the leaf folds. The rinse machine circulates the infested leaves in a hypochlorite
solution which dissolves the silk and separates the mites from leaves and silk.
3.11 Acaricide Testing Techniques
Bioassays of different pesticides against mites are conducted under laboratory con-
ditions through the following methods (Ebelling 1960).
3.11 Acaricide Testing Techniques 37
3.11.1 Slide-Dip Method
In this technique, first of all a piece of double surface sticky tape is pasted on the
glass slides tightly, avoiding any air spaces that would allow a toxicant to accumu-
late between the slide and tape. With the help of stereomicroscope, 50 adult female
mites are affixed to the tape on the dorsal part of their hysterosoma through a moist-
ened tip of fine brush. The prepared slides are dipped in different concentrations of
the toxicant solution and gently agitated for 5 s. Then the slides are dried for 15 min
and placed horizontally in trays under constant temperature and relative humidity
conditions. Excess toxicant is blotted off the slide with filter paper. After 24 h of
treatment, survivors are counted. Mites that respond movement to prodding with a
fine brush are considered alive.
3.11.2 Leaf-Dip Method
This method involves taking of young leaf discs (2.5 cm × 2.5 cm) of bean/other

hosts, and onto them mite-infested leaves are pinned, and the mites were allowed to
transfer on the leaf discs. The pinned leaves are removed after 24 h or shorter peri-
ods, and the selected leaves or leaf discs are dipped in an acaricide solution for 5 s.
Treated leaves (leaf discs) are observed at different intervals for observing toxicity
of different concentrations of different pesticides. A minimum of three replications
for each concentration are kept. Intervals of observations are decided keeping in
view convenience and earlier toxicity information of the candidate pesticide.
3.11.3 Residual Leaf Disc Assays
Selected leaf discs are sprayed with toxicant solutions through Potter’s tower or
other suitable spray technique. Then mites from the culture are released on the
treated leaf discs, and observations regarding toxicity of pesticides are recorded
after suitable intervals. These tests should be conducted under suitable temperature
and humidity conditions.
3.11.4 Ovolarvicide Tests
First of all eggs are obtained on leaf discs after releasing female mites from the mite
culture and kept for 24 h. Eggs are observed after 24 h of releasing mites, and
females are removed. These leaf discs are sprayed/dipped in different concentra-
tions of pesticides, and per cent mortality of eggs and larvae is observed.
3.12 Conclusions
Mites are ubiquitous and have successfully colonized almost every known terres-
trial, freshwater and marine habitats and have parasitic relations on animals and
mankind in a variety of habitats. They are microscopic (0.08–20 mm) and studies on
their different aspects can only be done if some specialized techniques are adopted.
A brief account of collection techniques like sweeping, beating, aspirators and
washing in water is discussed here. Techniques for studies on different aspects of
phytophagous, soil and water living, animal parasitic and storage mites are dis-
cussed briefly. Different methods used for collection, killing and preservation of
mites and ticks in different habitats are mentioned. Techniques used for preparation/
mounting on slides for temporary mounts or for permanent slide mounts of mites
and ticks are also mentioned. Techniques for low-temperature electron microscopy
studies (LT-SEM) and for different molecular studies are also mentioned. LT-SEM
technique serves to produce high-quality magnified images for knowing micro-
details of mites and ticks on plants and insect hosts which is not possible through
traditional microscopy. Methods for packing and sending mites and ticks to experts
for confirmation are also given. Rearing techniques, different sampling methods and
techniques for testing efficacy of different pesticides against them are also discussed
for convenience of persons interested in their working.
Further Reading
Abou-Setta MM, Childers CC (1987) A modified leaf arena technique for rearing phytoseiid or
tetranychid mites for biological studies. Fla Entomol 70:245–248
Achor DS, Ochoa R, Erbe EF, Aguilar JH, Wergin WP, Childers CC (2001) Relative advantages of
low temperature versus ambient temperature scanning electron microscopy in the study of mite
morphology. Int J Acarol 27:3–12
Argov Y, Amitai S, Beathie GAC, Gerson U (2002) Rearing, release and establishment of imported
predatory mites to control citrus mite in Israel. Biocontrol 47:399–409
Barlow VM (2009) Sampling methods for Varroa mites on the domesticated honeybee. Virginia
Cooperative Extension, Publication 444–103. Virginia Tech, Virginia State University
Bergh JC, French JV (2000) A method for harvesting and shipping live citrus rust mites (Acari:
Eriophyidae). Fla Entomol 83:370–374
Bush SE (2009) Field guide to collecting parasites. Natural History Museum, University of Kansas,
Lawrence, 16 pp
Chant DA, Muir RC (1955) A comparasion of the imprint and brushing machine for estimating the
numbers of fruit tree red soider mite. Metatetranychus ulmi (Koch) on apple leaves. Rep E
Malling Res Stat for 1954: 141–145
Colloff MJ (2009) Dust mites. CSIRO Publishing/Springer, Dordrecht
Ebelling W (1960) Testing acaricides. In: Methods of testing chemicals on insects, vol II. Burgess,
Minneapolis, pp 156–192
Edwards CA, Fletcher CE (1971) A comparison of extracting methods for terrestrial arthropods.
In: Methods of study in quantitative soil ecology. Population, production and energy flow, IBP
handbook No. 18 V. Blackwell, Oxford, pp 150–185
Evans GO (1992) Principles of acarology. CAB International, Wallingford, 563 pp
Evans GO, Browning E (1955) Techniques for the preparation of mites for study. Ann Mag Nat
Hist 8:631–635
Further Reading 39
Gupta SK, Dhooria MS, Sidhu AS (1972) Comparative evaluation of methods for sampling of
mite, Tetranychus cucurbitae Rahman and Sapra. Entomol News (USA) 86:33–36
Gupta SK, Dhooria MS, Sidhu AS (1973) Development of sampling techniques for estimating
population of Tetranychus neocaledonicus Andre infesting brinjal. Acarologia 17:489–492
Helle W, Overmeer WPJ (1985) Rearing and handling. In: Spider mites: their biology, natural
enemies and control, vol 1A. Elsevier, Amsterdam, pp 161–170
Hillis DM, Mable BK, Larson A, Davis SK, Zimmer FA (1996) Nucleic acids IV. Sequencing and
cloning. In: Hillis DM, Monts C, Mable BK (eds) Molecular systematics. Sinauer, Sunder land,
pp 321–384
Krantz GW, Walter DE (eds) (2009) A manual of acarology, 3rd edn. Texas Tech University, Texas,
USA
Leigh TF, Magi VL, Wilson LT (1984) Development and use of a machine for recovery of arthro-
pods from plant leaves. J Econ Entomol 77:271–276
Norton RA, Kethley JB (1988) A collapsible, full-sized Berlese-Funnel System. Entomol News
99:41–47
Ochoa R (2000) Mites get frozen, photographed and identified. Agri Res 9:175–193
Saito Y, Osakbe M (1992) A new fixation method for preparing mite specimens for optical and
SEM microscopic observations. Appl Entomol Zool 27:427–436
Singer G (1964) A simple aspirator for collecting small arthropod directly into alcohol. Ann
Entomol Soc Am 57:796–798
Singer G (1967) A comparison between different mounting techniques commonly employed in
acarology. Acarologia 9:475–484
Sircom J (2002) Photographic sampling: a photographic sampling method for mites on plants. Exp
Appl Acarol 24:55–61
Steykstal G, Murphy W (1986) Collecting and preserving insects and mites: tools and Techniques.
ARS. USDA Misc. Publication No. 1443
Tashiro H (1967) Self-watering acrylic cages for confining insects and mites on detached leaves.
J Econ Ent 60:354–356
Thind BB (2005) A new versatile and robust mite trap for detection and monitoring of storage
mites in the cereal and allied industries. Exp Appl Acarol 35:1–15
Travis BV (1968) Glyptal – a useful slide ringing compound. J Med Entomol 5:24
1. https://www.youtube.com/watch?v=e83shxkObNA Tracheal Mite Dissection

2. https://www.youtube.com/watch?v=mkY-BNZ3Jf4 Strawberry diagnostics: Using a mite brush
to sample for two-spotted spider mites
3. http://www.entsoc.org/PDF/Pubs/Periodicals/AE/AE-2008/Summer/Acorn.pdf Mite Brushing
Machine
4. https://www.youtube.com/watch?v=PUuC42_h1jQ Use of a Table Top Scanning Electron
Microscope
5. http://jenny.tfrec.wsu.edu/opm/displaySpecies.php?pn=5010 Mite Grid
6. https://www.youtube.com/watch?v=rSDwu6f-wDo How to Make an Insect Collection: Aquatic
7. https://www.youtube.com/watch?v=9ztZW5-Elo4 Insect Aspirator
8. https://www.youtube.com/watch?v=J5rGo3uBFIU How to Make a Berlese Funnel
Morphology and Anatomy of Acari
4
Abstract
Mites and ticks are potentially the largest group of chelicerate arthropods and are
assemblages of different forms along varying, yet distinctly phylogenetic lines.
Mites rival insects in adaptability to terrestrial and aquatic habitats but are more
closely related to spiders than insects. They share the characteristics of jointed
legs and a chitinous exoskeleton but also possess the arthropodan features of an
open circulating system, ventral nerve cord, alimentary canal and striated mus-
cles and may also possess Malpighian tubules. Unlike insects and many other
arthropods, Acari lack antennae and mandibles. Mouthparts of spiders (to whom
they are closely related) are always piercing and sucking type, but it is only in
Acari that exhibit parasitism, predaceous nature and phytophagous, mycopha-
gous and saprophagous nature in different groups. The acarine body is divided
into gnathosoma and idiosoma. Various distinct structures found in these two
regions are discussed in detail. The body covering or integument in acarines is
thoroughly discussed. Typical structures such as chelicerae and palps on gnatho-
soma and development of different shields, plates and legs are discussed through
comparative accounts in different families. Salient features of the digestive sys-
tem, central nervous system, circulatory system, excretory system and reproduc-
tive system are mentioned.
Keywords
Gnathosoma • Idiosoma • Chelicerae • Palps • Integument
The Acari are potentially the largest group of chelicerate arthropods and are assem-
blages of different forms derived along varying, yet distinctly phylogenetic lines.
Mites rival insects in adaptability to terrestrial and aquatic habitats but are more
closely related to spiders than insects. They share the characteristics of jointed legs
and a chitinous exoskeleton. Acari also possess the arthropodan features of an open

DOI 10.1007/978-981-10-1594-6_4
42 4 Morphology and Anatomy of Acari
circulatory system, ventral nerve cord, alimentary canal and striated muscles and
may also possess Malpighian tubules (in many cases) which collect and sequester
excretory products for their elimination. Unlike insects and many other arthropods,
Acari lack antennae and mandibles. Mouthparts of spiders (to whom they are closely
related) are always piercing and sucking type, but it is only in Acari that exhibit
parasitism, predaceous nature and phytophagous, mycophagous and saprophagous
nature in different groups.
4.1 External Morphology
The body tagma of the Acari comprises two regions: the gnathosoma (jaw body)
and the idiosoma (distinct body). Mites are the only arachnids in which the chelic-
erae are located in the body region – the gnathosoma – which is separate from the
region with the legs attached, the idiosoma (Andre and Remacle 1984; Beard et al.
2012).
4.1.1 Integument
The body covering or integument in the acarines is complex in structure and diverse
in function. Integument provides a protective exoskeleton and sites for muscle
attachment as well as lining the tracheal system and alimentary canal. The high
degree of permeability of the integument plays an important role in restricting water
loss from the body. Although showing considerable flexibility, the sclerotized non-
cellular component of the integument is limited in the extent to which it can expand
to allow for the increase in size. In order to accommodate growth and size, it is shed
periodically and replaced during the moulting cycle (Norton and Behanpelletier
1991; Norton et al. 1997; Pugh 1987; Romosor 2004).
4.1.1.1 Composition
The cuticle consists of a thick procuticle, which may be differentiated into an
endocuticle (=hypostracum) and a superficial wax cement or secretion layer, the
tectostracum or cerotegument. The relative thickness of the procuticle and epicuti-
cle varies from 0.25–2.0 μm to 0.05–1.5 μm, respectively, in case of two-spotted
spider mite, Tetranychus. The epicuticle comprises two layers: the inner epicuticle
(having homogeneous innermost layer) containing lipids, phenols and protein and
an outer epicuticle, lying immediately below the wax layer. The outer epicuticle is
the first part of each new cuticle to be secreted and covers almost the entire integu-
mentary surface of the acarines. The outermost cement layer, overlying the epicu-
ticle, protects the underlying wax layer. Hardening and subsequent darkening of the
outer procuticle result in sclerotization of the body (Fig. 4.1). Immature forms are
usually less sclerotized than adults, and there is a tendency during ontogenetic
development for a progressive fusion of shields to form more extensive area of
sclerotization. In general the epicuticle is uniform, whereas the procuticle and the
4.1 External Morphology 43
Fig. 4.1 Acarine cuticle (From Norton et al. (1997)). (a) Cross-section. (b) Epicuticular layers
secretory layer show functional adaptations. In haematophagous parasites, which

ingest large quantities of fluid, there is generally a reduction in the areas of sclero-
tized cuticle and an extensible folding of the epicuticle of the soft cuticle of the body
so that its surface appears corrugated. During growth the thickness of the cuticle
takes place chiefly in the endocuticle region (Evans 1992; Krantz and Walter 2009).
Heavily armoured species of the Oribatida and Mesostigmata have a uniformly
thick cuticle with the endo-, and exocuticular areas of the procuticle clearly defined.
Some species of Tetranychidae have the ridges of the cuticle provided with numer-
ous lobes which may serve as evaporative structures in feeding mites (absent in
diapausing females) or to function in increasing the rigidity of the cuticle. The sur-
face of the sclerotized areas of the cuticle can be variously ornamented particularly
in Mesostigmata and Ixodida. Other most common forms found are reticulate–areo-
late pattern and foveate, granulate, punctate and tuberculate forms.
Hard and soft, when describing ticks, reflect the extent to which the ticks are
covered by sclerotized, i.e. hardened cuticle. Ixodid ticks are referred to as hard
ticks because of their sclerotized (hardened) capitulum, scutum, appendages and
other small areas. On the other hand, argasid ticks are leathery or soft ticks because
sclerotized areas are relatively small. The cuticle, being an external skeleton, deter-
mines the maximum size and the shape that a particular developmental stage can
attain, and further growth is possible only by moulting, i.e. shedding the cuticle and
replacing it by a larger one. Development of cuticular outgrowths and depressions
of the cuticle for the protection of legs are found in the Oribatida and Uropodina
(Sanders and Norton 2004). The prodorsum of the Oribatida is sometimes provided
with blade or flange-like processes which protect the distal part of the anterior legs
when these are withdrawn to the body. Some cells of the epidermis are specialized
to perform special functions. For example, dermal glands and its ducts in argasid
ticks secrete the cement layer of the cuticle a few hours after the moulting. But in
ixodid ticks, the cement layer is lacking. Dermal or cuticular glands are also com-
monly found on the body and legs in Mesostigmata (Coons and Alberti 1999;
Griffiths et al. 1990; Hackman 1982).
4.1.1.2 Colouration
In many mites and ticks, typical body colouration is found. The pale colour of some
species may be due to pigments of ingested food. Similarly the green colour in phy-
tophagous mites and reddish-brown colour in haematophagous species are also
because of the liquid food they suck while feeding. Very little is known about the
pigments. Some colours have a physical origin and they change on immersion in
fluids. Sclerotization imparts a brown colour to the cuticle and so the capitulum,
scutum and appendages are dark in colour. Representatives of family Bdellidae,
Trombidiidae and many water mites are normally brightly coloured.
4.1.1.3 Moulting
During the latter part of the life of an instar, the body may become distended fol-
lowed by the cessation of feeding and locomotion. The exuvial phase is always a
pharate previous instar. A quiescent phase is a feature of the developmental cycle of
most Acari with the apparent exception of the Mesostigmata and argasid ticks. In
the Oribatida, about one-third time for total development may be spent in the quies-
cent state. The emergence of an instar from the cuticle of the preceding instar is
facilitated by the splitting of the old cuticle along a definite line of weakness known
as the ecdysial cleavage line. The rupture of the cuticle is largely achieved by hydro-
static pressure resulting from muscular contractions of the body. Ixodid ticks
undergo two moults in their progress from larvae to nymphs to adults. Argasid ticks
undergo three or more moults. Some argasid species have two nymphal instars, oth-
ers up to six or more (Hackman 1982; Woodring and Cook 1962).
4.1.2 Differences Between Hard Ticks and Soft Ticks
Hard ticks (family Ixodidae) and soft ticks (family Argasidae) are differentiated
from each other as given below (Krantz and Walter 2009; Sonenshine and Roe
2014):
(i) Scutum. In soft ticks, the scutum (dorsal shield) is absent. But in hard ticks, the
scutum is present in larva, nymphs and adults; male ticks have large scutum but
females have small scutum.
(ii) Mouthparts (capitulum). In soft ticks mouthparts are not visible from above in
nymphs and adults, but are seen in larvae. But in hard ticks, mouthparts are
visible dorsally in larvae, nymphs and adults (ER1 and ER2).
(iii) Cheliceral sheath. Cheliceral sheaths are smooth in soft ticks, but in hard ticks
sheaths are denticulate.
(iv) Palps. In soft ticks, palps look like legs. But in hard ticks, palps are
club-shaped.
(v) Coxal organs. Present in soft ticks, but are absent in hard ticks.
4.1.3 Body Divisions
Segmental amalgamation, or tagmosis, in the Acari has resulted in the formation of

two major body divisions: an anterior gnathosoma (commonly known as capitulum
in ticks and water mites) and the posterior idiosoma. The two tagmata articulate by
a circumcapitular furrow which may be obscured in some groups by secondary
somatic development (Evans 1992; Hoy 2011). A dorsosejugal furrow (also referred
to as sejugal furrow) may be found at a level between legs II and III in some opilioa-
cariform and acariform mites, and a postpedal furrow may also be found just behind
legs IV (ER17). The following nomenclature, however, is commonly used to clas-
sify the divisions of the acarine body:
Gnathosoma
(Segments of the mouth and its appendages)
Propodosoma
(Segments of the first and second legs)
Metapodosoma
(Segments of the third and fourth legs)
Opisthosoma
(Segments posterior to the legs)
Gnathosoma and propodosoma combined is also referred to as proterosoma,

while metapodosoma and opisthosoma combined are referred to as hysterosoma.
Propodosoma and metapodosoma combined have been referred to as prosoma in
literature (Fig. 4.2). In Parasitiformes, gnathosoma and idiosoma are the principal
regions and rarely any other division is found. Contrastingly, most of the Acariformes
mites have a well-defined sejugal furrow that divides the body into a propodosoma
(or proterosoma) and hysteosoma (Hoy 2011; Wooley 1988). In some of these
mites, a postpedal furrow divides the body into a narrowed opisthosoma behind the
fourth pair of legs (as noticed in families Demodicidae and Halarachnidae).
4.1.3.1 Gnathosoma
The gnathosoma of the Acari resembles the head of a generalized arthropod body in
that the mouthparts are appended to it (Fig. 4.3). But the brain lies in the idiosoma
region behind the gnathosoma, and the ocelli, if present, are situated dorsally or
Fig. 4.2 Major body

divisions
dorsolaterally on the propodosoma. Gnathosoma has also been referred to as the

capitulum (false head) in some literature especially while describing ticks. So gna-
thosoma cannot be called a head, even though it is head-like (as in Ixodida,
Erythraeidae and water mites) because the brain (synganglion) is situated in between
the legs. Gnathosoma is never fused with the idiosoma; rather, it is movable at the
intersegmental connection (ER8, ER11, ER13 and ER16).
Hypostome Ventrally the extensions of the palpal coxae meet in the midline to
form the hypostome and deuterostome in gamasids and other mites. The endites of
the palpal coxae become the hypostome and anteriorly may be divided into internal
malae and corniculi (=external malae). In Ixodid ticks, hypostomal development is
specialized in the form of a tonguelike structure with many retrorse (recurved) teeth
and forms the ventral floor of the buccal cavity. Ventral aspects of gnathosoma are
also known as infracapitulum, subcapitulum and hypognathosoma (ER9, ER10 and
ER 12).
The tick’s body consists of the capitulum and the idiosoma. The mouthparts –
chelicera, palps and hypostome – are found in the capitulum. The two outermost
structures of the mouthparts are the two highly mobile palps; between them are the
paired chelicerae which in turn protect the hypostome which is the central rod-
shaped structure (ER3 and ER5). The hypostome is the primary organ for attach-
ment to the host’s skin. It is covered with numerous recurved teeth that are placed
into the hosts’ skin. The teeth located on the hypostome progressively get smaller
and fewer towards proximal regions. The hypostome itself is hollow except for the
Fig. 4.3 (a) Gnathosoma and tritosternum in Opilioacaridae (ventral aspect). (b) Chelicera and
palps in spider mites. (c) Gnathosoma (venter) of oribatid mites. (d) Gnathosoma (ventral aspect)
of ixodid ticks
distal end. Deep groove on the dorsal side of the hypostome allows the blood from
the host to be brought into the mouth and pharynx (Krantz and Walter 2009)
Gnathosoma may be shielded by a hood and can be retractable. The anterior
dorsal prominence is referred to as epistome (= tectum capitulum); ventrally the
dorsal shield is formed into a cavity (into which the mouthparts can be retracted)
and is known as camerostome (vaulted chamber) as in Uropodida and Oribatida.
The infracapitulum or subcapitulum forms the anterior part of the gnathosoma, and
its external walls are formed by the enlarged coxae of the palp (= pedipalp) which
meet and fuse ventrally (Baker 1990). The sterna of the gnathosoma have different
parts, namely, presternum associated with the chelicerae and the deutosternum
associated with the palps. The deutosternum sometimes is extended to become part
of the hypostome. In Pyemotidae, the gnathosoma is cuboidal in shape with stubby
palps and stylettiform chelicerae. Square-shaped gnathosoma is found in family
Tarsonemidae. An elongated gnathosoma is found in snout mites (Bdellidae and
Cunaxidae) and in lizard mite family Pterygosomidae. The gnathosoma may be in
the form of a truncated tunnel-like tube as in Cryptognathidae (Wooley 1988).
Chelicerae
The chelicerae are the first and only preoral pair of appendages that lies dorsal to the
mouth. They are usually three-segmented in both Acariformes and Parasitiformes.
The primitive state of the chelicerae is chelate–dentate in majority of the mites.
With the pincer-like distal end, the chelicerae are capable of grasping, pinching,
shearing and sometimes even penetrating their hosts. In more specialized chelic-
erae, there is a reduction to two segments that results in the piercing organ, but
without the capability to grasp. Chelicerae of mites usually terminate in a chela that
is composed of a dorsal fixed digit and ventral movable digit. In males of many
Mesostigmata, the movable digit is modified for sperm transfer (spermadactyl). In
ticks, the chelicerae are the cutting organs and are used to penetrate the hosts’ skin
and access to its blood. The movable digits are equipped with broad heavily sclero-
tized cusps or spinelike teeth. All of the cusps are oriented in the horizontal plane so
that most of the cutting action is lateral. In essence tick chelicerae can cut but cannot
grasp as they cannot rotate or pronate. The digits are used to rip and tear the host’s
skin to expose the dermis. When the digits are not being used, they are retracted
within the cheliceral sheaths that surround the elongated shaft. Some mechanosen-
sory, chemosensory and thermosensory sensilla are found on the digit spines.
Together these sensilla provide information on the biochemical characteristics of
the wound site environment.
The shape of the chelicerae depends on the feeding habits of the mites. For exam-
ple, in predaceous mites the chelicerae are quite massive with large teeth. In para-
sitic mites (Rhinonyssidae), the chelicerae are quite reduced and edentate. Chelicerae
are modified for grasping (Eviphididae), needle-like or stylettiform (Tetranychidae
and Tuckerellidae); strongly hooked and sickle-shaped (Anystidae); serrated pierc-
ing organs (Trombiculidae), or may be formed into cutting tips (Ixodida). In
Collembola feeding mites (Veigaiidae), elongate and dentate chelicerae are found.
But in fungivorous mites, chelicerae are quite massive with closely set teeth.
Gamasid mites that feed on nematodes have short cheliceral units and long teeth.
However, most of the Gamasida, Astigmatina and Oribatida possess chelate–dentate
chelicerae.
Palps
The palps are articulated to the palpcoxal region of the gnathosoma, but in majority
of the mites, they are free and leglike. Four to six segmented palpi are found in dif-
ferent orders of mites. There are six movable segments – the trochanter, femur,
genu, tibia, tarsus and apotele. The palps are generally simple but exhibit many
variations in different families. Most mites lack a claw on the palp tarsus. In families
Rhinonyssidae and Halacaridae, the palps are much reduced and are closer to the
mouth. Palps may be used for sensory purposes and grasping the food and some-
times are also used for cleaning the chelicerae. In Ixodida, usually segmented palps
are found; individual segments are of different lengths and are modified in shape
enclosing the hypostome. But in soft ticks (Argasidae), segments of the palp are
subequal and normally looking. Palpi of gamasid and holothyrid mites are compa-
rable to soft ticks.
Great variations in the form of palpi are found in Acariformes; they are very
small in members of Pyemotidae, Tarsonemidae, Eriophyidae and Demodicidae.
The palp in case of Pyemotidae consists of only one segment. Palps are simple in
families Bdellidae and Tydeidae. Distinctively the palp tarsus is relatively swollen
in Rhagidiidae, but in Bdellidae the palp tarsus is rectangular in shape and serves as
a useful character for their identification. Palps are much reduced in Astigmatina
and are held close to the subcapitulum. The palps in bdellid mites are of tactile and
chemosensory nature. Palps are raptorial in predatory mite families Halacaridae,
Cheyletidae and Cunaxidae. Palps bear a distinctive thumb–claw complex in many
spider mites (Tetranychidae), rake-legged mites (Caeculidae) and many trombidi-
form mites. The tibiotarsus or tarsus region of the palp bears some chemo- and
mechanoreceptors (Nuzacci and DiPalma 2002).
4.1.3.2 Idiosoma
The idiosoma comprises the entire body posterior to the gnathosoma and hence
assumes the functions parallel to those of the abdomen, thorax and parts of the head
of insects and other arthropods. A great diversity of form, chaetotaxy and degree of
sclerotization of idiosoma is found in different groups of acarines. The primary
external structures of the idiosoma are involved in locomotion, respiration, sperm
transfer, sensory reception, secretion, and osmoregulation.
Legs
Legs are attached to the idiosoma in different locations. In most mite groups, both
adults and nymphal stages possess four pairs of jointed legs but the larva has three
pairs. Fourth pairs of legs appear with the first nymphal stage. However, some
exceptions do occur in different prostigmatid and astigmatid mites. In family
Eriophyidae, wormiform mites possess only two pairs of legs in all stages after
hatching. Males of some members of family Podapolipidae possess only three pairs
of legs (Larvacarus spp. and Podapolipus spp.), but females have only one pair of
legs.
Typically the legs comprise seven primary segments – the coxa, trochanter,
femur, genu (also called patella), tibia, tarsus and pretarsus (Fig. 4.4). But in follicle
mites (Demodicidae), legs are short, are telescoped and are four segmented. In some
groups of mites, primitive secondary division of trochanters III–IV is retained while
basitarsal and basifemoral features are found. Amalgamation of the femur and genu
or of the tibia and tarsus also sometimes results in the formation of a femorogenu or
a tibiotarsus. The terminal pretarsus when present typically comprises a basilar
sclerite and a set of distal structures – a median empodium, a pad or sucker-like
Fig. 4.4 Leg segments

(generalized)
Fig. 4.5 Generalized
acarine ambulacrum
pulvillus and a pair of claws. The pretarsus is known as an ambulacrum when

median elements are present. The ambulacrum itself is a highly variable complex in
which some elements may be greatly modified or lost, for example, the empodium
may be clawlike or padlike. The tarsi of legs are variously modified and serve as an
important diagnostic character and significantly contribute for identification at fam-
ily, genera and even species level. Claws, caruncles, apoteles and ambulacra are
important features in differentiating different acarine orders or other subdivisions
(Fig. 4.5). In ticks, legs are divided into six segments: coxae, trochanter, femur,
tibia, metatarsus and tarsus. The coxae articulate with the body wall through mem-
branous connections which allow some mobility in retraction and extension. Distally
tarsus ends in a pair of claws and a pulvillus (Ixodidae) or claws only (Argasidae).
Haller’s organ is a typical sensillum present on the dorsal surface of first legs in both
hard and soft ticks (ER14).
Legs are primarily locomotory in function, but in some species may be modified
to serve for grasping, anchoring, jumping and swimming purposes. Sometimes legs
may be equipped with some spinelike or spur-like apophysis which may help in
capturing prey. Legs may also be modified for clasping hairs as in some parasitic
mites (Myobiidae and Listrophoridae). In nasal mites (Rhinonyssidae), claws of
legs are used as surrogate chelicerae for piercing and tearing the mucous nasal
membrane of their bird hosts.
Other Idiosomal Structures

Many times sclerotized shields or plates are present on the different dorsal or ventral
areas of the idiosoma (Figs. 4.6 and 4.7). These shields may be weakly tanned and
are even difficult to observe in some acariform mites, but the shields are strongly
developed and heavily tanned in many parasitiform mites. The degree of develop-
ment of different shields in most acarines usually increased through ontogenetic
development. Idiosomal shields provide a degree of protection from predation and
environmental extremes and also serve as sites for muscle attachment. In ticks the
dorsal shield may be covering only a small portion of the propodosoma, or it may
extend laterally and posteriorly to cover all of it. In Mesostigmata, prodorsal shields
are extended to cover the entire podonotum region and are known as podonotal
shield. Other sclerotized shields based on their position on the idiosoma are desig-
nated as opisthonotal shield, mesonotal shield and pygidial shield. Sometimes in
Oribatida, the proterosoma is covered by a proterosomatic shield that completely
obscures the gnathosoma dorsally. The opisthosomatic shield not only forms a cov-
ering dorsally but also extends lateroventrally to provide a shell-like circumgastric
‘notogaster’. Fusion of various podonotal or hysteronotal shields many times is
found in Mesostigmata. Dorsal and ventral hysterosomatic shield complexes may
be entirely absent in many Prostigmata (including representatives of Eupodides and
Raphignatha). Dorsal shields generally display patterns of ornamentation which
sometimes prove as diagnostic characters. Ventrally the idiosoma may be virtually
unsclerotized or it may carry a variety of shields. In general basal leg articulations
and genital anal apertures are almost invariably provided with some surrounding
sclerites. Ventral shield expansion, reduction and fusion into more or less fixed pat-
terns are commonly found in the Holothyrida, Mesostigmata and Oribatida and
serve as useful taxonomic characters for higher categories or even for the identifica-
tion of species. Festoons – small portions of the ventral posterior body margin – are
marked by delicate grooves which are usually rectangular and are found in some
ixodid ticks.
4.1.4 Sensory Receptors
Many types of sensory receptors like mechanoreceptors that respond to tactile stim-
uli are found on different areas of the body (Fig. 4.8). Most mechanical receptors
are simple and spinose, or they may be ornamented, often forming distinctive pat-
terns in the idiosoma serving as useful characters in identification of different
Fig. 4.6 Sclerotized ventral structures of mesostigmatid mites
taxonomic categories. Setae may serve as a measure of protection from potential

predators. Specialized tactile setae, namely, trichobothria, are found in small num-
bers on the body and on leg appendages of some Opilioacarida. Trichobothria may
be hair-like and are greatly elongated in some Acariformes (Bdellidae). One or two
pairs of simple, pectinate or clavate bothridial setae are found on the prodorsum of
many prostigmatid, endeostigmatid and oribatid mites. These setae may perform
anemotactic or acoustic function. In acariform family Tydeidae and in chiggers
(Trombiculidae), a pair of sensilla may arise from the propodosoma. In family
Fig. 4.7 Ventral shields

(ixodid ticks)
Fig. 4.8 Acarine setae (From Krantz and Walter (2009)). (a) Tactile seta. (b) Solenidia. (c) Famuli.
(d) Trichobothria
Tarsonemidae and Pyemotidae, a characteristic paired sensilla also is found

(Crossley 1960).
Proprioceptors – distinctive pore-like openings which serve as sensory struc-
tures – are present on the legs, palpi, chelicerae (in Opilioacariformes and
Parasitiformes) and idiosoma throughout the Subclass Acari. Proprioceptors may be
in the form of small, membrane-covered clefts, slit sense organs or poroids.
Chemosensory organs are located on the terminal segments of legs I–II and on
the palpi. Eupathides (acanthoides and pseudacanthoides) identified by the symbol
ζ (zeta) are spinose, apically rounded or divided structures found on the extremities
of tarsi I–II (rarely on tarsi III) and on the tip of palptarsus. Eupathidia may function
as contact or as gustatory chemoreceptors. A special hollow seta found at the tip of
web-spinning tetranychine spider mites (Tetranychidae) is generally considered as
a eupathidium, although it functions in delivery of silk received from an associated
underlying gland or series of glands. Neither palpal glands nor terminal hollow
palpal hairs are present in tetranychid subfamilies that do not produce silk.
Eupathidia are present at the terminus of the larval and protonymphal palptarsi of
some Anystidae. The famulus designated as ε (epsilon) is similar to the eupathidia
in having a pore on tarsus I (sometimes on tarsus II) in the Acariformes. In addition
to eupathidium and famuli, the legs and palpi of acariform mites sometimes carry
porose and thin-walled solenidia. The solenidia may be terminally rounded or
pointed setiform or peg-like structures that lack the actinopilin core or sheath which
is a characteristic of eupathidia and famuli. Solenidia are found on the tarsus of palp
and on the tarsus and tibia of legs. Solenidia are considered olfactory in function.
Unlike the Acariformes, all of the normal setae and setiform sensory structures of
the Parasitiformes are optically inactive (Evans 1992; Ivanov and Leonovich 1983;
Kettle 1995; Krantz and Walter 2009).
Tarsus I of ticks (Ixodida) carries a distinctive dorsal sensory field commonly
known as Haller’s organ – this complex comprises an anterior pit and a more poste-
rior capsule, each of which contains a variety of sensilla which responds to different
stimuli (Ivanov and Leonovich 1983; Klompen and Oliver 1993). Porose, thin-
walled olfactory receptors are located in the deep capsule portion of Haller’s organ.
A sensory organ similar to Haller’s organ has also been reported in some
Mesostigmata and in holothyrid family Allothyridae.
In many acariform mites, a pair of urstigma or Claparede organs is present in
larvae on the ventral region of coxae I–II. In case of acarine orders Opilioacarida,
Holothyrida, Ixodida and Prostigmata, one to two pairs of simple ocelli may be
present on the anteromedian or anterolateral aspect of the prodorsum region. In
Bdelloidea, Tetranychoidea and Anystoidea, one pair of lateral eyes is commonly
found, but unpaired in acariform mites belonging to family Halacaridae and
Hydryphantidae. The median eye or pair of eyes is commonly found on the under-
side of a ‘naso’ – an anteromedial prodorsal projection in many prostigmatid and
oribatid mite families (Krantz and Walter 2009; Wooley 1988).
In Oribatida and some Prostigmata, genital papillae in protonymph (one pair),
deutonymph (two pairs), tritonymphs (three pairs) and adults are present. In
Hydrachnidiae, more than 100 genital papillae are found. In earlier literature,
4.2 Internal Anatomy (Fig. 4.9) 55
genital papilla has been referred to as acetabula as in water mites (e.g. Parasitengona).
In some recent literature, urstigmata and genital papilla are reported to have a role
in respiration (Evans 1992).
4.2 Internal Anatomy (Fig. 4.9)
Most of the information given below is based on the publications of Sauer and Hair
(1986), Evans (1992) and Hughes (1954). A technique for examination of internal
anatomy of ixodid ticks is nicely shown in ER15.
4.2.1 Digestive System
The embryogenesis and functional subdivisions of the acarine alimentary canal

closely resemble that of insects. There are three basic gut regions: the foregut and
hindguts are of ectodermal origin and are lined with a chitinous intima; the midgut
originates endodermally. In the fully developed alimentary canal, the mouth opens
into a muscular pharynx, followed by the oesophagus which passes through the
ganglionic mass. The midgut often is diverticulate, i.e. it has several fingerlike
Fig. 4.9 Dorsal internal

aspect (Mesostigmata)
(From Evans (1992))
projections for food storage especially in parasitic mites, and is capable of great
extension especially in blood-sucking ticks. Most digestion and absorption occur in
the midgut region. Some mites lack a connection between the midgut and hindgut;
such mites feed only on fluids but do not defecate. The hindgut in these mites is
transformed into an excretory organ for elimination of nitrogenous wastes. Other
mites with entire guts may have Malpighian tubules, extending from the junction of
the midgut and hindgut, as excretory organs. Coxal glands, salivary glands, genital
papillae and Claparede organs and haemocytes are the cells that serve as the immune
system of Acari (Agbede and Kemp 1985; Akov 1982; Alberti et al. 2003; Brody
et al. 1976; Cones and Roshdy 1973; Hughes 1950; Mothes 1981).
4.2.2 Central Nervous System
The central nervous system is composed of a ganglionic mass or synganglion (brain)

that surrounds the oesophagus. The ganglionic mass is divided by the oesophagus
into two regions: the pre-oesophageal or the supra-oesophageal and post-oesophageal
or sub-oesophageal. The pre-oesophageal region innervates the photoreceptors and
the gnathosoma, while the post-oesophageal region innervates the remaining parts
of the body. Mites and ticks also possess a variety of sensory structures on their
body surface, similar to sensory structures in insects (Alberti and Crooker 1985;
Blauvelt 1945; Coons et al. 1974).
4.2.3 Circulatory System
Acari have an open circulatory system in which the more or less clear haemolymph
circulates in the haemocoel and bathes the internal organs. The haemolymph distrib-
utes food materials and waste products and contains haemocytes, the cells that serve
as the mite’s immune system. But it does not contain oxygen-binding proteins as
found in blood of vertebrates and in some arthropods (Binnington and Obenchain
1982).
4.2.4 Excretion
Malpighian tubules that open into the hindgut are present in several groups of Acari
and eliminate several nitrogenous wastes. In acarines that lack Malpighian tubules,
a portion of the midgut carries out the necessary desired function of excretion.
Water and ion balance involve several glands, e.g. coxal glands, salivary glands,
genital papillae and Claparede organs. In ixodid ticks, much of the excess water in
blood meals is excreted back into the host via the salivary glands (Blauvelt 1945;
Romosor 2004).
4.2 Internal Anatomy (Fig. 4.9) 57
4.2.5 Respiratory System
Ventilation is achieved by a system of the trachea that opens to the outside via open-
ings known as stigmata. In some groups, namely, Endeostigmata, Oribatida and
Prostigmata, respiration is cuticular, but in other groups internal exchange of oxy-
gen and carbon dioxide is facilitated through a branched tracheal system that opens
externally through stigmata. The presence or absence of stigmata and their relative
positions is the major diagnostic character for identification of different orders and
suborders of subclass Acari. Plastron respiration is also found in mites living in
marine conditions.
4.2.6 Reproductive System
The internal reproductive system typically consists of a single ovary (but paired in
the Astigmata) in the female and paired testes in the male. Females typically possess
spermatheca for sperm storage after insemination. Both sexes may have various
accessory glands. Both the germinal tissues (testes and ovaries) are mesodermal in
origin and lead to the outside by means of ducts (oviducts in females and vasa def-
erentia in males): vagina in females and ejaculatory duct in case of males which are
ectodermal in origin. In most cases the external genital opening or gonopore is
located ventrally. In males there may be a seminal vesicle inserted between the vasa
deferentia and the ejaculatory duct, a portion of which may be elaborated into an
intromittent organ or aedeagus. A distinct aedeagus may be present in males of
many Acariformes families particularly Cheyletidae, Myobiidae, Demodicidae,
Acaridae and Podapolipidae. Accessory glands may open into the seminal vesicle or
ejaculatory duct and contribute to the production of seminal fluid. Accessory glands
are commonly found in Ixodida but are absent in Oribatida, most Prostigmata and
Astigmata. In female, a receptacle may be associated with the vagina and accessory
glands may be present. However, in some acarines there may be a sperm receptacle
or spermatheca. Sperms, usually within a spermatophore, are introduced into the
female by direct deposition or are deposited on the substrate and then are taken up
by the female (Akimov and Yastribtrov 1984; Blauvelt 1945; Dunlop and Alberti
2007; Lees and Beament 1948; Nitalinski et al. 1990; Woodring and Carter 1974).
4.2.7 Musculature
In acarines muscles are striated and a complex skeletal musculature is involved

regarding the movements of the various appendages. Both intrinsic and extrinsic
types of muscles are found in mites. Intrinsic muscle actions extend over the joints
of leg segments and are flexors. Intrinsic muscle actions result in flexion. Extrinsic
muscles consist of dorsoventral oblique rotator and elevator muscles. Extrinsic
muscles originate and insert over the body wall. The chelicerae in most mites and
ticks are retracted by the action of skeletal muscles through the haemolymph by
contraction of muscles in the idiosoma. Longitudinal, circular and irregular net-

works of visceral muscles affect peristalsis in various tubular organs such as the
alimentary canal (Evans 1992; Wooley 1988).
4.3 Conclusions
Significant volume of literature dealing with taxonomy, biology crop or animal

damage, respectively, by phytophagous and animal parasitic mites is found. The
research has generated a great deal of valuable information about their impact on
several crops and animals. Many morphological studies are found regarding the
structure and function of different types of integument, setae and other sensory
structures present on the dorsal surface of body (both gnathosoma and idiosoma)
regions. Patterns of different body shields and sclerotization of region of append-
ages are commonly reported on different Parasitiformes mites. In ticks, which are
quite big in size than mites, more detailed studies regarding their morphology and
anatomical details are reported. In case of Acariformes mites, many types of stria-
tion patterns on the body and other patterns of tubercles are found. These studies
have definitely helped in identification at higher levels and even up to species and
generic levels. Such structures help in maintaining water balance. In Acariformes,
more studies on morphological and anatomical respects have been reported for the
most important families like Tetranychidae, Eriophyidae, Tenuipalpidae,
Trombiculidae, Sarcoptiformes, Acaridae and Psoroptidae, and their relations in
managing them have also been reported. In case of ticks which are serious pests of
farm animals and human beings, feeding on blood as well as transmitting various
bacterial diseases as vectors, more detailed studies with modern techniques are
reported. Detailed physiological studies on digestion, respiration, the nervous sys-
tem, the reproductive system and other organ systems at molecular levels still
remain to be done to find answers for their proper management under different habi-
tats. More intensive studies on different aspects of morphology through better
known techniques like scanning electron microscopy utilizing LT-SEM and other
modern techniques are needed which may help in identification at higher levels and
even up to species and generic levels.
Further Reading
Achor DS, Ochoa R, Erbe EF, Aguilar H, Wergin WP, Childers CC (2001) Relative advantages of
low temperature versus ambient temperature scanning electron microscopy in the study of mite
morphology. Int J Acarol 27:3–12
Agbede RIS, Kemp DH (1985) Digestion in the cattle tick Boophilus microplus (Canestrini): light
microscope study of the gut cells in nymphs and females. Int J Parasitol 15:147–157
Akimov IA (1989) Muscular system and elements of the skeleton of Nothrus palustris (Acariformes:
Oribatida). Zool Zh 48:57–66
Akimov IA, Yastribtrov AV (1984) Reproductive system of Varroa jacobsoni I. Female reproduc-
tive system and oogenesis. Verten Zool 6:61–68
Further Reading 59
Akov S (1982) Blood digestion in ticks. Curr Them Trop Sci 1:197–211
Alberti G, Crooker AR (1985) Internal anatomy. In: Halle E, Sabelis MW (eds) Spider mites: their
biology, natural enemies and control, vol 1A. Elsevier Science Publishers B.V., Amsterdam,
pp 29–62
Alberti GA, Seniczak A, Seniczak S (2003) The digestive system of fat body of an early- derivative
oribatid mite, Archegozetes longisetosus Aoki (Acari: Oribatida: Trhypochthoniidae).
Acarologia 43:149–219
Andre HM, Remacle C (1984) Comparative and functional morphology of the gnathosoma of
Tetranychus urticae (Acari: Tetranychidae). Acarologia 25:179–190
Baker AS (1990) A survey of external morphology of mites of the superfamily Europodoidea 1894
(Acari: Acariformes). J Nat Hist 24:1227–1261
Balashov YS (1972) Blood sucking ticks (Ixodoidea) – vectors of diseases of man and animals.
Misc Publ Entomol Soc Am 8:161–376
Beard JJ, Ochoa R, Bauchan GR, Welbourn WC, Pooley C, Dowling APG (2012) External mouth-
part morphology in the Tenuipalpidae (Tetranychoidea): Raoiella a case study. Exp Appl
Acarol 57:257–269
Binnington KC, Obenchain FD (1982) Structure and function of the circulatory, nervous and neu-
roendocrine systems in ticks. Curr Them Trop Sci 1:351–398
Blauvelt WE (1945) The internal morphology of the common red spider mite (Tetranychus telarius
Linn.). Mem Cornell Univ Agric Exp Station 270:1–35
Brody AR (1970) Observation on the fine structure of the developing cuticle of a soil mite Oppia
coloradensis (Acarina: Cryptostigmata). Acarologia 12:421–431
Brody AR, McGrath JC, Wharton GW (1976) Dermatophagoides farinae: the digestive system.
J NY Entomol Soc 80:152–177
Cones LB, Roshdy MA (1973) Fine structure of the salivary glands of unfed Dermacentor varia-
bilis (Say) (Ixodoidea: Ixodidae). J Parasitol 59:900–912
Coons LB, Alberti G (1999) The Acari – ticks. In: Harrison FW, Foelis R (eds) Microscopic anat-
omy of invertebrates, vol 8B, Chelicerate Arthropoda. Wiley, New York, pp 267–514
Coons IB, Roshdy MA, Axtell RC (1974) Fine structure of the central nervous system of
Dermacentor variabilis (Say), Amblyomma americanum (L.), and Argas arboreus Kaiser,
Hoogstraal and Kohls (Ixodoidea). J Parasitol 60:687–698
Crossley DA (1960) Comparative external morphology and taxonomy of nymphs of the
Trombiculidae (Acarina). Univ Kans Sci Bull 40:135–321
Dunlop JA, Alberti G (2007) The affinities of mites and ticks: a review. J Zool Syst Evol Res
46:1–18
Evans GO (1968) The external morphology of the postembryonic developmental stages of
Spinturnix myoti Kol. (Acari: Mesostigmata). Acarologia 10:589–608
Evans GO (1992) Principles of acarology. CAB International, Wallingford
Griffiths DA, Atyeo WT, Norton RA, Lynch CA (1990) The idiosomal chaetotaxy of astigmatid
mites. J Zool (Lond) 220:1–32
Hackman RH (1982) Structure and function of tick cuticle. Annu Rev Entomol 28:75–95
Hoy MA (2011) Agricultural acarology: an introduction to integrated mite management. CRC
Press, New York, 410 pp
Hughes TE (1950) The physiology of the alimentary canal of Tyrophagus farinae. Q J Microscop
Sci 91:45–61
Hughes TE (1954) The internal anatomy of the mite Listrophorus leuckarti (Pagenstecher). Proc
Zool Soc Lond 124:239–256
Ivanov VP, Leonovich SA (1983) Sensory organs. In: Balashov YS (ed) An Atlas of Ixodid tick
ultrastructure. Entomological Society of America Special Publication, Annapolis
Jongejan L, Uilenberg E (2004) The global importance of ticks. Parasitology 12:53–514, UK:
Cambridge University Press, (DOI:10-1017/S0031 18200–4005967)
Kettle DS (1995) Medical and veterinary entomology, 2nd edn. CABI Publishing, New York,
720 pp
Klompen JSH, Oliver JH (1993) Haller’s organs in the soft ticks (Acari: Parasitiformes: Ixodida).
J Parasitol 79:591–603
Krantz GW, Walter DE (eds) (2009) A manual of acarology, 3rd edn. Texas–Tech University Press,
Lubbock, 807 pp
Lees AD, Beament JWL (1948) An egg waxing organ in ticks. Q J Microscop Sci Lond
98:291–332
Lindquist EE (1996) External anatomy and notation of structures. In: Lindquist EE, Sabelis MW,
Bruin J (eds) Eriphyoid mites: their biology, natural enemies and control, vol 6, World crop
pests. Elsevier, Amsterdam, pp 3–31
Mills LR (1973) Morphology of glands and ducts in the two spotted spider mite. Tetranychus urti-
cae Koch. Acarologia 15:218–236
Mitchell RD (1962) The structure and evolution of water mite mouthparts. J Morphol 110:41–59
Mitchell RD (1964) The anatomy of an adult chigger mite Blankaartia acuscutellaris (Walch).
J Morphol 114:373–391
Morgan CVG (1960) Anatomical characters distinguishing Bryobia arborea and B. praetiosa
(Acarina: Tetranychidae) from various areas of the world. Can Entomol 92:595–604
Mothes U (1981) Functional microscopic anatomy of the digestive system of Tetranychus urticae
(Acari: Tetranychidae). Acarologia 22:257–270
Nitalinski W, Szlandak E, Boczek J (1990) Anatomy and ultrastructure of the reproductive system
of Acarus siro. Exp Appl Acarol 10:1–31
Norton RA, Behanpelletier VM (1991) Calcium carbonate and calcium oxalate as cuticular hard-
ening agents in the oribatid mites (Acari: Oribatida). Can J Zool 69:1504–1511
Norton RA, Alberti G, Weigman G, Woas S (1997) Porose integumental organs of oribatid mites
(Acari: Oribatida). I. Overview of types and distribution. Zoologica (Stuttgard) 146:1–31
Nuzacci G, DiPalma A (2002) Mouthparts of a tydeid mite: an ultrastructural and functional inves-
tigation. Entomologica (Bari) 36:71–91
Proctor HC (ed) (2006) Aquatic mites: from genes to communities. Kluwer, Dordrecht
Pugh PJA (1987) A comparison of the structure and function of the cerotegument in two species of
Cryptostigmata (Acarina). J Nat Hist 21:603–616
Romosor WS (2004) Introduction to arthropods: structure, function and development. In: Eldridge
BF, Edman JD (eds) Medical entomology. Kluwer Academic Publishers, Dordrecht, pp 13–52
Roshdy MA, Hefnawy T (1973) Functional morphology of Haemaphysalis spiracles (Ixodoidea:
Ixodidae). Z Parasitenkd 42:1–10
Sanders FH, Norton RA (2004) Anatomy and function of the ptychoid defensive mechanism in the
mite Euphthiracarus cooki (Acari: Oribatida). J Morphol 259:119–154
Sauer JR, Hair JA (eds) (1986) Physiology of ticks. Ellis Harwood, Chichester, 508 pp
Sonenshine DE, Roe RM (2014) Biology of ticks, vol I, 2nd edn. Oxford Univesity Press, London
Van der Hammen L (1982) Morphology and postembryonic development of Rhizoglyphus echino-
pus (Fumouze & Robin) (Chelicerata: Actinotrichida). Zool Meded Leiden 56:237–258
Wharton GW, Brody AR (1972) The peritrophic membrane of the mite Dermatophagoides fari-
nae: Acariformes. J Parasitol 58:801–804
Woodring JP, Carter SC (1974) Internal and external morphology of the deutonymph of
Caloglyphus boharti (Arachnida: Acari). J Morphol 144:275–295
Woodring JP, Cook EF (1962) The internal anatomy, reproductive physiology and moulting pro-
cess of Ceratozetes cisalpinus (Acari: Oribatei). Ann Entomol Soc Am 55:164–181
Wooley TA (1988) Acarology: mites and human welfare. Wiley, New York
1. http://www.cvbd.org/en/tick-borne-diseases/about-ticks/general-morphology/ Morphology
and Anatomy of Ticks
Further Reading 61
2. http://www.discoverlife.org/mp/20q?search=Amblyomma+maculatum&guide=Arachnida
External Morphology of Ticks
3. http://smithsonianscience.si.edu/2015/02/tickstick/ How do ticks…tick?
4. http://us-tick-key.klacto.net/ Interactive Identification Key for the Hard Ticks (Ixodidae) of the
Eastern U.S.
5. https://www.youtube.com/watch?v=7sGvuoHIjjo Tick Anatomy
6. https://www.youtube.com/watch?v=OJrYOYbmF0M (Different ticks and mites)
7. http://www.cvbd.org/en/tick-borne-diseases/ Tick-Borne Diseases
8. https://www.youtube.com/watch?v=3JNprSWv96w Introduction to the Acari
9. http://www.discoverlife.org/mp/20q?search=Amblyomma+maculatum&guide=Arachnid
Tick anatomy
10. https://www.youtube.com/watch?v=WpOyWesQtgc Tick animation
11. https://www.youtube.com/watch?v=knweGM7yZDg Tick Morphology
12. www.afrivip.org/sites/default/files/identification_complete_1.pdf Ticks: Tick identification
13. http://www.ars.usda.gov/SP2UserFiles/person/333/External%20mouthpart%20morphol-
ogy%20in%20the%20Tenuipalpidae%20Raoiella%20a%20case%20study.pdf External
mouthpart morphology in the Teunipalpidae (Tetranychoidea): Raoiella a case study
14. http://www.cvbd.org/en/tick-borne-diseases/about-ticks/general-morphology/locomotion/
Leg of Tick and locomotion
15. http://www.midsouthentomologist.org.msstate.edu/pdfs/Vol2_1/vol2-1_004.pdf Examination
of the Internal Morphology of the Ixodid Tick
16. https://www.youtube.com/watch?v=knweGM7yZDg (Morphology of Tick)
17. http://itp.lucidcentral.org/id/mites/invasive_mite/Invasive_Mite_Identification/key/0_
Glossary/Mite_Glossary.htm#Vertical-scapular system Glossary of Acarine Terms
Classification of Subclass Acari
5
Abstract
Mites are the most diverse and abundant of all arachnids, but because of their
very small size (usually less than a millimetre in length), we rarely see them with
the unaided eye. Mites are ubiquitous and have successfully colonized nearly
every known terrestrial, marine and freshwater habitat including polar and alpine
extremes, tropical lowlands and desert barrens and surface and mineral soils.
More than 55,000 species of mites have been described – a small fraction (about
5 %) of the total number of animal species alive today. Subclass Acari is divided
into two superorders – Parasitiformes and Acariformes. More than 15,000 spe-
cies are reported under Parasitiformes, while more than 40,000 species have been
described under Acariformes. Superorders have been further classified into
orders and suborders mentioning salient diagnostic characters under each cate-
gory. Comparative status of orders, suborders and cohorts as described in differ-
ent systems of classifications has been clearly given in this chapter. More than
896 species of ticks have been authentically described in family Ixodidae and
family Argasidae under order Ixodida. Acariformes are more diverse of the two
superorders and have been reported under 351 different families.
Keywords
Acariformes • Parasitiformes • Ixodida • Trombidiformes
Mites are the most diverse and abundant of all arachnids, but because of their small
size (usually less than a millimetre in length), we rarely see them. The ticks are an
exception, in that they are filled with blood. Mites are also among the oldest of all
terrestrial animals with fossils shown from the early Devonian (nearly 400 million
years ago). Mites are truly ubiquitous – they have successfully colonized nearly
every known terrestrial, marine and freshwater habitat including polar and alpine
extremes, tropical lowlands and desert barrens and surface and mineral soils (Dunlop
1996, 2010). More than 55,000 species of mites have been described – a small

DOI 10.1007/978-981-10-1594-6_5
64 5 Classification of Subclass Acari
fraction (perhaps 5 %) of the total number of animal species estimated to be alive

today. A higher classification of subclass Acari into different superorders, orders
and suborders is discussed below as per Krantz and Walter (2009). A brief account
of economically important families is discussed in a separate chapter.
5.1 Superorder Parasitiformes
Alternative names for superorder Parasitiformes found in earlier literature are order
Parasitiformes (Krantz 1978), superorder Anactinotrichida (Van der Hammen 1972,
and Evans 1992) and cohort Parasitiformes (Wooley 1988).
Members of superorder Parasitiformes possess one to four pairs of dorsolateral
or ventrolateral stigmata posterior to coxae II; coxae of legs free, usually movable;
tarsi of legs II–IV with peripodomeric fissure associated with slit organs; and tarsi
of leg I with dense dorsal cluster of solenidiform setae subdistally. Many species are
parasitic, predaceous and living in the soil litter, rotting wood and house-dust; some
species also feed on fungus and pollen. There are over 15,000 described species of
Parasitiformes mites, and the total estimates found may be between 100,000 to
200,000.
Classification Superorder Parasitiformes is classified into four orders mainly

Opilioacarida, Holothyrida, Mesostigmata and Ixodida. Brief account of these
orders is given below (ER4, ER5 and ER6).
5.1.1 Order Opilioacarida
Alternative names for this are suborder Opilioacarida (Krantz 1978), order
Opilioacariformes (Johnston 1982; Walter and Proctor 1999), order Opilioacarida
(Wooley 1988; Van der Hammen 1989) and order Notostigmata (Evans 1992).
There is only one superfamily Opilioacaroidea in this order.
Palp tarsus with one or two terminal claws; adult opisthosoma with four pairs of
dorsolateral stigmata (without peritremes) posterior to level of coxae III; anus ter-
minal; trochanters III and IV divided into two podomeres. Females with well-
developed extrusible ovipositor terminating in two to three lobes. They are widely
considered as primitive as they retain six pairs of eyes and possess abdominal seg-
mentation. The anterior portion of the body bears two to three pairs of simple eyes,
and the posterior portion has many pores with about 12 secondarily developed seg-
ments; terminal apotele a pair of claws; rutella present; III–IV legs with subdivided
trochanter; tritosternum large and divided. They are large mites (1.5–2.5 mm) and
are rarely found under stones and other debris in semiarid habitats. These mites are
carnivorous/omnivorous and seem to be of no economic importance.
5.1 Superorder Parasitiformes 65
5.1.2 Order Holothyrida
Alternatively known as suborder Holothyrida (Krantz 1978; Walter and Proctor

1999), suborder Holothyrina (Johnston 1982) and order Holothyrida (Wooley 1988;
Van der Hammen 1982, 1989; Evans 1992).
Adult holothyrids are large (2–7 mm) with a heavily sclerotized body (Gr. Holo
meaning whole and thyr means shields) and may be having one pair or without
ocelli. Tectosternum is present, and hypostome is with corniculi. A gland opens on
the lateral dorsal shield posterior to the peritreme (Thonnes’ organ). There is only
one superfamily Holothyroidea. These mites live in mesic litter habitats on islands
in Caribbean Sea, in Indian Ocean, in Southern and Central America and in
Australasia and are found in leaf litter, in mosses and under stones in moist forests
from near sea level to about 2000 metres in elevation.
5.1.3 Order Mesostigmata
Alternatively these mites have been placed in earlier literature as suborder Gamasida
(Krantz 1978), suborder Mesostigmata (Johnston 1982), order Gamasida (Wooley
1988; and Van der Hammen 1989) and order Mesostigmata (Evans 1992). They are
commonly known as mesostigmatan or mesostigmatid mites. Representatives are
very small mites (0.2–4.5 mm long) with a pair of stigmatal openings above legs
III–IV and are usually associated with a distinct peritrematal groove; chelicerae are
usually chelate–dentate, but produced as elongate stylets in some parasites; anterior
margin of subcapitulum (hypostome) usually with a pair of horn-like or bidentate
corniculi (rarely membranous), never with a field of retrorse spines; base of subca-
pitulum with a median groove with rows of one to many denticles; stalk-like trito-
sternum with two to three processes present at base of median groove; coxae freely
articulating with body; without prodorsal trichobothria; males never with an
aedeagus, but often with chelicerae modified for sperm transfer (spermatodactyl,
spermatotreme); female sperm receiving structure primary or secondary. Mostly
they are free-living predators (family Phytoseiidae and Laelapidae) or may be para-
sitic on vertebrates and on social insects. Measure from 200 to 4500 μm. Many
members are phoretic on insects as deutonymphs; some mites feed on pollen, nectar
and fungi. Most mesostigmatid mites are fluid feeders.
Classification Order Mesostigmata is further divided into three suborders, namely,

Monogynaspida, Trigynaspida and Sejida.
5.1.3.1 Suborder Monogynaspida

They can be recognized by the single pair of spiracles positioned laterally on the
body. Adult females have a single genital shield covering the ovipore and bearing
zero to one (rarely two) pairs of genital setae. Genital shield is truncate or rounded
posteriorly. Capitulum exposed or hidden under vertex. Most Monogynaspida are
found as parasites of birds (including poultry birds), mammals (e.g. rat mites),
reptiles (e.g. snake mites) and bees. Many species are found on flowers, as predators
of stored products and also predaceous on soil mites and small collembolans. Some
mites of this group are vertebrate parasites (superfamily Dermanyssoidea). Several
species of superfamily Phytoseioidea are used as natural enemies of different phy-
tophagous mites. Superfamilies Rhodacaroidea, Eviphidoidea, Diarthrophalloidea
and Europodoidea also contain many representatives having great economic impor-
tance. Monogynaspides account for most of the 12,000 described species of
Mesostigmata.
5.1.3.2 Suborder Trigynaspida

Genital region in adult female possesses two to four shields or their remnants.
Separate podonotal and opisthonotal shields are found; and venter usually with
large ventrianal and genitoventral shields. Chelicerae chelate–dentate to edentate;
movable digit with one to three brush-like structures; spermatodactyl absent.
Superfamilies Cercomegistoidea, Antennophoroidea, Fedrizioidea,
Coelaenopsoidea, Megisthanoidea and Parantennuloidea are all covered under sub-
order Trigynaspida. Most members of the Trigynaspida are associated with insects
(especially beetles and ants), millipedes or centipedes. A few are found on reptiles
(snakes and lizards). Some families of this suborder are free-living mites mostly in
tropical to subtropical forest litter.
5.1.3.3 Suborder Sejida

Alternatively known as cohort Sejina (Krantz 1978). Female mites have a large
genital shield and large sac-like spermatheca. The body has numerous gland open-
ings and is often densely tuberculated. Adult female with dorsal podonotal and
pygidial shield that may bear horns. Peritremes and peritremal shields are usually
well developed. Males tend to have more consolidated body shields. Palp apotele is
two tined; chelicerae are chelate–dentate. Superfamilies Sejoidea and
Heterozerconoidea are covered under suborder Sejida. They are common litter
inhabitants in moist forests in the subtropics and tropics especially in association
with rotten wood. Some members are even predaceous on small arthropods and
nematodes.
5.1.4 Order Ixodida
Alternatively known as suborder Ixodida (Krantz 1978; Johnston 1982; Walter and
Proctor 1999) and order Ixodida (Wooley 1988; Van der Hammen 1982, 1989;
Evans 1992). Representatives are known as ticks. Ixodid in Greek means sticky, and
Ixodes in Latin means ‘hard ticks’. Subcapitulum with hypostome modified into a
piercing organ with retrorse teeth but without corniculi; tarsus of palp without a
tined apotelic claw; opisthosomatic stigmata without elongated peritremes; dorsum
of tarsus I with a sensory complex (Haller’s organ) consisting of a deep posterior pit
and a less well-defined anterior depression, both with sensory setae.
5.2 Superorder Acariformes 67
Ticks are actually a highly distinctive group of relatively large mites; body length
varies from 1 to 5 mm as unfed adults but up to 20 mm when fully engorged. There
are about 896 species of ticks described from the world. Order Ixodida comprise
only one superfamily Ixodoidea which is divided into three families, namely,
Ixodidae, Argasidae and Nuttalliellidae. Brief characteristics and important genera
found in each family are given in a separate chapter which includes important fami-
lies of subclass Acari (Baker and Murrell 2002; Black and Piesman 1994; Lindquist
1984).
Economic Importance of Ixodida Ticks are obligatory blood feeders on verte-

brate hosts and depend entirely on blood for food and survival. They parasitize
mammals, birds and reptiles. No tick species feed on plant juices or prey on other
arthropods. Their feeding results in blood loss and inflammation, irritation and
allergy of the skin. Tick feeding may also cause a toxic reaction in the host resulting
in ‘tick paralysis’. Ticks serve as vectors of a broad range of microorganisms, pro-
tozoa, rickettsiae and viruses. Ticks and tick-borne diseases affect human health,
poultry, veterinary animals and zoo animals and result in significant economic
losses. Diseases transmitted by ticks to livestock are an additional major constraint
to animal production particularly in subtropical areas of the world.
5.2 Superorder Acariformes
Acariformes are more diverse of the two superorders of subclass Acari. Acariformes
are alternatively known as order Acariformes (Krantz 1978), cohort Acariformes
(Wooley 1988) and superorder Actinotrichida (Evans 1992). Mites belonging to
superorder Acariformes are without visible stigmata posterior to coxae II but never
behind the legs; coxae of their legs are integrated with venter of podosoma. Short
peritremes are sometimes present on anterior plate or variously elaborated on the
cheliceral bases. Gnathosoma with ring-like band of cuticle surrounding cheliceral
bases exposed dorsally, fused into a stylophore (separate from subcapitulum) or
integrated with subcapitulum into a head-like capsule; palps are well developed
(five free segments) or variously reduced or apparently absent. Venter of gnatho-
soma of these mites is entire or provided with articulations, but never with a median
denticulate groove or with dense field of retrorse spines. One to two pairs of ocelli
may be present, but many taxa are blind. There are over 40,000 described species in
351 families (Baker et al. 1958; Krantz and Walter 2009; Dabert et al. 2010; Pepato
et al. 2010), and an estimated total of 440,000–929,000 species including unde-
scribed species may be found (ER 6).
Classification Superorder Acariformes is divided into two orders, namely,

Trombidiformes and Sarcoptiformes. Brief characteristics and further classifi-
cation of each order are given below.
5.2.1 Order Trombidiformes
Chelicerae are rarely chelate; fixed digit often regressed and movable digit usually
a hook-, needle- or stylet-like structure; cheliceral bases sometimes fused medially;
palpi are simple or modified into a thumb–claw process; subcapitulum without
rutella; ambulacra of at least legs II and III usually with two lateral claws and with
or rarely without a median empodium which may be pad-like or rayed and often
armed with tenent hairs. Tracheal system with one pair of stigmata opening between
bases of chelicerae, or on anterior prodorsum usually present and sometimes associ-
ated with peritremes on the cheliceral bases or on the anterior margin of prodorsum.
Order Trombidiformes is further divided into two suborders: Sphaerolichida and
Prostigmata (Cunliffe 1955; Dabert et al. 2010).
5.2.1.1 Suborder Sphaerolichida

Tracheal system absent; prodorsum with three or six pairs of setae including two
pairs of filamentous bothridial sensilla, sometimes in a common pit. Chelicerae che-
late and with elaborate dentition; coxal fields contiguous; naso nude. Suborder
Sphaerolichida includes superfamilies Lordalychoidea and Sphaerolichoidea.
5.2.1.2 Suborder Prostigmata

Alternatively known as suborder Actinedida (Krantz 1978) and order Actinedida
(Wooley 1988). Mites of the suborder Prostigmata have tracheal system with one
pair of stigmatal opening between bases of chelicerae or an anterior prodorsum usu-
ally present (secondarily absent in Eriophyoidea, Stigmaeidae and some
Dolichocyboidea); prodorsum usually with four or fewer pairs of setae (sometimes
including one to two pairs of bothridial sensilla); chelicerae rarely chelate (e.g.
Rhagidiidae and some Bdellidae); usually with fixed digit sheath-like or completely
regressed; coxal fields contiguous or II–III separated. Many fall within size limits of
300–500 μm, but some parasitic species of Heterostigmata may measure less than
100 μm in length. However, some members of family Trombidiidae can exceed
12 mm in length.
Suborder Prostigmata includes 36 superfamilies; Anystoidea, Arrenuroidea,
Bdelloidea, Cheyletoidea, Eriophyoidea, Erythraeoidea, Eupodoidea, Halacaroidea,
Hydrachnoidea, Hygrobatoidea, Pyemotoidea, Pygmephoroidea, Pterygosomatoidea,
Raphignathoidea, Scutacaroidea, Tetranychoidea, Trombiculoidea and
Trombidioidea are important superfamilies. Representatives include a variety of ter-
restrial, aquatic and marine predators and phytophagous and saprophagous
parasites.
5.2.2 Order Sarcoptiformes
Alternatively known as suborder Acaridida and Oribatida (Krantz 1978) and order
Astigmata and Oribatida (Evans 1992; Wooley 1988). In these mites, functional
chelicerae are present (often highly modified) and may be usually chelate–dentate;
palps present; stigmatal openings and peritremes absent, linear or elaborated on

cheliceral bases; prodorsal plate usually present or absent and often bearing a pair
of trichobothria. Subcapitulum usually with rutella or pseudorutella; empodium of
pretarsus often pad-like, or covered with processes, or produced as tenent hairs, and
rarely claw-like. Lateral claws usually present at least on legs II–III (leg I pretarsus
may be claw-like or absent, leg IV pretarsus sometimes absent); legs III–IV present
or absent. Opisthosomal glands absent; genital opening longitudinal, transverse or
absent; genital papillae present or absent; order Sarcoptiformes is divided into two
suborders, namely, Endeostigmata and Oribatida.
5.2.2.1 Suborder Endeostigmata

Small soft-bodied mites; leg tarsi with unpaired empodial claw. Opisthosoma with-
out lateral glands and with five to six pairs of prodorsal setae of which one to two
pairs usually are bothridial. Some mites are elongate to worm-like (often with annu-
lations), usually with reduced prodorsal setation and no bothridia. Bimichaeloidea,
Nematalycoidea, Oehserchestoidea, Terpnacaroidea and Alicorhagoidea are impor-
tant superfamilies.
5.2.2.2 Suborder Oribatida

Alternatively known as order Oribatida (Balogh 1992; Wooley 1988; Evans 1992).
Also commonly known as oribatid mites, beetle mites, armoured mites, astigmatans,
cryptostigmatans, moss mites, acarids, etc. Size ranges from 0.1 to 2.0 mm long.
Minute to large mites usually with a distinct prodorsal shield or fully sclerotized
prodorsum and sometimes with extensive idiosomatic sclerotization. Genital aper-
ture exposed or partially covered by paragenital flaps and may be inversely V- or
U- shaped with usually two pairs of genital papilla in the adult. Palpi with only two
segments, rarely three. Adult males with a sclerotized aedeagus and often with a
pair of copulatory adanal suckers. Larva usually with urstigma. Most oribatids are
fungivores or detritivores and mostly inhabit soil-litter system; only a few are minor
pests of plants. Some oribatids spread fungal spores and other microbial propagules.
About 6000 species described in 100 genera. Important superfamilies in suborder
Oribatida are Palaeacaroidea, Ctenacaroidea, Brachychthonioidea,
Hypochthonioidea, Parhypochthonioidea, Phthiracaroidea, Perlohmannioidea,
Oppioidea and Ceratozetoidea. The role of oribatid mites is discussed separately in
a chapter on soil mites as they seem to play good role in soil formation, as predators
of microarthropods and as intermediate hosts of cestodes which are disease trans-
mitting in animals. Superfamilies Histiostomatoidea, Hemisarcoptoidea,
Glycyphagoidea, Acaroidea, Analgoidea and Sarcoptoidea form part of Oribatida
(which includes cohort Astigmatina earlier known as Astigmata, Acaridiae or
Acaridida).
5.3 Conclusions
The Acari is tentatively assumed to be monophyletic because of the primitive shar-

ing of a variety of presumably apomorphic traits by its composite lineages, the
Parasitiformes and the Acariformes and because there is no convincing evidence as
yet that these two lineages (superorders) are more closely related to any other group
of Arachnida than they are to each other. Some more relevant references are reported
under ‘Further Reading’ of this chapter for better understanding the relationships
with different groups of Arachnida. At higher category, the level of cohort is often
placed between the levels of class and order. However, some authors have used
cohort between order and family level, but in modern classifications, suffixes for
supercohort and cohort names are standardized throughout the subclass (-ides and
-ina, respectively). Wherever used, the subcohort names are given the ending -ae.
Considering phylogenetic perspective, certain acarine groups have been overly
ranked in previous classifications, but in the classification scheme followed in the
present text (Lindquist et al. 2009), some modifications are made. For example, the
Astigmata previously ranked as separate Order is lowered to cohort rank
(Astigmatina) under oribatid supercohort Desmonomatides, while the Parasitengona
and Tarsonemina (or Heterostigmata) which were previously ranked as separate
suborders in some earlier classification schemes are lowered to cohort rank as sub-
sets of the supercohorts Anystides and Eleutherengonides, respectively, in the
scheme adopted in this text.
Further Reading
Baker SC, Murrell A (2002) Phylogeny, evolution and historical zoogeography of ticks. Exp Appl
Acarol 28:55–68
Baker EW, Wharton GW (1952) An introduction to acarology. Macmillan, New York
Baker EW, Caunix JH, Cunliffe F, Wooley TA, Yunker CE (1958) Guide to the families of mites.
Institute of Acarology, College Park
Balogh J (1992) The oribatid mites genera of the world, vol I. Hungarian Natural History Museum,
Budapest
Black WC, Piesman J (1994) Phylogeny of hard- and soft-ticks (Acari: Ixodida) based on mito-
chondrial 16S rDNA sequences. Proc Natl Acad Sci U S A 91:1034–1038
Cunliffe F (1955) A proposed classification of the trombidiforme mites. Proc Entomol Soc Wash
57:209–218
Dabert M, Witalinski W, Kazmierski A, Olszanowski Z, Dabert J (2010) Molecular phylogeny of
acariform mites (Acari: Arachnida): strong conflict. Mol Phylogenet Evol 56:222–241
Dunlop JA (1996) Evidence for a sister group relationship between Ricinulei and Trigonotarbida.
Bull Br Arachnol Soc 10:193–204
Dunlop JA (2010) Geological history and phylogeny of Chelicerata. Arthropod Struct Dev
39:124–142
Evans GO (1992) Principles of acarology. CAB International, Wallingford, 563pp
Johnston DE (1982) Acari. In: Parker SP (ed) Synopsis and classification of living organisms.
McGraw–Hill, New York, pp 111
Krantz GW (1978) Manual of acarology, 2nd edn. Oregon State University Bookstores, 509 pp
Further Reading 71
Krantz GW, Walter DE (2009) A manual of acarology, 3rd edn. Texas Tech University Press,
Lubbock, pp 97–103, 807 pp
Lindquist EE (1984) Current theories on the evolution of major groups of Acari and on the rela-
tionship with other groups of Arachnida, with consequent implications for their classification.
In: Griffiths DA, Bowmann EE (eds) Acarology VI, vol I. Wiley, New York, pp 28–62
Lindquist EE, Krantz GW, Walter DE (2009) Classification. In: Krantz GW, Walter DE (eds) A
manual of acarology, 3rd edn. Texas Tech University Press, Lubbock, pp 97–103, 807 pp
Pepato AR, da Rocha CEF, Dunlop JA (2010) Phylogenetic position of the acariform mites: sensi-
tivity to homology assessment under total evidence. BMC Evol Biol 2010(10):235.
doi:10.1186/1471-2148-10-235
Van der Hammen L (1972) A revised classification of the mites (Arachnida, Acarida) with diagno-
sis, a key, and notes on phylogeny. Zool Meded Leiden 47:273–292
Van der Hammen L (1989) An introduction to comparative arachnology. SPB Publishing, The
Hague
Walter DE, Proctor HC (1999) Mites, ecology, evolution and behaviour. CAB International,
Wallingford, 322pp
Wooley TA (1988) Acarology: mites and human welfare. Wiley, New York, 484pp
1. https://www.youtube.com/watch?v=3JNprSWv96w. Introduction to the Acari

2. https://www.youtube.com/watch?v=NOwnSW4v-Qs. Introduction to Acariform Morphology
3. http://itp.lucidcentral.org/id/mites/invasive_mite/Invasive_Mite_Identification/key/Whole_site/
Acari_Mites.htm. Mites or Acari
4. http://eol.org/pages/2586535/names?all=1. Acari: About Higher Groups & Families
5. http://www.discoverlife.org/20/q?search=Arachnida. Arachnida: Spider, Ticks & Mites
6. http://itp.lucidcentral.org/id/mites/invasive_mite/Invasive_Mite_Identification/key/0_Glossary/
Mite_Glossary.htm#Vertical-scapularsystem. Glossary of Acarine Terms
Important Acarine Families
6
Abstract
The subclass Acari (mites and ticks) belong to the assemblages of terrestrial
chelicerates in class Arachnida. They are quite small in size: the smallest are
among the plant mites (about 80 μm long), and the largest are among the preda-
tors (about 13 mm long). Most mite species are 400–800 μm long in adult stage.
Their varied modes of life influenced diversified morphology and functions.
Among arachnids, mites are the only phytophagous and parasitic. More than
55,000 species of different Acari have been described under 124 superfamilies
and 540 families. Among Acari, new species of order Ixodida are found only
infrequently; however, greatest increases in new species are among the Astigmata,
Mesostigmata, Oribatida and Prostigmata. Suborder Opilioacarida is compara-
tively less diverse and lesser known. Not more than 10 % of the total Acari are of
significant economic importance. Diagnostic characters of different families
with the help of figures are discussed in more than 100 families mentioning
important species under each family, but details of most important mites are
given under different relevant chapters. However, general account of ticks, tet-
ranychid mites and eriophyid mites is discussed in this chapter as these groups
needed more attention considering their utmost economic importance.
Keywords
Astigmata • Mesostigmata • Oribatida • Prostigmata • Trombidiformes
The Subclass Acari belong to the assemblages of terrestrial chelicerates in

Arachnida, whose representatives have secondarily adapted to an aquatic habitat.
Mites are quite small in size: the smallest are plant mites (about 80 μm long), and
the largest are predators (about 13 mm long). Most mite species are 400–800 μm
long in adult stage. Their varied modes of life influenced diversified morphology
and functions. Mites are the only phytophagous and parasitic arachnids. More than
55,000 species of different Acari in 540 families and 124 superfamilies have been

DOI 10.1007/978-981-10-1594-6_6
74 6 Important Acarine Families
described, and many new species are being described routinely from different geo-
graphic regions. Among Acari, new species of Ixodida are found only infrequently.
But greatest increases in new species are among the Astigmata, Mesostigmata,
Oribatida and Prostigmata. Suborders Opilioacarida and Holothyrida are compara-
tively less diverse and less known too. Not more than 10 % of the total Acari are of
significant economic importance (Krantz and Walter 2009; Walter and Proctor
2013). In this chapter brief characteristics of different families to which the impor-
tant species belong are discussed under different superfamilies. Economically
important species along with their hosts/habitats are given under each family, but a
detailed account of each most important species is given under relevant chapters.
However, ticks, tetranychid mites and eriophyid mites are being discussed in more
detail in this chapter as they needed more attention considering their utmost eco-
nomic importance.
6.1 Superorder Parasitiformes
6.1.1 Order Holothyrida
6.1.1.1 Superfamily Holothyroidea

Adults are fully sclerotized with dome-shaped holodorsal shield with short, broad
peritremes above legs. Tritosternum is flagelliform, if present; hypostome with cor-
niculi; chelicerae three segmented; palpi five segmented. They are relatively large
mites (>2 mm). The superfamily Holothyroidea consists of three families, namely,
Allothyridae, Holothyridae and Neothyridae. Adult holothyrids are large (2–7 mm),
heavily sclerotized and found in leaf litter, in mosses and under stones in moist
forests. Some species are scavengers and even predaceous in habit.
6.1.2 Order Opilioacarida
6.1.2.1 Superfamily Opilioacaroidea

They have leathery cuticle with purple or bluish markings but without distinct plates
and four pairs of dorsolateral spiracles with no peritremes; females have well-
developed extrusible ovipositor terminating in two to three lobes, paired rutella and
discoid or biramous With’s organ lateroventrally. Leg coxae free; trochanters III–IV
and tarsi I–IV are divided in tritonymphs and adults. Opilioacarids are covered in
only one family, Opilioacaridae. They are 1500–2300 μm in length, with long legs,
having leathery cuticle with distinctive blue or purple pigment patterns on the body
and legs. They live in caves, litter or under rocks in both semiarid and forest litter in
tropical and warm temperate regions. Opilioacarids have been reported to feed on
pollen, fungal spores and fragmented arthropod remains. Caribeacarus, Neoacarus
and Phalangiacarus are important genera.
6.1.3 Order Ixodida
6.1.3.1 Superfamily Ixodoidea

Idiosoma with a few or no sclerites but with a thick cuticle, with or without a
podonotal shield. Integument folded or striate, leathery or mammillated/tubercu-
lated. Festoons may or may not be present on the posteromarginal region.
Gnathosoma anterior or anteroventral in position. Hypostome modified as holdfast
organ; palpi simple, telescoped or normally extended with three to four segments
but without palpal apotele. Postlarval stages with dorsolateral stigmatic plates
between III and IV or behind coxae IV ((Sonenshine 1993; Sonenshine et al. 2002);
Sonenshine and Roe 2014). Haller’s organs on the distidorsal aspect of tarsus I; all
tarsi with claws. Divided into three families, namely, Ixodidae, Argasidae and
Nuttalliellidae (ER10 and ER11 and ER12).
Family Ixodidae (Fig. 6.1a, b) The members are commonly known as ‘hard ticks’.
Scutum (shield) present in female larvae, nymphs and adults is small, while in males
it is large. Mouthparts (capitulum) are visible dorsally in larvae, nymphs and adults.
Palps are club shaped; chelicerae have denticulate sheaths and no coxal glands. It is
generally accepted that the Ixodidae are divided into two lineages, the Prostriata
which consists of a single genus Ixodes and the Metastriata (all other genera). Brief
account of different genera belonging to hard ticks is given below (ER13).
Amblyomma Anal groove lies below anus, palpi longer than wide, ornate ticks, eyes
present, festoons present, rectangular basis capituli and oval idiosoma. A. variega-
tum Fabricius and A. hebraeum Koch cause major economic losses in cattle
industry.
Dermacentor Palpi wider than long, ornate, eyes present, festoons present and
basis capituli rectangular. D. marginatus Sulzer infests cattle and sheep and is
worldwide in distribution.
Haemaphysalis Anal groove below anus, eyes are absent, palpi wider than long, not
ornate and festoons present. H. punctata (Canestrini and Fanzago) is an important
pest of sheep.
Hyalomma Eyes present, palpi longer than wide, not ornate and festoons present.
Important species are H. aegyptiacum Linnaeus (tortoise ticks) and H. marginatum
Pomerantsev (Mediterranean ticks).
Ixodes Anal groove above anus, eyes absent, palpi longer than wide, not ornate and
no festoons. Ixodes ricinus Linnaeus (castor bean tick) is the most important and is
found worldwide. It parasitizes sheep, deer, horses and humans.
Rhipicephalus Anal groove below anus, palpi wider than long, eyes present, not
ornate, festoons present and basis capituli hexagonal. R. appendiculatus Neumann
(brown ear ticks) and R. sanguineus Latreille (brown dog tick) are important
species.
Family Argasidae (Fig. 6.1c, d) The members included in this family are com-
monly known as ‘soft ticks’. Scutum (shield) is absent; mouthparts (capitulum) are
not visible dorsally in nymphs and adults but are seen in larvae. Palps of soft ticks
look like legs. Chelicerae in these ticks have smooth sheaths; coxal organs are pres-
ent. Brief characteristics of important genera of soft ticks are given below.
Antricola Dorsal integument tuberculated, hypostome scoop-like and exclusively

associated with cave dwelling bats. The capitulum of these ticks cannot be seen in
dorsal view because it lies within a groove or depression called a camerostome. A.
marginatus (Banks) is a parasite of bats.
Fig. 6.1 Ixodidae: (a) Dorsum. (b) Venter, Argasidae: (c) Dorsum. (d) Venter
Argas Provided with an obvious marginal sutural line separating dorsal and ventral
body surfaces. A. persicus Oken is an economically important pest of fowls.
Ornithodoros Dorsal integument mammillated and hypostome usually with teeth

but never scoop-like. Mainly associated with bats. O. coriaceous Koch infests
mainly ears of cattle, but also attacks human beings.
Otobius Nymphs with dorsal integument beset with spines and adults with granular
integument and vestigial hypostomes. O megnini Duges, also known as spinose ear
tick, is an important species feeding within the ears of many mammals.
Family Nuttalliellidae (Fig. 6.2) This is a monotypic family characterized by fea-

tures, namely, intermediate to those of the two major tick families reported earlier.
Only one species, Nuttalliella namaqua Bedford, is reported from nests of swallows
from South Africa and Tanzania. These ticks have a leathery, mammillated or granu-
lar integument and lack a dorsal scutum. Hypostome is short with few denticles;
palpi consist of three segments.
Important Website on Ticks Considering the great economic importance of ticks

as most important parasites and disease transmitters in human beings, veterinary
animals, reptiles and birds, affecting their health badly, a website TicksBase (http://
www.icttd.nl/) has been initiated at Utrecht University, the Netherlands. This web-
site provides taxonomic catalogue of all known ticks (Acari: Ixodida) of the world
and is facilitated by the international consortium on ticks and tick-borne disease
Fig. 6.2 Nuttalliellidae: (a) Dorsum. (b) Venter

(ICTTD). Currently, it comprises 889 valid species (702 hard ticks, 186 soft ticks
and one nuttalliellid tick) including their synonyms, distribution data and common
names.
Tick Life Cycle Ticks have complex life cycles involving several blood meals with
the same or different animal hosts. Soft ticks are usually associated with nests, dens,
burrows or roosts of their animal hosts; they usually mate when they are not on a
host. Hard ticks are usually not associated so closely with their hosts but instead are
free-ranging and come in contact with animal hosts only for blood feeding. Hard
ticks usually mate when they are on a host. Because soft ticks take small blood
meals for a short feeding period, the female lays only a few hundred eggs during her
lifetime, with eggs laid at intervals. Hard ticks take large blood meals and lay 6000
or more eggs at one time (ER6 and ER10).
Ticks have four generalized life cycles which are known as one-host, two-host,
three-host and multi-host life cycle. Ticks are often referred to by the kind of life
cycle they have, for example, the American dog tick is a three-host tick. The three-
host life cycle has been adopted by most of the ixodid ticks, but two-host and one-
host life cycle is also found in some species of hard ticks. However, the multi-host
life cycle is characteristic of nearly all species of soft ticks. In multi-host life cycle,
many nymphal moults occur and these moults are called intermediate stages.
Individual nymphs feed and moult several times before moulting to the adult stage.
The adult ticks may feed many times. The female ticks may lay small batches of
eggs after each blood feeding.
Tick Removal To remove feeding ticks, dab them with alcohol. If feeding has just
started, and mouthparts are not cemented in, ticks sometimes pull their mouthparts
out. Try covering them with a dollop of menthol shaving cream or ‘Vicks VapoRub’.
It is generally best to take tweezers, grasp tick at the skin level and pull steadily until
the tick is removed. If mouthparts are left in the skin, it causes wound if left
untreated. So always treat with an antiseptic to prevent secondary infection (ER12).
Attachment and Feeding A fully engorged tick can deposit 100–18,000 eggs on
the ground. Normally thousands of tiny six-legged larvae hatch from the batch of
eggs and crawl randomly up grasses, weeds, twigs or low vegetation or walking
over the ground to await or search for a host in the surrounding area. These ticks,
called seed ticks, suck blood. The engorged seed ticks drop off and moult to eight-
legged nymph and again find a host. After several days of feeding, the engorged
nymph drops off its host and moults again. Females of hard ticks may feed from
several days to more than a week (ER6).
The tick slides its pair of slender teeth painlessly into the hosts’ skin and so the
feeding attachment begins. The central holdfast organ (hypostome), covered with
recurved teeth or ridges, is then inserted. Blood sucking begins; secretions from the
ticks’ salivary glands are then painlessly injected into the wound; these secretions
form around the holdfast organ and glue it in place. At this point, the tick cannot
voluntarily detach until its feeding ceases and the secretions stop. The strength of
the holdfast organ helps the ticks resist being scratched or pulled off. When feeding
is complete, the engorged female drops off the host, lays eggs and then dies.
Losses Caused Ticks are extremely dangerous external parasites of warm-blooded

animals, birds, amphibians, reptiles, veterinary animals and man. Their bites are not
only annoying and occasionally painful but may result in localized skin inflamma-
tion, secondary infection and possible introduction of disease-causing pathogens.
More than 65 disease-causing pathogens are known to be transmitted by ticks. Some
ticks are venomous and produce very painful bites, and some ticks even cause tick
paralysis and lameness in people and animals (ER13).
Both the hard and soft ticks are vectors of over 30 diseases to man, pets, cattle,
sheep, goats and livestock. Many domestic and wild animals are killed by tick-
borne diseases, such as fowl spirochaetosis, Texas tick fever, relapsing fever, piro-
plasmosis and anaplasmosis. Many other animals are so weakened that they
succumb to other diseases. Argas persicus causes nervousness, weight loss, lowered
egg production and even death in poultry birds due to blood loss. Many species of
hard ticks are significant human disease vectors and are responsible for spread of
Lyme disease and Rocky Mountain spotted fever (RMSF). Tick-borne encephalitis
(TBE) can lead to meningitis and, in serious cases, results in paralysis and deaths.
Management of Ticks No single strategy can control or manage ticks. So the fol-
lowing options need to be tried for their successful management:
• Ticks prefer still, damp, shady areas, so turn on the lights and dehumidifier and
fans.
• Look for ticks where pets (dogs, cats, etc.) are living and sleeping. Search areas
under the edge of rugs, under furniture, and in cracks around baseboards, win-
dows and door frames, or dog boxes.
• Since most ticks only slowly climb to the top of the grass, medium height vegeta-
tion and/or low bushes to await a passing host. So cultivate the land or at least
keep your grass and bush cut low especially around walks, paths, fences, sheds,
trees, shrubs, play areas and other potentially dangerous locations. Keeping the
grass short in tick-infested areas increases tick desiccation during hot weather.
• Protect yourself by wearing hats, long-sleeved pants and shorts, and tuck shirts
into light coloured pants and pants into socks or boots when going into tick-
infested or suspect areas.
• Repellents. Use any chemical recommended as repellent, permethrin
(Permanone) – a synthetic pyrethroid pesticide, azadirachtin or neem oil and/or
menthol or sulphur powder. Since these chemicals are also toxic, use them
cautiously.
• Monitoring and Surveying for Ticks. Examine the volunteer persons who wear
protective clothing and walk through tick-infested areas, for the attached ticks.
Also monitor by dragging white cloth over relatively open ground or ‘flagging’
low-level vegetation (i.e. moving the cloth in a waving motion over and through
vegetation) in densely bushy ground. Ticks that are lining up, waiting or questing
for passing hosts cling to the cloth and can be removed for identification and for
counting. Sampling should be done when vegetation is not wet and when ambi-
ent temperature is above 10 °C. Dry ice (solid form of carbon dioxide) can also
be used for most efficient collection of ticks. CO2 vaporizing from the dry ice
attracts ticks onto a white cloth panel on which they are easily visible and can be
removed periodically, or onto a platform lined with double-sided sticky tape on
which they get trapped.
• Dusting. Dust the open grass lawn when it is 6 cm or less with powdered sulphur,
lime, talcum powder or ground up chalk or calcium carbonated, or treat the area
as a last resort with an appropriate labelled pesticide poison, e.g. wettable pow-
der and/or microencapsulated formulations.
• Biological Control. Releases of chalcid parasitic wasp, Ixodiphagus hookeri (=
Hunterellus hookeri), have been found somewhat effective. Female wasp lays
eggs inside an engorged nymphs of the tick and parasitize them. Many species of
spiders and ants reported to eat ticks if they can find them. Several species of
fungi and nematodes are also reported feeding nymphal ticks while they are
moulting on the soil.
• Mechanical Controls. Increase areas of open lawn and sunlight penetration.
Keep lawn mower to a height of 7.5 cm or less as it lowers the humidity at ground
level. Get rid of bush, weeds, leaf litter, and other debris since these vegetations
can attract ticks and their host. Use wood chips or gravel to create a barrier
between wooded areas (where ticks are common) and your lawn.
• Habitat Management. Controlled burning of habitat may also help reduce tick
numbers. Promote areas that receive lots of direct sunlight as these are not con-
ducive to high tick populations. A 45 cm perimeter border of gravel or limestone
may prevent movement of ticks from grass areas into buildings. Cracks and crev-
ices around the base of buildings should be sealed with caulk. In the indoors,
routinely thorough vacuum cleaning of cracks, furniture, ceilings, floors/carpets,
wall behind pictures, bookshelves and drapes should be done. Spot treatment
with talcum powder or medicated body powder, and also crack and crevice treat-
ment should be done with residual dusts or silica gel or food grade diatomaceous
earth.
6.1.4 Order Mesostigmata (Krantz and Walter 2009)
6.1.4.1 Superfamily Ascoidea
Family Ameroseiidae (Fig. 6.3) Females measure 300–500 μm. The dorsal shield
is entire and usually bears 29 pairs of setae. The sternal shield of female bears two
to three pairs of setae; the genital shield is usually wedge shaped. Podal sperma-
Fig. 6.3 Ameroseiidae
theca is present. The corniculi are conspicuous and usually forked distally. They are
found in cracks, on the surface of baled hay, or feed on moulds which grow under
damp situations. Kleemania plumosa (Oudemans) is a cosmopolitan inhabitant of
stored food and in nests of social insects where it feeds on fungi.
Family Melicharidae (Fig. 6.4) Dorsal shield is entire; fixed cheliceral digit with
pilus dentilis modified into a hyaline flap, found as predators. Many species have
adapted to feed on fungi, pollen and nectar. Melichares agilis (Hering) is commonly
found in cultivated soil and in insect-infested stored grains and on dried fruits.
Family Ascidae (Fig. 6.5) Members are small to medium sized (300–500 μm) and
pale yellow to brown in colour. Chelicerae are dentate, serrate or rarely with
regressed fixed digit. Palp tarsus possesses two-tined apotele. The idiosoma has one
shield or is divided into two shields. Genital opening in males is found at the base
of tritosternum. The legs are long terminating in a pair of claws. They are preda-
ceous on soil mites and plant mites. Some species are very good biocontrol agents
of soil-inhabiting pests in greenhouses. Lasioseius althiasae (De Leon) is a good
predator of soil arthropods and nematodes. Blattiscocius tarsalis (Berlese) is
reported to feed on eggs of moth, Ephestia kuehniella, in flour mills.
6.1.4.2 Superfamily Phytoseioidea
Family Otopheidomenidae (Fig. 6.6) Otopheidomenidae is closely related to the

family Phytoseiidae. They have a reduced fixed digit of chelicerae, a terminally
located anus and a reduced number of setae and are found in the thoraco-abdominal
and wing axillary areas of noctuid and sphingid moths. Otopheidomenis spp. and
Fig. 6.4 Melicharidae
Fig. 6.5 Ascidae
Fig. 6.6 Otopheidomenidae
Noctuiseius spp. are found in tympanic recess of moths. Heavy infections of

Hemipteroseius indicus (Evans) sometimes may paralyze red cotton bugs.
Family Podocinidae (Fig. 6.7) Measures 430–600 μm. Leg I greatly elongated
with genual, tibial and tarsal segments attenuate, subequal; palp tarsal claw usually
three tined, with one to two apical, whip-like seta. They possess a phytoseiid-type
spermathecal system. They inhabit humus and litter habitats throughout the tropics.
Podocinum pacificum Berlese is reported from rodent nests, ant nests and grain field
debris. P. sagax (Berlese) is sometimes reported voraciously feeding on
collembolans.
Family Blattiscocidae (Fig. 6.8) Fixed cheliceral digit with setiform pilus dentilis;
movable cheliceral digit lacking a ventral mucro, peritrematic shield broadly fused
posteriorly to exopodal plate curving behind coxae IV; epigyneal shield usually
truncate posteriorly. Members have adopted to a broad spectrum of terrestrial, arbo-
real and sub-aquatic habitats. Blattisocius species are common predators of acarid
mites and insect eggs and larvae in stored food and in nests of small mammals and
birds. B. keegani Fox is reported feeding on bagworm pupae.
Family Phytoseiidae (Fig. 6.9) They are medium-sized (250–400 μm) mites with
relatively long legs. Many species are pale, yellowish, orange or dark brown.
Fig. 6.7 Podocinidae
Idiosoma is covered by a single shield (rarely two shields) with not more than 20
pairs of setae. Leg I terminates in well-developed ambulacra. The stigma opens
between legs III and IV. The male chelicerae bear a spermatodactyl, the shape of
which has diagnostic value.
Life History and Biology The life cycle consists of egg, larva, protonymph,
deutonymph and adult stages. Eggs generally require very high humidity for hatch-
ing. Phytoseiid mites develop faster than spider mites; they can reproduce through-
out the year in tropical and subtropical conditions and in greenhouses under
temperate areas. Under field conditions in temperate areas, phytoseiids overwinter
in protected habitats as fertilized females. Phytoseiid mites feed on a variety of food
and have developed different feeding habits. They are known to be common inhabit-
ants of leaf domatia. Neoseiulus, Amblyseius, Galendromus, and Phytoseiulus are
the well-known genera. Most phytoseiids feed on a variety of products and small
arthropods (including thrips, whiteflies, scale crawlers). Phytoseiid mites are mainly
predators of spider mites; some species have also been reported feeding on nema-
todes, fungus and pollen. Some species are reported useful as predators of thrips
particularly under greenhouse conditions. They are fast-running mites. Phytoseiids
are the best known predators among the Acari and are being commercially used for
Fig. 6.8 Blattiscocidae: (a) Dorsum. (b) Venter
biological control of spider mites and are sold by many insectaries particularly for
control of T. urticae in greenhouse crops (cucumbers, peppers, tomatoes, strawber-
ries and some ornamentals).
In general, phytoseiid mites have a high intrinsic rate of increase (rm). Species
like Phytoseiulus persimilis are usually specialized predators of Tetranychus spe-
cies. These predators have been most widely used in applied biological control of
spider mites. They have relatively low food requirements for development and
reproduction, and this quality accounts for their efficiency even at low prey
densities.
6.1.4.3 Superfamily Dermanyssoidea
Family Dermanyssidae (Fig. 6.10) Elongate second cheliceral segment of female

far exceeding basal cheliceral segment I length. In case of male chelicerae, second
segment of chelicerae is with normal length. Idiosoma is broadly rounded posteri-
orly. Dermanyssids are parasites of mammals and birds. Dermanyssus gallinae (De
Geer) and Ornithonyssus sylviarum (Canestrini and Fanzago) are important pests of
poultry.
Fig. 6.9 Phytoseiidae: (a)

Dorsum. (b) Venter
Family Laelapidae (Fig. 6.11) Members of this family have dorsal shield with
more than 23 pairs of setae; female genital shield is flask shaped. Stigma is located
between legs III and IV, without straight peritremes. Leg I is usually with ambula-
crum. Soilborne species, Androlaelaps and Stratiolaelaps, serve as natural control
of root worms. Another soilborne predator Hypoaspis aculeifer (Canestrini) is
reported feeding on plant nematodes and scarabaeid beetles. Tropilaelaps clareae
Delfinado and Baker is an important ectoparasite of honeybees.
Family Varroidae (Fig. 6.12a) Fixed digit of chelicerae absent; peritremes of

female short, looped medially or apically and confined to level of coxae III or III–
IV. Varroa destructor Anderson and Trueman is an important parasite of honeybees,
Apis mellifera, and is discussed in detail in a separate chapter concerning bee para-
sitic mites.
Halarachnidae (Fig. 6.12b) Epigyneal shield absent or rudimentary; sternal shield

usually present; and stigmata are located ventrally or are lateroventral in position.
They are parasites of respiratory tracts of terrestrial and marine mammals. Many
members of the family, particularly Pneumonyssus simicola Banks and P. caninum
Chandler and Ruhe, infest the sinuses and nasal passages of monkeys and dogs.
Fig. 6.10 Dermanyssidae: (a) Dorsum. (b) Venter
Fig. 6.11 Laelapidae: (a) Dorsum. (b) Venter

Fig. 6.12 (a) Varroidae

(b) Halarachnidae
Family Spinturnicidae (Fig. 6.13) Tritosternum absent or represented by tritoster-

nal base remnant; and epigyneal shield narrowed or otherwise reduced. Members
are parasites of bats’ wing membrane. Platyrrhinus and Sturnia are important
genera.
Family Entonyssidae (Fig. 6.14) Sternal and epigyneal shield well developed but
often weakly sclerotized; stigmata are lateral in position. Entonyssus asiaticus Fain
is a lung parasite of snakes.
Fig. 6.13 Spinturnicidae: (a) Venter. (b) Dorsum
Fig. 6.14 Entonyssidae:
(a) Venter. (b) Dorsum.
(c) Chelicerae
Fig. 6.15 Rhinonyssidae
Fig. 6.16 Macronyssidae: (a) Dorsum. (b) Venter

Family Rhinonyssidae (Fig. 6.15) Sternal shield reduced or absent but with distinct
sternal setae; epigyneal shield well developed or reduced; and stigmata are dorsal in
position. Members are parasites of respiratory tracts of birds. Ptilonyssus mimi George
and Sternostoma tracheacolum (Lawrence) are important nasal parasites of birds.
Family Macronyssidae (Fig. 6.16) Members have edentate but elongate chelic-
erae. Corniculi are membranous; palp trochanter often with a raised mediolateral
keel and with a large non-setigerous spur on leg coxae II; other coxae without spurs
but occasionally with small ridges. They are parasites of mammals, birds and rep-
tiles. Ornithonyssus bacoti (Hirst) and O. sylviarum (Canestrini and Fanzago) are
important parasites of birds and reptiles.
Family Hystrichonyssidae (Fig. 6.17) Second cheliceral segment normally devel-

oped, considerably shorter than greatly elongated basal cheliceral segment.
Hystrichonyssus turneri Yunker and Baker are reported as parasites of porcupines
and snakes.
6.1.4.4 Superfamily Parasitoidea
Family Parasitidae (Fig. 6.18) In case of males, chelicerae with a spermatotreme –

a sperm holding process that is coalesced distally with movable digit. Males usually
with highly developed, sexually dimorphic, spine-like setae, spurs and apophyses
on legs II. Many representatives are predaceous, free-living or associated with
insects. Parasitellus species are phoretic on bumblebees; and Parasitus spp. are
common phoretics on carabid and silphid beetles.
6.1.4.5 Superfamily Rhodacaroidea
Family Rhodacaridae (Fig. 6.19) Chelicera usually with arthrodial brush.

Palptarsal apotele three tined. They are free-living predators. Commonly found in
surface and subsurface soils. Rhodacarus and Rhodacarellus are important genera.
Family Digmasellidae (Fig. 6.20) Palptarsal apotele two tined. They are predators
and insect associates. Dendrolaelaps fallax (Leitner) are phoretic on sciarid flies.
6.1.4.6 Superfamily Veigaioidea
Family Veigaiidae (Fig. 6.21) Palptarsal apotele three or four tined and with adja-
cent hyaline scale-like process; hypostome with internal malae typically elaborated,
often bilobed and moustache-like. They are free-living predators and are found in
soil and decaying organic matter. Veigaia and Gamasolaelaps species are reported
from upper soil region as predators.
Fig. 6.17 Hystrichonyssidae
Fig. 6.18 Parasitidae
Fig. 6.19 Rhodacaridae:
(a) Dorsum. (b) Venter
Fig. 6.20 Digmasellidae: (a) Dorsum. (b) Venter

Fig. 6.21 Veigaiidae
6.1.4.7 Superfamily Uropodoidea
Family Uropodidae (Fig. 6.22) Female genital shield located between coxae II–
IV. Uropoda spp. are found in highly organic, insular deposits of manure and com-
post. Uroobovella marginata is a slow-moving mite found commonly in poultry and
cattle manure.
6.1.4.8 Superfamily Diarthrophalloidea
Family Diarthrophallidae (Fig. 6.23) Deutonymphs and adults with a single dor-
sal shield that is never flanked by marginal shields; dorsal shield with several pairs
of greatly elongated marginal setae; paranal setae also markedly elongated; palpal
claw absent. Troctognathus spp. and Brachytremella spp. are found on passalid
beetles.
Fig. 6.22 Uropodidae
Fig. 6.23 Diarthrophallidae
6.2 Superorder Acariformes
6.2.1 Order Trombidiformes
6.2.1.1 Suborder Prostigmata
6.2.1.1.1 Superfamily Tetranychoidea
Family Tetranychidae (Fig. 6.24) Members are commonly known as spider mites
as unlike spiders many species spin a web on plants’ leaves, feed and mate inside it.
Members of this family measure 200–900 μm. Morphology of the ambulacrum and
dorsal chaetotaxy are the main characteristics used for the classification of the
Tetranychidae. Leg chaetotaxy and its ontogeny offer additional potential and
appear to be of good phylogenetic value. The structures of the termination of the
‘peritremes’ are sometimes considered important for identification. The tarsi and
tibiae of the legs often carry some ‘tactile’ and fewer sensory ‘setae’. The shape of
the sensilla is often important for species identification. The dorsum of the tarsus I
of most tetranychids has two pairs of closely associated setae termed as ‘duplex
setae’. A single pair of duplex setae is present on the dorsum of tarsus II. Each pair
of the duplex setae consists of a long distal sensory member and short proximal
tactile member. The empodium and tenent hairs (homologous with true claws) vary
between different genera of tetranychid mites (Baker and Tuttle 1970, 1994; Bolland
et al. 1998; Helle and Sabelis 1985). Dorsal body integument pattern, leg length and
shape of the aedeagus many times prove useful in identification (ER3 and ER7).
Bryobiinae, Eurytetranychinae and Tetranychinae are the important subfamilies
under family Tetranychidae. A wide variety of dorsal body integument pattern
occurs in the Bryobiinae, whereas the dorsal body integument pattern of the
Fig. 6.24 Tetranychidae
web-spinning Tetranychinae tends to have more or less uniform striation (ER14). In

subfamily Bryobiinae, empodium is without tenent hairs; females with two or three
pairs of anal setae; and males with five pairs of genito-anal setae. But in subfamily
Tetranychinae, empodium is absent, or if present then without tenent hairs; females
with one or two pairs of anal setae and males with four pairs of genito-anal setae.
Subfamily Bryobiinae is divided into three tribes – Bryobiini (with true claws unci-
nate and empodium pad-like), Hystrichonychini (with true claws and empodium
pad-like) and Petrobiini (claws are pad-like and empodium uncinate). Bryobia,
Bryobiella, Hystrichonychus and Porcupinychus are important genera under sub-
family Bryobiini. Subfamily Tetranychinae is divided into three tribes,
Eurytetranychini (empodium claw-like when present; tarsus I with loosely associ-
ated setae or with one pair of duplex setae) and Tenuipalpodini (empodium claw-
like or split distally, tarsus I with two pairs of duplex setae and tarsus II with one
pair and opisthosoma with f1 in normal dorsal position or absent). Tenuipalpoides
and Tenuipalponychus (tribe Tenuipalpodini); Eurytetranychus, Eutetranychus and
Aponychus (all of tribe Eurytetranychini); and Schizotetranychus, Oligonychus,
Panonychus and Tetranychus (all of tribe Tetranychini) are important genera.
In Tetranychini, legs are never longer than the body, but long legs are found in
some Bryobiinae and in less derived Tetranychinae such as tribe Eurytetranychini.
In the other tetranychid families, a moderate leg length is the rule. In Bryobiinae,
aedeagus is of simple shape. But in Tetranychini, the aedeagus is more complicated
and is formed of a shaft and a knob.
Spinning Behaviour The use of silk and webbing is a major difference between
the Bryobiinae and the Tetranychinae. Little or no webbing is found in
Eurytetranychinae and Bryobiinae, but moderate webbing (Schizotetranychus spp.
and Oligonychus spp.) to complicated webbing (Eotetranychus spp. and Tetranychus
spp.) is found in Tetranychinae. But in many Eurytetranychinae (Panonychus spp.),
the females spin a network of guy ropes over each egg.
Habitat In case of Bryobiinae, most species live on the twigs or on the upper side
of leaves, but many Eurytetranychini mites live on the upper surface of leaves only.
In subfamily Tetranychini, members of most advanced genera (Eotetranychus and
Tetranychus) feed exclusively on the undersurface, but some members of
Oligonychus and Schizotetranychus live on both the upper and lower surfaces of the
leaves. In Panonychus species, immatures feed on lower surface, but adults live on
both the surfaces (Holtzer et al. 1988).
Host Range Many species live on only a few taxonomically close plant species; a
few are monophagous; others are extremely polyphagous. In general, members of
family Tetranychidae infest a wide range of plants belonging to different botanical
groups. Almost all species of Schizotetranychus live on monocotyledonous plants.
Tendency of polyphagy is observed in species living on herbaceous plants, being an
unstable habitat. Highest polyphagy is observed in Tetranychus species belonging
to subfamily Tetranychinae.
Population Dynamics The intrinsic rate of natural increase, rm, found from life
table data is helpful in determining the potential of increase in a population. Intrinsic
rate of increase is large when the duration of the generation is short and fertility
high; rm of spider mites is dependent on the breeding conditions like nature of the
host plant, surface available to each individual, temperature and humidity. The high-
est rm is found in different spider mites belonging to Tetranychini, particularly
among Tetranychus species which are most prolific. An increase in rm leads to an
increase in the number of annual generations and destruction of its host plants.
Moreover, these species have much more effective means of dispersal. On the con-
trary, most of the mites belonging to Bryobiinae have low rm, and these mites live
on perennial plants having a more stable habitat.
Diversity In family Tetranychidae, about 1272 species have been described in 95

genera. More than one hundred species can be considered as pests under different
agroclimatic conditions. About ten species are considered as major pests of different
crops, fruit trees, etc. The most well-known and widespread species of spider mite
is commonly known as two-spotted spider mite, Tetranychus urticae Koch.
Websites on Spider Mites Dr. Phillip in France has maintained a special website
named as Spider Mites Web (www.montpellier.inra.fr/CBGP/spinweb/). On this
website, taxonomic data relating to history of nomenclature, the geographical distri-
butions (with maps), and the host range of 1272 valid species recorded till November
2011 have been given. In addition about 1350 references on spider mites found from
the year 1758 till date are also listed on the website for use of the acarologists and
entomologists.
Injury by Spider Mites Spider mites injure leaves by piercing cells and sucking
their cell contents. The injury produces a white or yellow spots or ‘stipplings’ that
are heaviest on the undersurface of the leaves. The leaves loose photosynthetic sur-
face as mite feeding continues. Both photosynthetic rate and leaf water status
decline with increasing levels of spider mite injury. As colonies grow and feeding
intensifies, entire leaves progress from greyish green to yellow, brown or coppery
and eventually drop off. Damage begins in the lower canopy and progresses
upwards. Heavily stippled upper leaves may exhibit deformations reminiscent of
herbicide injury. If the infestation is unchecked by disease, predators or miticides,
spider mites may kill the entire plant. Many times, progression of symptoms from
silvering, yellowing, browning, lower leaf loss and death of plants may be mistaken
from the drought symptoms. Drought triggers spider mite outbreaks in many crops
by upsetting balance of spider mite by natural enemies, weather and host quality
(ER3 and ER5).
Spider mites cause widespread changes in their host at cytological and histologi-
cal levels. Leaf surfaces are damaged including punctured and collapsed epidermal
cells, disrupted cuticles and stomata. Mesophyll tissues including vascular bundle
sheaths, palisade and spongy parenchyma are also affected.
The effective micro-environment of a small arthropod such as a spider mite

seems very different from representative nearby conditions. Temperature and
humidity also have been shown to affect the behaviour of spider mites in several
ways that may have important consequences for their population dynamics. These
include modifications of dispersal and web building; air movement may also influ-
ence these factors. Prolonged direct contact with free water and from rain or simu-
lated rains also has detrimental effects on spider mites.
Factors Promoting Mite Outbreaks Generally spider mite outbreaks were

uncommon in situations where productivity was low, since the mite populations
remained low due to natural regulation by predators, disease and poor nutrition from
low-quality host plants. Biological disruption from use of fertilizers and pesticides
and good nutrition speeds up the developmental rate and increased fecundity and
longevity of mites. Under such modern techniques of crop production, spider mite
outbreaks are quite common.
Rainfall may slow down the infestations, but it will not destroy all the popula-
tions. Spraying is the best option to control spider mites. Pyrethroid insecticides
perform poorly against mites and could even aggravate the situation by ‘flaring’
their populations. Eggs are difficult to kill with insecticides/miticides, so reinfesta-
tion is likely to occur in 7–10 days after treatment as a result of egg hatching. The
reinfestation is frequently heavy because natural enemies have been reduced or
eliminated. A second application may be necessary to kill newly hatched mites
before they mature and deposit more eggs. Details about pesticides being used
against these pests are given in a separate chapter of this text.
Family Tenuipalpidae (Fig. 6.25) Members of this family are commonly known
as false spider mites, flat mites and red tea mites. They are variously coloured; have
different shapes like ovoid, round and elongate with flat venter or flat dorsum; and
measure about 0.3–0.4 mm. Cheliceral bases adnate, fused mesally into a stylo-
phore (sometimes withdrawn into the body). Chelicerae with fixed digit reduced
and movable digit whip-like. Simple palpus which lacks a thumb–claw complex on
the penultimate segment, palpal segmentation often reduced. Adults with three or
four pairs of legs; tarsal claws usually with tenent hairs on both claws and empo-
dium. Genital papillae absent; males with an intromittent organ (Smiley and Gerson
1995).
Importance They are generally reddish looking, slow moving and normally feed
near the midrib or veins on the underside of leaves. Tenuipalpid species Raoiella
indica Hirst, Brevipalpus californicus Banks, B. obovatus Donnadieu, B. phoenicis
Geijskes and Tenuipalpus punicae Pritchard and Baker are important pests. They
are commonly encountered on different plants and are of high quarantine impor-
tance. They feed on stems, fruits, flowers and leaves (often on the lower surface).
Some species like Larvacarus transitans (Ewing) produces twig galls on jujube
trees in India, and Obdulia tamaris Pritchard and Baker on Tamarix trees in Mideast.
Fig. 6.25 Tenuipalpidae: (a) Tenuipalpus. (b) Brevipalpus
Serious damage is caused to many crops (e.g. citrus, tea, grapes, fruit trees, orna-
mentals, orchids, pineapple and grasses). Several tenuipalpid species are confirmed
vectors of virus or virus-like diseases of plants. Brevipalpus phoenicis vectors cof-
fee ring spot and passion fruit green spot viruses in Brazil and Costa Rica. Some
Brevipalpus spp. have been identified as vectors of rhabdovirus that cause diseases
such as ‘citrus leprosis’. Mite-vectored diseases reduce the production as well as
life span of the plants.
Family Tuckerellidae (Fig. 6.26) Also known as ‘peacock mites’ because of the
elaborate ornamentations adorning the dorsal surface of their bodies. These are
brightly coloured having fan-like dorsal body setae and also possess five to seven
pairs of whip-like caudal setae. Empodium with tenent hairs; most dorsal setae
expanded leaf-like; posterior region with a row of flagelliform setae. Cheliceral
bases adnate, fused medially into a stylophore; chelicerae with fixed digit reduced
and movable digit whip-like. Tuckerella is the only genera of this family. They are
rarely found, but all reported species are phytophagous in nature and are found on
the stems of woody plants, usually in the cracks on small twigs. Some species are
associated with grasses. Tuckerella pavoniformis (Ewing) is reported commonly on
papaya.
Fig. 6.26 Tuckerellidae
Family Linotetranidae (Fig. 6.27) Members are commonly known as cryptic false
spider mites. They are rarely found in soil and on grasses in dryland habitats but not
as pests. They are usually colourless, slender and less than one-third of a millimetre
in length. Resemble other elongate tetranychids. Linotetranidae have also been
reported from moss and rotting trees. Empodium of these mites is provided with
tenent hairs; prodorsum with four pairs of setae. Cheliceral bases adnate, fused
mesally into a stylophore (sometimes withdrawn into the body); chelicerae with
fixed digit reduced and movable digit whip-like. Chambered peritremes present at
the dorsolateral surface of the prodorsum. Eyes are absent and palps five segmented,
usually with thumb–claw process. Linotetranus is an important genus, but none of
the species are pests of plants.
6.2.1.1.2 Superfamily Bdelloidea
Family Bdellidae (Fig. 6.28) Gnathosoma is cone shaped; mouthparts are snout-
like possessing long palpi which terminate with strong setae and lack the palpal
thumb–claw complex, measuring 450–3500 μm in length. They are reddish in
colour and fast-moving mites found on plants and in soil preying on phytophagous
mites and small insects like Collembola. Bdella depressa Ewing and B. longicornis
Linnaeus are found to feed on spider mites; another species B. lapidaria Womersley
is reported as an efficient predator of lucerne flea, Sminthurus viridis. B. distincta
Fig. 6.27 Linotetranidae
Fig. 6.28 Bdellidae
Fig. 6.29 Cunaxidae
Baker and Balock is reported feeding on eggs and crawlers of armoured scale
insects.
Family Cunaxidae (Fig. 6.29) They are small- to medium-sized (350–500 μm),
red- or brown-coloured and fairly fast-moving mites. The chelicerae are indepen-
dent and elongated; the movable digits are short and hooked, whereas the fixed
digits are reduced. The palps are raptorial, three to five segmented and often armed
with long spines on the internal margins. The stigmata are located at the bases of
chelicera without the peritremes. The prodorsum has two pairs of prominent
trichobothria. These are cosmopolitan and occur in soil, leaf litter, compost and
stored products. Members are general predators of small arthropods and nematodes.
Cunaxa capreolus Berlese and C. setirostris Hermann, Cunaxoides parvus Ewing
and C. olivieri Schruft are important predators (Meyer and Ryke 1959).
6.2.1.1.3 Superfamily Halacaroidea
Family Halacaridae (Fig. 6.30) Body rounded with enlarged palps; legs with long
setae; legs I and II anteriorly directed and legs III and IV posteriorly directed. They
are small-sized mites (220–800 μm in length). Halacaridae are the most diverse
family ecologically among all the Acari. Their biological diversity has enabled
them to invade habitats not only in salt water but also in freshwater, lakes, streams
and springs, as well as intertidal and cavernicolous habitats in fresh and saline water.
Fig. 6.30 Halacaridae
There are phytophagous forms and carnivorous forms (predators, parasites and even
scavengers). Many halacarid species are found as both external and internal para-
sites on animals and other hosts; some species are reported from intertidal or shal-
low subtidal habitats and also reported from ocean depths of 400–1400 m.
Copidognathus, Acarothrix, and Peregrinacarus are predominantly inhabitants of
diluted brackish water and freshwater (Cook 1967; Prasad and Cook 1972; Proctor
2004).
6.2.1.1.4 Superfamily Eupodoidea
Family Eupodidae (Fig. 6.31) Anal opening terminal; intercoxal region with 15 or
more pairs of setae; femora IV normal or strongly swollen, used in jumping. They
are commonly encountered in soil, litter, low-growing vegetation, lichens or moss
in temperate, boreal and polar latitudes. They are also reported from habitats that
range in elevations from sea level to high mountains (>2200 m). Some occur in
coastal intertidal zones and others have been reported from caves and under stones.
Cocceupodes species have swollen femora IV and are capable of rapid backward
movements when threatened. Leg I of eupodids is typically elongate (e.g. Eupodes
longisetosus Strandtman), but those of Linopodes species are unusually long and
may be encountered in mushroom houses and cause economic losses.
Family Rhagidiidae (Fig. 6.32) Palptibia with three setae; tibia I with two or more
solenidia; prodorsum with one pair of bothridia and three pairs of setae. Rhagidiids
Fig. 6.31 Eupodidae
Fig. 6.32 Rhagidiidae
Fig. 6.33 Penthalodidae
are unusual in having fields of modified recumbent dorsal solenidia on tarsi I–II that
are distinctive to be diagnostic at the generic level. These rhagidial organs are usu-
ally inserted in individual or compound grooves or troughs and typically are modi-
fied so that each solenidion takes the form of a pick hammer, with the offset stem
representing the shortened handle.
Members are soft-bodied and rapidly moving mites. They are generally opaque
white, but some are yellow, orange or red in colour. Larval and nymphal stages of
rhagidiids use spin from the podocephalic gland system to spin a web around them-
selves prior to moulting. Adults of Rhagidia longisensilla Shiba also spin silk for
catching prey for communal refuge and protection of the eggs. Reported from
alpine/high mountain regions, some are found at lower elevations. Rhagidiids com-
monly occur in the uppermost layers of soil and litter or within the soil column and
seem to prefer moist, dark cool niches. They are predaceous in nature. Rhagidia and
Coccorhagidia species are commonly reported.
Family Penthalodidae (Fig. 6.33) They are coloured and generally strongly scler-
otized; dorsum with ‘V’- or ‘Y’-shaped furrow and an anterior epirostrum project-
ing over the gnathosoma and ornamented with reticulate or punctate forms. Genus
Penthalodes is holarctic in distribution, while genus Stereotydeus is found world-
wide on moss and litter in temperate and subtropical habitats.
Family Penthaleidae (Fig. 6.34) Chelicerae separate; fixed digit regressed.

Movable digit sickle-like to prong-like – an adaptation for phytophagous behaviour.
Anal opening dorsal or dorsoterminal; femora IV similar to others. The stigmata are
Fig. 6.34 Penthaleidae
located between the bases of chelicerae. Red-legged earth mite, Halotydeus destruc-
tor (Tucker), is an important pest of many low-growing crops especially annual
broad-leaved plants and grasses. Blue oat mite, Penthaleus major (Duges), also
infests cereals, grasses, vegetables and ornamental plants in temperate and subtropi-
cal regions.
6.2.1.1.5 Superfamily Tydeoidea
Family Ereynetidae (Fig. 6.35) Presence of a peculiar sensory complex of

unknown function, the ‘Ereynetal organ’ on the tibia of leg I. Ereynetids seem to be
restricted to humid or wet habitats. They are rarely found in exposed foliar habitats
or arid regions. Some members are commonly found under bark and in dung, leaf
litter, moss and lichens, while some species are intimately associated with terrestrial
arthropods or gastropods. Riccardoella species are found on the surface and within
the mantle cavity of snails and slugs. Several Ereynetes spp. are associates of bark
beetles or phoretic associates on dung-inhabiting beetles and flies.
Family Tydeidae (Fig. 6.36) Weakly to heavily sclerotized striate or reticulate

mites. Tydeids are very small to small mites (150–500 μm) and have reduced fixed
cheliceral digits, fused cheliceral bases, and short movable cheliceral digits. Mites
are oval in shape and fast moving and can be found both singly and in congrega-
tions. The members live in a wide range of habitats and are predatory, fungivorous,
and scavenging species. Some species are considered pests or beneficial on agricul-
tural crops. Lorryia, Orthotydeus, Pronematus and Tydeus are important genera of
this family. Homeopronematus anconai (Baker) is reported as a good control of
eriophyid mite, Aculops lycopersici.
Fig. 6.35 Ereynetidae
Fig. 6.36 Tydeidae
Family Iolinidae (Fig. 6.37) Adults with only one pair of genital papillae, or papil-
lae absent; moveable digit of chelicerae stylet-like or whip-like. Palptarsus usually
with three or four elongated setae; males typically with an aedeagus. Proctotydeus
spp. are found associated on the tympana of noctuid moths. Pronematus ubiquitous
Fig. 6.37 Iolinidae
(McGregor) feeds on honey dew, fungi and dead insects. Homeopronematus anconi
(Baker) is reported as a good control of eriophyid mite, Aculops lycopersici (Wolff.)
6.2.1.1.6 Superfamily Eriophyoidea

Eriophyoidea, also known as the Tetrapodili or four-legged mites, are worldwide in
distribution, occurring on a variety of primarily perennial hosts. All post-embryonic
instars lack third and fourth pair of legs (ER16). All the members are obligatory
phytophagous mites. They are very small to moderate in length (100–150 μm in
general, but range is 80–300 μm) (Jeppson et al. 1975).
Family Eriophyidae (Fig. 6.38) They are tiny, microscopic mites, yellowish to
pinkish white to purplish in colour. The rostrum is usually small relative to the body,
with the short form oral stylets. The distal recurved portion of the stylet is usually
much shorter than the stylet base. They are found on a wide range of plants; and
many species are major pests causing economic losses to crops, fruit trees and low-
ers quality of fruits and flowers. Some species are used as biological agents to con-
trol weeds and invasive plant species.
Family Diptilomiopidae (Fig. 6.39) Also known as big-beaked mites. Gnathosoma

is large compared to the body; chelicerae are abruptly curved and are bent down
near the base. The big-beaked plum mite, Diptacus gigantorhynchus (Nalepa), dam-
ages apple and some other Prunus species.
Family Phytoptidae (Fig. 6.40) The members are commonly known as gall mites
or rust mites. The cheliceral stylets are slightly evenly curved form of small to mod-
erate size like in Eriophyidae.
Fig. 6.38 Eriophyidae
Fig. 6.39 Diptilomiopidae
Fig. 6.40 Phytoptidae
Body divisions, life cycle and damage by eriophyid mites

Body divisions: There are three main body divisions – the rostrum, the dorsal shield
or cephalothoracic shield and the abdomen.
Rostrum An important structure in the classification at the family level, it is of a

beak-like form and directed downwards. In the family Diptilomiopidae, the rostrum
is large in relation to the body and is apically attenuate. In the Phytoptidae and
Eriophyidae, on the other hand, the rostrum is usually small relative to the body.
Cephalothoracic Shield Also sometimes known as the propodosomal or dorsal

shield or simply shield, it is triangular or semicircular and bears a pattern of lines,
granules or broken lines which are distinctive for individual species. The pattern of
lines may vary slightly or differ in intensity in any given species, but it is an impor-
tant diagnostic feature. The cephalothoracic shield sometimes has a projection – the
anterior shield lobe, which is an important character at subfamily and genus level.
In a few instances, the lobe is thin- and flap-like. The lobes’ variation in shape and
prominence is of importance in species identification.
The Abdomen The principal part of the eriophyid body, usually tapers towards the
rear and is worm-like, is divided transversely into a number of superficial rings
which in bud and gall mites are regular dorsoventrally. Free-living forms are less
worm-like and may have flattened abdomens with longitudinal ridges or grooves or
lateral structures. They may also have the body clearly divided dorsoventrally into
tergites and sternites. Some species have a row of a white, waxy secretion on the
dorsal surface of the abdomen or may cover themselves with flocculent wax. The
body rings are frequently studded with elongate oval or rounded structures, called
micro-tubercles, which may completely cover the body, or occur on the sternites
only. The form, number and arrangement of micro-tubercles are useful diagnostic
characters. The abdomen bears a limited number of setae. The relative length of the
setae is a valuable specific and generic character.
Genitalia The transverse genital opening is located anteriorly, just behind the hind
coxae. The male genital opening is ‘V’ shaped, but in females genital opening is
covered by a semicircular hinged flap, the cover flap, which may have longitudinal
markings or scoring. The degree and nature of the cover flap markings are of signifi-
cance in the generic and specific placement of some species.
Legs All eriophyid mites have only two pairs of legs. The forelegs and hindlegs are
essentially similar. Leg setation is an important character for identification. The
claws or ‘featherclaws’ which arise from the distal part of the tarsus are important
structures. The rock-like claw may be straight or curved and often terminates as a
knob. The length of the claws is of use for species identification. The featherclaw is
usually of simple type which has a central stem from which arise branches or rays.
The number of rays is an important species character. The second type is divided
featherclaw or having some modification. Featherclaw structure is important in
genus and species placement.
Life Cycle Two main types of the life cycle occur in eriophyid mites. The first,
sometimes called a simple life cycle, involves only one type of female. The female
lays eggs, there are two nymphal stages and finally the adults develop. The nymphs
differ from the adults in being smaller, in lacking external genitalia and in the extent
of body micro-tuberculation. All known eriophyid mites are obligate plant parasites.
Males do not mate with females, but fertilization occurs from the contact of females
with sperm sacks laid down on the host by males.
The second type of life cycle sometimes called a complex life cycle involves two
types of females: the protogyne or primary form, which is associated with males
and structurally is very similar to them, and the secondary female or deutogyne,
which is not associated with males and is quite different in appearance to the proto-
gyne. Frequently protogynes occur on the leaves of the host plant, whereas deuto-
gynes usually appear in response to leaf hardening or the onset of winter and
hibernate in bark crevices or lateral buds, emerging in the following spring to lay
eggs on the new foliage. The eggs give rise to protogynes and males. Deutogynes
preserve the species during less favourable times of the year when food is not freely
available. Dispersal of these slow-moving mites is by wind, water, birds, insects or
humans. Commerce/trade of plants or their vegetative parts (in horticulture particu-
larly) is probably the major reason in introducing these mite species into distant
places previously un-infested areas. So any unknown plant abnormality should be
critically examined for the presence of these tiny eriophyid mites. Existence of deu-
togynes and protogynes of a species is helpful in identification at species and generic
level and is also helpful in pest management of these mites as deutogynes which
usually have different habits from the protogynes.
Types of Eriophyid Mites The eriophyid mites are often divided into several
groups.
• Russeting (Rust) Types. Feeding produces a brownish discolouration of the epi-

dermis. Heavy fine stippling introduces a blemishing or staining of leaf and fruit
surfaces. ‘Silvering and bronzing’ of mite-infested areas may also result.
• Gall Former Types. A number of certain eriophyid mites produce galls and other
plant abnormalities by injecting growth regulators into the host tissues. Modified
leaf provides shelter as well as food. There are many types of galls (bladder,
pouch, finger, nail head, etc.), but they all have an exit hole at the bottom (usually
undersurface of the leaf). Most galls are found on the leaves, but some are on
flowers, petioles, green stems and even roots.
• Erinea Forming Types. The feeding by certain eriophyid mites induces the pro-
duction of hairy patches, i.e. erineum (pl. erinea), on the surfaces of leaves and
petioles. Feeding within the thick hairs offers the mites both anchorage and pro-
tection. Some erinea on the undersurface are domed up and distort the upper leaf
surface.
Damage Caused by Eriophyid Mites (ER15)

• Leaf-Edge Rolling. In some cases, feeding by eriophyid feeding produces a roll-
ing of the leaf margin.
• Witches’ Broom. Some eriophyid mites cause the tip of branches to develop
broom-like bunches of twigs, commonly known as ‘witches’ broom’ which
result in shortening of internodal growth with resultant clustering of buds.
• Blistering and Tissue Breakdown. Eriophyids, with their small mouthparts, feed
on individual plant cells and do not usually deeply penetrate plant tissues. Mite
feeding results in loosening buds and raised blisters that dry out later in the sum-
mer leaving dead areas on the expanded leaf blades.
• Bud Destruction. Feeding by some bud-inhabiting mite species causes a blasting
of the buds. The feeding causes the interior embryonic parts of buds to swell
producing the symptom commonly known as ‘big bud’. The bud dies after the
mites leave the cavities. Partial damage results in distorted new growth that looks
like possible herbicide injury.
• Virus-Like Symptoms. The mite-injected growth regulators sometimes may cause
the entire physiology of the host plant to be disturbed (failure of fruit bud forma-
tion, loss of vigour, early death) or scattered symptoms (concentric ring blotch)
suggestive of a virus disease.
• Virus Transmission. Sometimes mites act as vectors of virus diseases and trans-
mit them by feeding. Some mosaic types of virus diseases are caused by mite-
infected viruses.
6.2.1.1.7 Superfamily Anystoidea
Family Anystidae (Fig. 6.41) Also known as whirligig mites. They are bright red,
long legged, large (500–1500 μm) and soft bodied, possessing few dorsal setae and
no prodorsal sensilla. They possess a palpal thumb–claw complex whose tarsus is
longer than claw. Most of the members are predaceous in habit. Anystis baccarum
Linnaeus is reported quite effective against mites infesting soybeans. A. agilis
Banks is also reported effective on grapevines against spider mites and also reported
feeding on larvae and adults of citrus thrips, Scirtothrips citri.
6.2.1.1.8 Superfamily Hydryphantoidea
Family Hydryphantidae (Fig. 6.42) Adults with two-segmented chelicerae; palpi

chelate with dorsotibial portion of tibia bearing a short, thick seta extending beyond
base of tarsus. Idiosoma usually with varying degrees of plate or platelet develop-
ment dorsally. Deutonymphs and adults are sluggish and are adapted for walking or
crawling in aquatic vegetation and detritus. Hydrophantes tenuabilis Marshall is
reported to feed on dragonfly eggs.
Family Thermacaridae (Fig. 6.43) Dorsal and ventral shields present; coxal plates
I with two rows of large setae extending posteriorly from capitular bay; palp femur
with two long medial setae. Reported only from hot springs. Thermacarus is an
important genus.
Fig. 6.41 Anystidae
Fig. 6.42 Hydryphantidae
6.2.1.1.9 Superfamily Eylaoidea
Family Eylaidae (Fig. 6.44) Prodorsal plate noticeably wider than long, con-
stricted near middle and bearing one pair of setae. Inhabits ponds and temporary
pools. Deutonymphs and adults feed on ostracods and cladoceran crustaceans and
are subelytral parasites of various aquatic Coleoptera and Hemiptera.
Fig. 6.43 Thermacaridae
Fig. 6.44 Eylaidae
Family Limnocharidae (Fig. 6.45) Prodorsal plate much longer than wide; geni-
tal acetabula scattered in ventral integument. Inhabits ponds and lakes. Limnocharis
is an important genera; adults feed on larval Chironomidae, and larvae parasitize
surface-dwelling aquatic Hemiptera (especially family Gerridae) and various
Odonata.
Fig. 6.45 Limnocaridae
6.2.1.1.10 Superfamily Hydrachnoidea
Family Hydrachnidae (Fig. 6.46) Adult gnathosoma typically with long, curved
pointed rostrum. Palpi chelate with dorsodistal tibial claw extending well beyond
base of tissues. Legs with five movable segments, with basifemur and telofemur
fused and with leg tarsi lacking paired claws. Dorsal plate bearing eight pairs of
setae. Larval Hydrachna are parasites of various aquatic Hemiptera and Coleoptera
in pools, ponds and lakes.
6.2.1.1.11 Superfamily Hygrobatoidea
Family Limnesidae (Fig. 6.47) Two pairs of ‘urstigmata’ borne between coxal
plates I and II; bearing seven to eight setae. Limnesia spp. and some other genera of
this family are found in streams, ponds and lakes.
Family Hygrobatidae (Fig. 6.48) Coxal plates I–III on each side all fused. Found
among the dominant water mites in flowing-water habitats. Larvae are parasites of
Chironomidae. Hygrobates nigromaculatus Lebert has been reported as a limiting
factor in limiting populations of chironomid larvae in lakes (Cook 1967; Prasad and
Cook 1972).
Family Unioncolidae (Fig. 6.49) Dorsal and coxal plates reticulate, coxal plates
bearing pointed projections posteriorly. Larvae are parasites of Chironomidae or
Trichoptera. Free-living species of Unioncola are essentially planktonic; many oth-
ers have obligate relationships with molluscs.
Family Aturidae (Fig. 6.50) Coxal plate III with pointed or lobed projections
posteriorly and bearing two pairs of setae. They are well-sclerotized and typically
Fig. 6.46 Hydrachnidae:
Fig. 6.47 Limnesidae
dorsoventrally flattened mites. Adults of many genera are strongly dimorphic. This
is a species-rich family of water mites. Larvae are parasites of chironomids.
Family Pionidae (Fig. 6.51) Genital acetabula numerous (more than 10 pairs),
genus of leg IV either concave on one side and with numbers of peg-like setae or
with median surface of palptibia bearing a peg-like setae at distal end, and coxal
Fig. 6.48 Hygrobatidae
Fig. 6.49 Unioncolidae
plates lacking well-developed projections associated with insertions of leg

IV. Members are found in springs, streams and standing water habitats. Piona con-
tains important species of water mites. Adults of many genera exhibit marked sex-
ual dimorphism.
Fig. 6.50 Aturidae
Fig. 6.51 Pionidae
6.2.1.1.12 Superfamily Arrenurroidea
Family Neoacaridae (Fig. 6.52) Well-sclerotized dorsoventrally flattened mites;

dorsal and ventral shields present; ventral shield suture lines between coxal plates
III and IV extending posteromedially to genital field region and well separated from
each other medially. Genital acetabula in single rows on each side. Neoacarus is an
important genera; larvae are known as parasites of Chironomidae.
Family Arrenuridae (Fig. 6.53) Adults with dorsal and ventral shields present; palpi
truncate; genital acetabula numerous and lying on acetabular plates extending later-
ally from gonopore. In Arrenurus genus most importantly, its members are found in
virtually all types of freshwater habitats. Many species feed on ostracods. Larvae of
Arrenurus are parasites of various nematocerous Diptera or Odonata (Cook 1967).
6.2.1.1.13 Superfamily Erythraeoidea
Family Erythraeidae (Fig. 6.54) Members are large, reddish mites with two pairs
of prodorsal sensilla; many dorsal setae and a thumb–claw process with long straight
chelae. Larvae usually parasitize other arthropods, whereas the nymphs and adults
are predators of bugs and other small insects. Blaustium putmani Smiley is reported
to feed on eggs and mobile stages of European red mite, Panonychus ulmi (Koch).
Family Smaridiidae (Fig. 6.55) Gnathosoma is entirely retractable into idiosoma;

idiosomal setae are usually setiform. Deutonymphs and adults are m oderately large
(1000–2000 μm). Active predators of small arthropods are primarily found in grass-
lands and litter habitats. Larvae of Smaris prominens (Banks) are reported as para-
Fig. 6.52 Neoacaridae
Fig. 6.53 Arrenuridae
Fig. 6.54 Erythraeidae
Fig. 6.55 Smaridiidae
sites of psocids; larvae of some other species are found on many other insects
especially scale insects.
6.2.1.1.14 Superfamily Trombiculoidea
Family Trombiculidae (Fig. 6.56) Commonly known as harvest mites or ‘chigger

mites’; larvae (170 μm) are commonly called ‘chiggers’, which are normally light
red and covered in hairs. There is a marked constriction in the front part of the body
in case of nymphal and adult stages. Chiggers attach to the host, pierce the skin and
inject enzymes into the bite wound that digest cellular contents. They do not burrow
into the skin or suck blood, but feed on skin causing itching.
They are commonly found on grasses and weeds and get transferred to their
hosts when they approach for grazing. They prefer damp areas in woodlands, grass-
lands, golf courses, parks and bushes along lakes and streams. Their feeding causes
intense ‘irritation’ or a wheal, usually with severe itching and dermatitis.
Leptotrombidium deliense Domrow is considered a dangerous pest and often may
carry tiny bacterium, Orientia tsutsugamushi, that causes scrub typhus or tsutsuga-
mushi. Trombicula alfreddugesi (Oudemans) and T. autumnalis (Shaw) are the most
prevalent species.
6.2.1.1.15 Superfamily Trombidioidea
Family Trombidiidae (Fig. 6.57) Idionotum with unpaired prodorsal plate;

palptibia usually lacking large spinose setae adjacent to claw and either lacking any
spinose setae that are not arranged in rows, when bearing large spinose setae
Fig. 6.56 Trombiculidae
Fig. 6.57 Trombidiidae
arranged in rows and then anterior edge of prodorsal plate concave. Measure 500–
4000 μm. Anal gnathosoma not retractable into idiosoma. They are active predators
of small invertebrates in habitats that range from s ub-aquatic to deserts. Trombidium
spp. parasitizes many species of insects and arachnids. Adults are usually found in
moist litter, or in moss in and around bark of tree trunk.
Fig. 6.58 Eutrombidiidae
Family Eutrombidiidae (Fig. 6.58) Idionotum bearing large plate posteriorly.

Deutonymphs and adults are large mites (500–4000 μm) and found in wide variety
of edaphic habitats worldwide. Postlarval instars of Eutrombidium locustarum
(Walsh) are predators of grasshopper eggs. Larvae of various Eutrombidium spp. are
parasites of various Orthoptera.
6.2.1.1.16 Superfamily Myobioidea
Family Myobiidae (Fig. 6.59) Chelicerae and subcapitulum fused into a gnathoso-
matic capsule that is retractable within the idiosoma; leg I modified for clasping
mammalian hosts. Ectoparasites in the fur of bats, rodents and various insectivores.
Myobia musculi (Schrank) attacks laboratory mice, and Radfordia ensifera (Poppe)
attacks laboratory rats and causes dermatitis in rodents.
6.2.1.1.17 Superfamily Pterygosomatoidea
Family Pterygosomatidae (Fig. 6.60) Chelicerae independently moveable, elon-

gate and with swollen bases and hook-like movable digit with one or more teeth;
empodia absent; peritremes emergent. Most species are reported from lizards and
tortoises. They are bright red mites found attached under scales or between toes,
feed on body fluids of their hosts and result in benign to severe pathological disor-
ders such as anaemia and intense skin irritation. Pimeliaphilus spp. are reported
parasitic on lizards and triatomine bugs.
Fig. 6.59 Myobiidae
Fig. 6.60 Pterygosomatidae
Fig. 6.61 Caligonellidae
6.2.1.1.18 Superfamily Raphignathoidea
Family Caligonellidae (Fig. 6.61) Measures about 250–550 μm. Basal segments
of stylate chelicerae fused with each other in midline to form a conical stylophore,
which bears on its dorsal surface a pair of sinuous peritremes. Palp five segmented;
palptibial claw single but well developed. Body without striations and shields,
found under the bark, mostly associated with bark beetles and from grain storages.
They are free-living predatory mites.
Family Stigmaeidae (Fig. 6.62) Body short (200–500 μm in length), orange or red
and ovoid or round in shape. Thumb–claw process present; chelicerae with stylet-
like movable digit. Dorsal shields frequently present; ventral shields absent. Coxae
I and II distinctly separated from coxae III. Genital and anal openings contiguous.
Stigmaeid mites are commonly found on leaves of plants and feed on different
stages of pest mites. Agistemus and Zetzellia are important genera, whose members
have been reported feeding on eriophyid mites, spider mites, false spider mites and
even on some soft-bodied insects (Meyer 1969).
Family Camerobiidae (Fig. 6.63) Body length ranges from 250 to 300 μm. Leg
tibia about three times longer than respective tarsi; oval to subcircular mites with
very long legs. Dorsal idiosomal setae in marginal area not whip-like. Idiosoma
nearly round in dorsoventral view. Gnathosoma often covered by prodorsum; che-
licerae fused to form a stumpy stylophore. Camerobia and Neophyllobius are
important genera of this family.
Fig. 6.62 Stigmaeidae
Fig. 6.63 Camerobiidae
Family Cryptognathidae (Fig. 6.64) Measures 300–400 μm in length. Fully

armoured mites with holodorsal shield produced anteriorly into a hyaline or orna-
mented collar into which the narrow elongate gnathosoma can be retracted, only the
bottom of chelicerae fused; dorsal and ventral idiosoma covered by a single shield.
Idiosoma oval in dorsoventral view, strongly sclerotized. Found in leaf litter, tree
bark, moss and lichens. Cryptognathous and Favognathous are important genera.
Fig. 6.64 Cryptognathidae
Fig. 6.65 Raphignathidae
Family Raphignathidae (Fig. 6.65) They are small red mites (300–500 μm);
round in shape without any suture between propodosoma and hysterosoma.
Propodosoma without any sensory setae; peritremes situated between chelicerae
and prodorsum. Tarsus with two claws and a rayed pulvillus. Tarsi I and II with a
single sensory organ. No genital suckers. May be found on low-growing herbs and
shrubs beneath tree bark, house-dust or in stored grains.
Family Eupalopsellidae (Fig. 6.66) Length varies from 250 to 400 μm. They have
four pairs of prodorsal setae, the empodia have one (rarely two) pair of tenent hairs
and palp tarsi are elongate. They are common in very dry regions. Saniosulus spp.,
Eupalopsis spp. and Eupalopsellus spp. are reported feeding on scale insects.
6.2.1.1.19 Superfamily Cheyletoidea
Family Cheyletidae (Fig. 6.67) Body oval to round (200–1600 μm in length);

palptibial claw present and well developed. Free-living predators, associated with
insects. In some studies, Cheyletus eruditus (Schrank) is considered effective in
controlling stored grain mites, Acarus siro Linnaeus. Some cheyletid mites are
found on plants feeding on crop pests, but some others are found in fur of
mammals.
Family Demodicidae (Fig. 6.68) Body elongated worm-like with numerous

annuli; legs strongly telescoped, highly reduced or modified into hooks. Measure
150–400 μm in length and are semi-transparent. Body is covered with scales for
anchoring itself to the hair follicles; possess pin-like mouthparts for eating skin
cells. Demodex folliculorum Simon is found commonly feeding in the eyelids,
cheeks and eye lashes of human beings. D. canis Leydig is found on domestic dogs
and cats and can occasionally infest humans.
Fig. 6.66 Eupalopsellidae
Fig. 6.67 Cheyletidae
Fig. 6.68 Demodicidae:
Family Cloacaridae (Fig. 6.69) Well-sclerotized mites without opisthosomatic

striae; gnathosoma reduced to a single segmented palpi, each with a fang-like stylet
apically. Legs without distinct claws. They are found as internal parasites in the tis-
sues of reptiles and birds. Pneumocephalus bubonic Fain and Smiley is reported
infesting connective tissues of lungs and bronchi of owls.
Family Psorergatidae (Fig. 6.70) All legs ending in paired lateral claws and a pad-
like or bilobed empodia; femora I–IV each with a strong ventral spur. Members are
parasites on the skin of mammals. Psorergates, Psorergatoides and Psorobia are
important genera and their infestations cause dermatitis, mange and follicular
infections.
Family Harpirhynchidae (Fig. 6.71) Legs I–II with empodial tenent hairs and
with or without lateral claws, legs III–IV without lateral claws or leg IV absent;
femora I–IV without strong ventral spurs. Infest the skin/feathers of birds found
under scales of reptiles. Ophioptera is an important genera.
6.2.1.1.20 Superfamily Pygmephoroidea
Family Siteroptidae (Fig. 6.72) Small mites (200–300 μm); chelicerae and subca-
pitulum are fused to the gnathosomal capsule, which has a pair of small cheliceral
stylets and reduced palps. The prodorsal sclerite of the female does not cover the
Fig. 6.69 Cloacaridae
Fig. 6.70 Psorergatidae: (a) Dorsum. (b) Venter
Fig. 6.71 Harpirhynchidae
Fig. 6.72 Siteroptidae: (a)

Dorsum. (b) Venter
entire propodosoma. The bases of legs II are twice as widely apart as the bases of
legs III–IV. Siteroptes spp. are pests of grasses and cereals and are responsible for
transmissions of pathogenic fungi. Siteroptes avenae is of economic importance
under greenhouse conditions. Pediculaster flechtmanni infests button mushrooms
and results in qualitative losses.
Family Pygmephoridae (Fig. 6.73) Gnathosoma of male longer than wide; tro-
chanter IV of female quadrangular. Prodorsum of females with stigmata round or
oval. Found associated with fungi, insects and animal nests. Pygmephorus sellnicki
Krczal has commonly been reported on cultivated mushrooms.
6.2.1.1.21 Superfamily Scutacaroidea
Family Scutacaridae (Fig. 6.74) Broad, tortoise-shaped bodies (120–370 μm in

length). Female with hood-like podonotal shield covering propodosoma and gna-
thosoma; margin of podonotal shield is striated. Scutacarus, Nasutiscutacarus and
Parascutacarus are important genera found commonly associated on halictid and
bumblebees, carabids and ants (Mahunka 1965).
Family Microdispidae (Fig. 6.75) They are small (80–315 μm) and possess greatly
enlarged and/or ornate dorsal setae. Idiosoma oval; distance between insertions of
legs II and III at least twice that between legs III and IV. Mostly free-living or asso-
ciated with insects and animal nests. Microdispus lambi (Krczal) is an important
Fig. 6.73 Pygmephoridae
Fig. 6.74 Scutacaridae
Fig. 6.75 Microdispidae: (a) Dorsum. (b) Venter
species found in soil, forest litter, decaying vegetation, rotting wood, moss beds and
other similar habitats.
6.2.1.1.22 Superfamily Pyemotoidea
Family Pyemotidae (Fig. 6.76) Female coxisternal plates II–IV separated from
each other medially by soft cuticle that bears a separate triangular sternal plate.
Male legs IV differentiated from legs II and III in being somewhat stouter and end-
ing with a single sessile claw. The chelicerae are minute and needle-like; the perit-
remes are barely absent and are dorsal in position directed anteriorly on the shoulders
of the propodosoma. Pyemotids reported as ectoparasites of insects and other
arthropods. Pyemotes anobii Krczal paralyses larvae and pupae of bees. P. herfsi
(Oudemans) is reported feeding on gall-making midges on oak leaves. P. tritici (La
Greze-Fossat and Montagne), straw itch mite, is associated with dermatitis in horses
and man and P. scolyti (Oudemans) with beetles.
Family Acarophenacidae (Fig. 6.77) Whitish-coloured adults measure about 140

μm in length; gnathosoma capsule, palps not distinct, chelicerae stylet-like. Body
covered by a series of sclerotized plates; prodorsum with anterior stigmatal
Fig. 6.76 Pyemotidae: (a)

Dorsum. (b) Venter
Fig. 6.77 Acarophenacidae
o penings; sensillum absent. Legs with strong claw. Acarophenax lacunatus Cross
and Krantz is an egg parasite of stored grain pest, Rhyzopertha dominica, on wheat.
Caraboacaridae (Fig. 6.78) Female with gnathosomatic capsule large, clearly

wider than long; female leg I lacking claws, smaller than legs II–III. Leg IV with
tibia and tarsus at least partially fused and with reduced claw. Mites are associated
with carabid beetles. Caraboacarus is an important genera.
6.2.1.1.23 Superfamily Tarsonemoidea
Family Tarsonemidae (Fig. 6.79) Females with four pairs of legs; legs IV three
segmented, usually much more slender than legs II–III. Males always with four
pairs of legs; legs IV inserted ventrally; male genital capsule caudal in position.
Members of Steneotarsonemus, Polyphagotarsonemus, Phytonemus, Florida and
Tarsonemus (Lindquist 1969) are known to feed on higher plants, while most tarso-
nemid species feed on the mycelia of fungi and alga. Polyphagotarsonemus latus
(Koch) and S. pallidus Banks are pests of agriculture. Acarapis woodi (Rennie) is an
endoparasite of bees (ER17).
Family Podapolipidae (Fig. 6.80) Females typically with one to three pairs of
legs; leg IV if present is five segmented and more slender than legs II and III. Male
genital capsule caudal or dorsal in position. Several species belonging to genera
Locustacarus and Podapolipoides are parasitic on insects and sometimes prove
detrimental.
Fig. 6.78 Caraboacaridae
Fig. 6.79 Tarsonemidae: (a) Dorsum. (b) Venter
Fig. 6.80 Podapolipidae
6.2.1.2 S
uborder Oribatida (Including Cohort Astigmatina)
(Balogh and Balogh 1992)
6.2.1.2.1 Superfamily Histiostomatoidea
Family Histiostomatidae (=Anoetidae) (Fig. 6.81) Chelicerae laterally flattened;

fixed digit typically comb-like with numerous finely spaced teeth; movable digit short.
Palpi short, simple and often with a membranous complex. Males are more heavily
sclerotized than females. Genital papilla ring-like, usually large, not using from a
common furrow and not directly associated with the genital opening; body relatively
larger and not dorsoventrally flattened. Histiostoma spp. are typical inhabitants of
dungs of vertebrates, and deutonymphs often specialize for phoresy on staphylinid
beetles. Histiostoma murchiei Hughes and Jackson is reported as a predator of earth-
worm egg cocoons. Anoetus spp. live in mutualistic association with halictid bees.
6.2.1.2.2 Superfamily Canestrinioidea
Family Canestriniidae (Fig. 6.82) Idiosoma with cuticle smooth, striated, reticu-
late, scaly or sometimes well sclerotized. Gnathosoma normally developed; palp
and chelicerae occasionally elongated. Females with genital valves fused to body
anteriorly, forming V-shaped oviparous. Members are moderate to large in size and
are very weakly sclerotized. They are external associates or parasites commonly
associated with many beetle families Carabidae, Scarabaeidae, Lucanidae,
Passalidae, Cerambycidae and Chrysomalidae. Passalophagus georglei is reported
associated with passalid beetles.
Fig. 6.81 Histiostomatidae
Fig. 6.82 Canestriniidae
6.2.1.2.3 Superfamily Hemisarcoptoidea
Family Hemisarcoptidae (Fig. 6.83) Adults with gnathosoma normally formed;

chelicerae chelate-dentate. Idiosoma with cuticle smooth, mammilate, microtri-
chous or occasionally strongly sclerotized female with oviparous in form of inverted
‘V’. Nanacarus minutus (Oudemans) is a common inhabitant of house-dust and
stored food products. Hemisarcoptid mites are common inhabitants of various arbo-
real and aquatic habitats. Hemisarcoptes spp. are obligate parasites of diaspidid
scale insects, and some species have been employed in biological control of scale
insects.
Family Chaetodactylidae (Fig. 6.84) Members have soft globose bodies with a
distinctly mammilate cuticle, lack sejugal furrow and have greatly enlarged empo-
dial claws that allow for movement on host pollen/nectar stores. Supracoxal gland
opening on an elongate, oval sclerotized region restricted to dorsal above leg I;
female pretarsus with paired elongate condylophores. Males with condylophores
fused along their length and emerging ventrobasally as a hook-like structure in the
base of the pretarsus. They are cosmopolitan in the distribution and are commonly
found in bee nests. Sennertia cerambycina Oudemans is found in nests of Xylocopa
and Ceratina bees. It is a kleptoparasite, destroying host eggs or larvae and feeding
on pollen and nectar stores.
Family Winterschmidtidae (=Saproglyphidae) (Fig. 6.85). Empodial claws pres-

ent; condylophorus usually fused into a V-shaped sclerite or a more elongate sclerite
Fig. 6.83 Hemisarcoptidae
Fig. 6.84 Chaetodactylidae
Fig. 6.85 Winterschmidtiidae
in base of ambulacral stalk. They are found in a wide variety of habitats. Saproglyphus
spp. are fungivorous in decaying materials, bark crevices and woody fungi.
Winterschmidtia spp. are found as predators of the eggs of bark beetles. It is
Procalvolia zacheri and Acalvolia squamata (Oudemans) sometimes infest stored
food products.
Family Carpoglyphidae (Fig. 6.86) Prodorsal sclerite absent; coxal apodeme I

fused medially with coxal apodeme II closing coxal fields I in both sexes.
Condylophores elongate and separate in female, asymmetrical in male. Carpoglyphus
lactis (Linnaeus) is a cosmopolitan pest of stored materials with a high sugar con-
tent such as dried fruit, powdered milk, honey and many other stored food material.
Their deutonymphs have also been found as phoretic on insects of order Lepidoptera
and Coleoptera.
6.2.1.2.4 Superfamily Glycyphagoidea
Family Glycyphagidae (Fig. 6.87) Ventral subcapitulum with a prominent pattern

of external transverse and oblique ridges; empodial claws small or absent; condylo-
phores very thin or absent. Female with an epigyneal apodeme and often with an
external copulatory tube; males without paranal suckers or modified setae on tarsus
IV. Commonly found in nests of vertebrates, stored products, house-dust or occa-
sionally on plant foliage. Aeroglyphus robustus (Banks) is commonly found in
stored food products.
Fig. 6.86 Carpoglyphidae: (a) Dorsum. (b) Venter
Family Chortoglyphidae (Fig. 6.88) Body without microtrichia; male with

paranal suckers and often with sucker-like setae on tarsus IV. Chortoglyphids have
highly reduced coxal apodemes III–IV and the males are only glycyphagids to pos-
sess paranal suckers. Dorsal setae are mostly spine-like, b ifurcate apically and ante-
riorly directed. Commonly found in nests of rodents, stored products and house-dust.
Chortoglyphus arcuatus (Troupeau) is many times reported on stored foods.
6.2.1.2.5 Superfamily Acaroidea
Family Acaridae (Fig. 6.89) Prodorsal sclerotization in the form of a rectangular

field that may be incised posteriorly, or prodorsal sclerotization absent; dorsal setae
variable but not usually both elongate and heavily barbed. Cuticle smooth or rarely
with small rounded protuberances. Acarus siro and Tyrophagus putrescentiae
(Schrank) are pests of stored grains, cheese and other stored food products.
Rhizoglyphus echinopus (Fumouze and Robin) is reported to feed on bulbs, corms
and tubers.
Fig. 6.87 Glycyphagidae
Fig. 6.88 Chortoglyphidae
Fig. 6.89 Acaridae
Family Lardoglyphidae (Fig. 6.90) Empodial claw bifurcate in females; males

with leg III enlarged, terminating in a large, straight empodial claw. Males with
paranal suckers and sucker-like setae on tarsus IV. Lardoglyphus konoi Sasa and
Asanuma and L. zacheri Oudemans are known as pests of dried meat and fish
products.
Family Suidasiidae (Fig. 6.91) Body distinctly longer than wide; dorsal setae fili-
form and unbarbed; tarsi with proral setae enlarged and claw-like. Deutonymphs
lack a pretarsus IV, but feeding stages possess a mammillate cuticle; sejugal furrow
is found. Suidasia and Neosuidasia species are known from stored food products
and in nests of bees, wasps and birds.
6.2.1.2.6 Superfamily Pterolichoidea
Family Freyanidae (Fig. 6.92) Body is generally broad; prodorsum with two ros-
tral setae or none; posterior legs ventrally inserted. Parasites of many orders of
aquatic birds. Freyana spp. are found associated with the wing and occasionally tail
feathers of aquatic birds.
6.2.1.2.7 Superfamily Analgoidea
Family Analgidae (Fig. 6.93) Ambulacral stalk inverted on the ventral tarsal apex
to form a distinctly convex surface; the ambulacral discs are relatively small and the
anterior legs bear apophyses and spines. Males of many genera exhibit polymor-
Fig. 6.90 Lardoglyphidae
Fig. 6.91 Suidasiidae
Fig. 6.92 Freyanidae
Fig. 6.93 Analgidae
phism, with the enlargement and the elongation of leg III. Large populations of
Megninia ginglymura (Megnin) mites result in weight loss and decline in egg pro-
duction of poultry birds.
Family Pteronyssidae (Fig. 6.94) Proximal enlargement of the ambulacral stalk as

a membrane along the ventral side of the tarsus; lateral sclerites of ambulacral disc
with distinct clear areas (lacunae). They are parasites on many bird species.
Pteroherpus is an important genus of feather mites.
Family Proctophyllodidae (Fig. 6.95) Most species are elongate and narrow, and
all lack propodosomatic setae and the solenidion on genu II. Females and many
males are bilobed posteriorly, with females possessing elongate projections from
the posterior lobes. Parasites of many hummingbird species. Allodectes and
Proctophyllodes are important genera containing important bird parasites.
Family Dermoglyphidae (Fig. 6.96) Elongated; both sexes with legs III and IV
ventrally inserted; anus terminal. They are parasitic in quills region of many bird
species. Female epigynial shield and male paranal suckers absent; and ambulacral
stalk convex ventrally.
Fig. 6.94 Pteronyssidae
Fig. 6.95 Proctophyllodidae
Fig. 6.96 Dermoglyphidae
Family Epidermoptidae (Fig. 6.97) Earlier known as Knemidocoptidae . Body

sac-like or elongate; prodorsal sclerite with a strongly developed internal apodeme
on each side. All legs very short. Infests feather follicles or skin lesions of numerous
bird species. Epidermoptes and Myialges are containing representatives ectopara-
sitic on birds and mammals. Knemidocoptes spp. are burrowing mites found on
poultry and domestic birds.
Family Syringobiidae (Fig. 6.98) Dorsal hysterosoma with a single, undivided

shield; adults are comparatively well sclerotized. They are inhabitant of quills of
birds. Plutarchusia spp. cause considerable damage and weakening of the feathers
(Proctor 2003).
Family Pyroglyphidae (Fig. 6.99) Tarsus II with solenidion ω (omega) inserted in

basal half of the segment. Largely unsclerotized dorsally except for a propodoso-
matic sclerites (e.g. Dermatophagoides) to being virtually completely sclerotized
(e.g. Pyroglyphus). Coxal fields are well sclerotized. The female epigyneal apodeme
ranges from a large inverted ‘U’ to a smaller almost transverse form. Pretarsal
ambulacral disc may be relatively large and sclerotized internally. Infests many bird
species or mammal nests, house-dust or stored food products.
Family Cytoditidae (Fig. 6.100) Chelicerae with fixed digit absent or entire che-
licerae vestigial; leg and body setae highly reduced. Body rounded; strongly
regressed chelicerae; strong reduction of sclerotizations. Loss of all paraproctal
setae and strong reduction in size of the remaining setae. Members inhabit the respi-
Fig. 6.97 Epidermoptidae
Fig. 6.98 Syringobiidae
Fig. 6.99 Pyroglyphidae: (a) Dorsum. (b) Venter

Fig. 6.100 Cytoditidae
ratory tract of birds. Cytodites nudus (Vizioli) inhabit the lungs, bronchi and air sacs
of poultry birds.
6.2.1.2.8 Superfamily Sarcoptoidea
Family Listrophoridae (Fig. 6.101) Commonly known as fur mite family.

Presence of a projecting tegmen that extends over the gnathosoma to contact the
host air. Propodosoma is sclerotized, and hysterosoma may also bear sclerites.
Parasites of many mammals. Listrophoroides cucullatus (Trouessart) is a wide-
spread fur mite of rats in tropical regions.
Family Chirodiscidae (Fig. 6.102) Strongly modified legs I–II terminate in flat,
plate-like attachment organs clasp host hairs; pretarsal elements absent from legs
I–II. They are parasites of mammals and rodents. Chirodiscoides caviae Hirst is a
common fur mite of laboratory guinea pig.
Family Psoroptidae (Fig. 6.103) These mites possess long legs with all the legs
projecting beyond the body margin. Anterior apodemes of coxal field not fused to
form a sternum; body rounded; pointed mouthparts; female opisthosoma rarely
bilobed; males with or without bilobed opisthosoma. They are mostly skin parasites
and sometimes also infest the inside of ears of many mammals. Psoroptes spp. and
Otodectes cynotis (Hering) feed on the skin or ears of sheep, livestock, horses and
domestic rabbits.
Fig. 6.101 Listrophoridae
Fig. 6.102 Chirodiscidae
Fig. 6.103 Psoroptidae
Family Sarcoptidae (Fig. 6.104) Also known as burrowing mites; short legs which
rarely project the body region. Females are typically globose, but some are elon-
gated dorsoventrally. Parasites of many mammalian hosts. Sarcoptes scabiei
(Linnaeus) is a parasitic mite of humans that form subdermal burrows parallel to the
skin surface in susceptible individuals.
Family Pneumocoptidae (Fig. 6.105) Rounded body with strongly abbreviated

opisthosoma; leg IV situated near the posterior end of the body and a well-devel-
oped sclerite that covers most of the dorsal hysterosoma. Pneumocoptid mites infest
the lungs of rodents. Pneumocoptes penrosei (Weidman) and P. banksi (Baker)
parasitize ground squirrels and dogs.
Family Gastronyssidae (Fig. 6.106) Dorsal hysterosoma without sclerites; tarsus

is very short and is equal in length or shorter than tibia. They are endoparasites of
nasal passages of bats.
Family Myocoptidae (Fig. 6.107) Legs III–IV of female and III of male strongly
modified for clasping hairs; rarely leg IV of female reduced. They have well-devel-
oped apophyses in leg segments. Body usually dorsoventrally flattened and grasp
one hair in legs III–IV on each side. Myocoptes musculinus Koch sometimes appears
in serious proportions in laboratory mouse colonies.
Fig. 6.104 Sarcoptidae
Fig. 6.105 Pneumocoptidae
Fig. 6.106 Gastronyssidae
Fig. 6.107 Myocoptidae
Further Reading 157
6.3 Conclusions
The Acari are quite small in size (80 μm to 13 mm) having varied modes of life.
More than 55,000 species of different acarine species are known (5 % of king-
dom Animalia) distributed in 540 families among 124 superfamilies. Subclass
Acari is divided into two superorders, namely, Parasitiformes and Acariformes.
There are over 40,000 species found under superorder Acariformes, while more
than 15,000 species have been reported under superorder Parasitiformes. In
superorder Parasitiformes, representatives belonging to orders Holothyrida and
Opilioacarida are comparatively less diversified and have lesser economic
importance. However, representatives belonging to order Ixodida to which ticks
belong are of great economic importance and have been studied in greater details
because of their more concern for mankind and veterinary animals. More than
896 species of ticks (including both hard and soft ticks) are described. Order
Mesostigmata includes many species of great economic importance under fami-
lies Phytoseiidae, Dermanyssidae, Varroidae, Parasitidae, Digmasellidae,
Eupodidae and Diarthrophallidae.
Superorder Acariformes is more diverse of the two superorders and includes over
40,000 described species in 351 families. Acariformes is divided into two orders,
namely, Trombidiformes and Sarcoptiformes. Order Trombidiformes is further
divided into two suborders, namely, Prostigmata (including almost all phytopha-
gous mites) and Sphaerolichida. Order Sarcoptiformes earlier known as Acaridida
and Oribatida is further divided into suborders Endeostigmata and Oribatida.
Endeostigmatans are soil inhabiting, but their importance is not well known.
However, suborder Oribatida has too much diversity of mites and includes impor-
tant representatives under superfamilies Histiostomatoidea, Hemisarcoptoidea,
Glycyphagoidea, Acaroidea and Sarcoptoidea.
Further Reading
Baker EW, Tuttle DM (1970) The false spider mites of Mexico (Acari Tenuipalpidae). USDA Tech.
Bull No. 1706. USDA, Washington, DC
Baker EW, Tuttle DM (1994) A guide to the spider mites (Tetranychidae) of the United States.
Indira Publishing House, West Bloomfield
Balogh J, Balogh P (1992) The oribatid mites genera of the world, vol 1. National Museum Press,
Budapest
Bolland HR, Gutierrez J, Flechtmann CHW (1998) World catalogue of spider mite family (Acari :
Tetranychidae). Koninklijke Brill, Leiden
Cook DR (1967) Water mites from India. Mem Amer Entomol Inst 9. Gainesvilla, FL
Helle W, Sabelis MW (eds) (1985) Spider mites: their biology, natural enemies and control. World
Crop Pests, vol 1B. Elsevier, Amsterdam
Holtzer TD, Norman JM, Perring TM, Berry JS, Heintz JC (1988) Effects of microenvironment on
the dynamics of spider mite populations. Exp Appl Acarol 4:247–64
Jeppson LR, Keiffer HH, Baker EW (1975) Mites injurious to economic plants. University of
California Press, 529 pp
Krantz GW, Walter DE (2009) A manual of acarology 3rd edn, pp 97–103. Texas Tech University
Press, 807pp
Lindquist EE (1969) Review of holarctic tarsonemid mites (Acarina: Prostigmata) parasitizing
eggs of pine bark beetles. Mem Entomol Soc Can 60. Ontario Entomol Soc Canada
Mahunka S (1965) Identification key for the species of the family Scutacaridae (Acari:
Tarsonemini). Acta Zool Acad Sci Hung 11:353–401
Meyer MKP (1969) Some stigmaeid mites from South Africa (Acari: Trombidiformes). Acarologia
11:227–271
Meyer MKP, Ryke PK (1959) Cunaxoidea (Acarina: Prostigmata) occurring on plants in South
Africa. Ann Mag Nat Hist Ser 132:369–384
Prasad V, Cook DR (1972) The taxonomy of water mite larvae. American Entomological Institute,
Gainesville
Proctor HC (2003) Feather mites (Acari: Astigmata): ecology, behaviour and evolution. Annu Rev
Entomol 48:185–209
Proctor HC (2004) Aquatic mites: from genes to communities. Kluwer Academic Publishers,
Dordrecht/London
Smiley RL, Gerson U (1995) A review of the Tenuipalpidae (Acari: Prostigmata) of Australia with
description of two new genera and four new species. Int J Acarol 21:33–45
Sonenshine DE (1993) Biology of ticks, vol 2. Oxford University Press, New York
Sonenshine DE, Roe BM (2014) Biology of ticks, 2nd edn. Oxford University Press, London
Sonenshine DE, Lane RS, Nicholson L (2002) Ticks (Ixodida). In: Mullen GR, Durden LA (eds)
Medical and veterinary entomology. Academic, San Diego, pp 517–558
Walter DE, Proctor HC (2013) Mites: ecology, evolution and behaviour. Springer, Dordrecht/
Heidelberg/New York
1. https://www.youtube.com/watch?v=NOwnSW4v-Qs. Introduction to Acariform Morphology

2. http://itp.lucidcentral.org/id/mites/invasive_mite/Invasive_Mite_Identification/key/Whole_
site/Acari_Mites.htm. Mites or Acari
3. https://www.youtube.com/watch?v=-yolQljY_6I. Destroy Spider Mites On Marijuana
Cannabis
4. https://www.youtube.com/watch?v=3JNprSWv96w. Introduction to the Acari
5. https://www.youtube.com/watch?v=2rz36ufAfEQ. Life cycle of spider mites
6. https://www.youtube.com/watch?v=kAQNoEp8P0o. Ticks up close
7. https://www.youtube.com/watch?v=zZigF4_t1Zk. Family Tetranychidae
8. http://idtools.org/id/mites/flatmites/index.php. Flat Mites of the World
9. http://eol.org/pages/2586535/names?all=1. Acari: About Higher Groups & Families
10. http://www.cdc.gov/nceh/ehs/Docs/Pictorial_Keys/Acarina.pdf. Acarina: Illustrated Key to
some common adult female mites and adult ticks
11. http://www.cdc.gov/nceh/ehs/Docs/Pictorial_Keys/Ticks.pdf. Ticks: Key to Genera of Ticks in
United States
12. https://www.youtube.com/watch?v=eATpttq6DfA&index=198&list=PLlRvaTnNBxia9NlhGf
dI5-zgytIv75qbG. The Tick Key
13. https://www.youtube.com/watch?v=QM0kBwSU3hc&index=145&list=PLlRvaTnNBxia9Nl
hGfdI5-zgytIv75qbG. Tick borne diseases of cattle
14. http://www.nappo.org/files/3714/3782/0943/DP_03_Tetranychidae-e.pdf. DP 03:
Morphological Identification of Spider Mites (Tetranychidae) Affecting Imported Fruits
15. https://www.unce.unr.edu/publications/files/nr/2004/FS0447.pdf. Ornamental Plant Damage
By Eriophyid Mites
Further Reading 159
16. https://www.youtube.com/watch?v=ecTauvwdeZg#t=1116.549386. Eriophyoidea

17. https://www.youtube.com/watch?v=UjtdMivX7x0&ebc=ANyPxKqTbw8oA1uKdtQf56PjXe
07YSkF2kai9Rz_ECNi0nrgQpn78-B82EiTNeDOWYVANk3QbO9MIi8FmfcFIh5XiDMMn
0dUfA. Family Tarsonemidae
18. http://itp.lucidcentral.org/id/mites/invasive_mite/Invasive_Mite_Identification/key/0_
Glossary/Mite_Glossary.htm#Vertical-scapular system.
Feeding, Development
and Reproduction 7
Abstract
Life cycle of different mites and ticks passes through seven developmental
stages, namely, eggs, prelarva, larva, protonymph, deutonymph, tritonymph and
adult. As with other arthropods, each stage is separated by a moult, and consider-
able variations in life cycles in different Acari are found. Acarines do not con-
tinue to moult once they have reached the adult stage. Comparative variations
found in life cycle of different groups particularly in mites and ticks are dis-
cussed for better understanding. Differences in feeding activities in mites and
ticks are particularly highlighted. Within ticks, feeding habits of hard ticks and
soft ticks are reported, and stages in the life cycle passed on a number of hosts
and egg laying patterns are explained. Blood feeding pattern and changes in body
shape after each feeding are reported. The type of feeding habits of plant feeding
mites and the extent to which tissue level they penetrated are noticed causing a
variety of damage to the plant. Feeding by spider mites, false spider mites, broad
mites and eriophyid mites is particularly reported. Mites exhibit a variety of
reproductive strategies in different groups. In several groups of acariform mites,
direct insemination is done through an intromittent organ, the aedeagus, in the
male. Secondary sexual dimorphism is also found in some groups. Sex determi-
nation mechanisms and reproductive modes also vary widely throughout the
Acari. Activities like maintenance of water balance and the occurrence of dia-
pause in mites and ticks are reported in good length.
Keywords
Life cycle • Diapause • Water balance • Feeding patterns

DOI 10.1007/978-981-10-1594-6_7
162 7 Feeding, Development and Reproduction
7.1 Development
Different mites and ticks during their life cycle pass through seven developmental
stages, namely, eggs, prelarva, larva, protonymph, deutonymph (or deuteronymph),
tritonymph and adult. As with other arthropods, each stage is separated by a moult,
and considerable variations occur in life cycles in different Acari. Since the nymphal
stages resemble one another, four distinct phases in the life cycle are recognized:
prelarva, larva, nymph and adult. The prelarva and larva have three pairs of legs,
while the nymphs and adults have four pairs of legs. The fourth pair of legs appears
in the protonymphal stage. Acarines do not continue to moult once they have reached
the adult stage (Dhooria 1981; Gupta et al. 1972). Different stages that occur in the
life cycle of mites and ticks are described briefly(ER 1 & ER 7).
7.1.1 Eggs
Many parasitic mites lay their eggs at places where they are protected from different
climatic hazards, and their access to the next host is ensured. In spider mites (family
Tetranychidae), the eggs may be suspended or tied with silken guy wires to the sur-
faces on which they are laid. Winter and summer eggs of spider mites differ in pig-
mentation and placement on the host plant. Eggs are glued to the leaf surface of the
host plant by a sticky substance (Van de Vrie et al. 1972). Parasitic mites of the
family Listrophoridae fix their eggs to the hairs of their lice host. In case of spider,
mites from 20 to 200 eggs per female are laid. The number of eggs laid by female
ticks varies considerably. Generally soft ticks lay less number of eggs per female,
but some species of hard ticks lay as much as 20,000 eggs per female. The eggs of
all ticks are laid in the physical environment but never on the host (Sonenshine and
Roe 2014).
7.1.2 Prelarva
Prelarva is a nonfeeding form that occurs in Acari and some other arachnids. In
some cases, this stage has also been referred to as deutovum. In some cases prelarva
appears to be little more than a featureless sac without legs or mouthparts. Extreme
inhibition of this type is referred to as calyptostasis and is found in some
Endeostigmata, Oribatida and certain higher Prostigmata. In other acariform groups,
the inactive prelarva may possess three pairs of legs, recognizable mouthparts and
setae, although mouthparts development and leg segmentation may be inhibited.
Some scientists have referred to incomplete calyptostasis as elattostasis. Extreme
form of elattostasis is found in families Nanorchestidae, Adamystidae and Anystidae.
Acariform prelarva share certain characteristics, including urstigma (Claparede’s
organ) and a non-functioning stomodeum. The prelarva moults to the larva (Krantz
and Walter 2009).
7.1 Development 163
7.1.3 Larvae
Larvae emerge from fully developed eggs and bear three pairs of legs (ER2). The
larval stage is characterized through little or no sclerotization and the lack of exter-
nal genitalia. Larvae may be inactive and nonfeeding (in Gamasida), voraciously
predaceous (in Cheyletidae, water mites) or parasitic (Trombiculidae). Larvae of
Trombiculidae are known as chiggers; and larval characters in Trombiculidae are
considered very useful in species determination. The urstigma present in some
acariform prelarvae persist in the larval stage when fully fed. Larvae of
Pterygosomatidae, Trombidiidae and water mites enter into a resting stage com-
monly known as nymphochrysalis which later emerges as nymphal stage after rup-
turing the old cuticle. In some groups larvae tend to be weak and sluggish nonfeeding
forms (as in many free living Mesostigmata). The larval stage of virtually all species
of argasid ticks differs from conspecific nymphal and adult stages in that it attacks
and feeds on vertebrate hosts for an extended period (days) rather than the shorter
attachment period (months to a few hours) of the nymph and adult. Because larval
stages attach to the host for the extended period, this life stage is typically collected
more commonly than conspecific post-larval stages, and many argasid species are
well known only from the larval stages (Evans 1992).
7.1.4 Nymphs
Active proto-, deuto- and tritonymphal stages occur in the Holothyrida, Oribatida,
Opilioacarida and in many Endeostigmata, Prostigmata and Astigmata. A maximum
of two nymphal stages occur in Mesostigmata (the protonymphs and deutonymphs),
while in some Prostigmata (most Heterostigmatina), there may be no free nymphal
stage. Only one active nymphal instar, the deutonymph, occurs in some Prostigmata,
with the protonymphal and tritonymphal stages passing as calyptostatic forms
within the skin of the preceding stages (ER2).
The nymphs of ticks have eight legs like the adult but have no genital opening.
Nymphal stages must undergo a critical waiting period for a suitable host. After
engorgement, the nymph drops from the host, moults and becomes an adult. Nymphs
may rest for long periods before becoming adults. Hard ticks have only one nymphal
instar, the nymph becoming an adult after moulting. Soft ticks may have several
nymphal instars.
7.1.5 Protonymph
The first nymphal stage following the larva is known as protonymphal stage. This
stage is usually free active instar that may or may not feed. It is usually found in and
adapted to the environmental location where the other nymphal stadia will occur.
Only one nymphal instar occurs in most of parasitengonine Prostigmata that develop
within the skin of the preceding instar. But there are two sequestered calyptostases:
one between larva and active deutonymphal instar (nymphochrysalis) and the other
between the deutonymph and adult (teleiochrysalis).
7.1.6 Deutonymphs
The deutonymph is the second nymphal stage in the life cycle and may differ from
the adult only in size, the patterns of sclerotization and setation. Phoretic
deutonymphs of Uropodina and parasitic Gamasida do not feed. Deutonymphs of
many Astigmatina are heteromorphic, i.e. are completely different morphologically
and in behaviour from preceding and succeeding stadia. Such heteromorphic
nymphs are called hypopi (= hypopodes) which occur sporadically. Hypopi lack
functional mouthparts and are principally phoretic stages. This special phenotype of
a deutonymph can survive bad environmental influences much better than the nor-
mal form as in family Glycyphagidae. In most cases, the deutonymphs moult to the
adult stage.
7.1.7 Tritonymphs
In most cases quiescent deutonymph moults to the adult stage, but in some groups
of mites, the third nymphal stage (i.e. tritonymph) develops. It is usually an active
stage, but may be pharate in some members of Astigmatina. Tritonymphal stage is
absent in the Mesostigmata and may be absent and represented by a pharate calyp-
tostatic stage (teleiochrysalis) in many Prostigmata. This stage is retained in some
Holothyrida.
7.1.8 Adults
Adult mites are comparatively larger in size and sexually mature. In most mites, the
adult is easily recognized by the appearance of fully formed genital regions and
often has distinctive sclerotized shields. Sometimes they have an ovipositor, sper-
matophoric organs or secondary sperm transfer systems. In case of ticks in addition
to egg stage, only three instars are recognized in life cycle, i.e. larva, nymph and
adult. There is only one nymphal instar in the ixodid tick life cycle, but varying
number of nymphal stages may occur in the argasid tick life cycles.
7.1.9 Life cycle
7.1.9.1 Mites
Most mites exhibit a fixed developmental pattern, passing through the same number
of instars regardless of how much food is available (ER 2). The time of development
from the egg to the adult varies from hours to a few days, to weeks, months and
7.1 Development 165
years. The life cycle duration is greatly affected by humidity, temperature and food
supply. Some species of family Macrochelidae complete their life cycle in 1.5 days,
but the tick, Dermacentor andersoni Stiles, completes its life cycle in 2 years. In yet
another tick species Ixodes uriae White, life cycle is completed in 4–5 years. Soil
mites (Oribatida) and water mites (Hydrachnidia) have very slow development
rates. Many species of family Phytoseiidae have proved as potential predators of the
pest mites, because of their rapid developmental rates than the pest mites (McMurtry
and Croft 1997). Many members of families Pyemotidae, Acarophenacidae,
Pygmephoridae, Histiostomatidae and Tarsonemidae have comparatively shorter
life cycles (4–8 days). In case of water mites, the larvae can be divided into three
types based on the habit of the parasitic stage in the life cycle. In the first type, lar-
vae do not leave the water but parasitize the aquatic insect as in some Hydrachnidae.
In the second type, parasitic larvae occur on insect living on the water surface or in
the air stores of aquatic insects in direct contact with water during the parasitic
phase (as in Limnocharidae and Eylaidae). In the third type, larvae parasitize insects
which live in air.
7.1.9.2 Ticks
Ticks have four generalized life cycles. These life cycles are related to the number
of individual animal hosts a tick will visit and feed on during its life from egg to
adult. These life cycles are commonly known as one-host, two-host, three-host and
multi-host life cycles. In ixodid ticks, three-host life cycle is most common. The
larvae find a host and feed for some days, then drop off and digest the blood and
moult to a nymphal stage. The nymph finds a new host (of the same or a different
species of animal), feeds on blood again, drops off and digests the blood. Then the
nymph moults to the adult stage. These ticks quest for a new third host and feed and
mate on this new third host. After fully feeding, the females drop off, digest the
blood and lay eggs. Males stay on the host, often do not feed and die after mating.
Many ticks have the eggs and moulting stages in soil or vegetation in the environ-
ment in which their hosts graze or hunt. The ticks crawl onto vegetation and wait for
their hosts to pass by. Thus in the ixodid ticks such as Rhipicephalus, the larvae,
nymph and adults will quest on vegetation. The ticks grasp on to the host using their
front legs and then crawl over the skin to find a suitable place to attack and feed.
Adult ticks of the genera Amblyomma and Hyalomma are active hunters and will run
across the ground to seek hosts and are resting nearby. Other ticks such as nearly all
argasid ticks and many Ixodes species spend their life cycle near the nests or in
shelters of their hosts. Carbon dioxide released by hosts stimulates questing in ticks.
Three-host life cycles may take years to complete depending on the environmen-
tal and climatic conditions in the area. The two-host life cycle also occurs in some
species of hard ticks which are extremely important pests of animals in some parts
of the world. The one-host life cycle occurs in some ixodid tick species. Larvae,
nymphs and adults all feed upon the same animal host without dropping off from it
to moult. One host ticks are important pests of domesticated animals in many parts
of the world. The adult female lays many eggs (2000–20,000) in a single batch in
their physical environment but not on hosts.
The multi-host life cycle is characteristic of nearly all of the species of soft ticks.
Multi-host life cycles take place in areas where host animals dwell (in burrows and
nests). In this life cycle, many nymphal moults occur and such nymphs are known
as intermediate stages. Then they detach from the host, stay in the shelter, digest the
blood and moult to the adult stage. The adult ticks may feed many times and ovi-
posit in small batches of eggs after each blood feeding. Most argasids are multi-host
ticks, but Otobius megnini (Duges) has a one-host life cycle.
7.2 Feeding
The nature and type of feeding by ticks and mites on their hosts differs and is dis-
cussed separately for each group below.
7.2.1 Ticks
Mode and patterns of feeding by both hard and soft ticks vary significantly and are
discussed below.
7.2.1.1 Hard Ticks

In ixodid or hard ticks, life cycle comprises the egg and three active stages: larva,
nymph and adult. There is only one nymphal instar in the life cycle of these ticks.
Hard ticks feed slowly from several days to as long as 2 weeks. Immature and adult
ticks each take a blood meal except for the nonfeeding males of some species. After
crawling onto their hosts (ER 3), these ticks embed their mouthparts into the skin of
the host and secrete cement from their salivary glands into and around the wound
site to anchor themselves (ER 5 & ER7). The cement binds the ticks firmly in place
and makes them very difficult to remove. During blood feeding, the ticks secrete
potent anticoagulants and anti-inflammatory agents, which suppress host wound
healing and facilitate blood flow. Often blood meals from 10 to 100 times of their
original body weight are consumed by a tick. Females feed only once; mating
occurs during feeding. Following mating, females suck blood rapidly for 24–48 h
and swell enormously. Then the replete females drop from their hosts, find a shel-
tered location and lay thousands of eggs. Following oviposition, the female dies. In
contrast to females, males swell only slightly during feeding. Male ticks remain on
the host and will attempt to mate with many females during repeated feedings. The
females mate only once before they are ready to engorge fully with the blood. In
prostriate ticks (Ixodes spp.), males have vestigial hypostomes, and these ticks
always mate off the host. In general, more than 90 % of the life cycle of hard ticks
is spent off the host(ER 8).
When host seeking and feeding occur in all three parasitic stages, the pattern is
termed a three-host life cycle. A few ixodid species exhibit a two-host or three-host
life cycle. For example, in the camel tick, Hyalomma dromedarii Koch, both larvae
and nymphs feed on the same host (two-host life cycle), and in the cattle tick
7.2 Feeding 167
Boophilus annulatus (Say), all stages feed, moult and even mate on the same host
(one-host life cycle).
7.2.1.2 Soft Ticks

Feeding is very rapid among soft ticks. Once they have crawled onto a host, the tick
embeds their mouthparts in the same way as hard ticks but without secreting cement.
Bloodsucking commences quickly, and as feeding progresses, the bloated ticks
excrete copious amounts of a clear, colourless coxal fluid. By eliminating excess
water and salts via the coxal fluid, these ticks can concentrate their blood meals and
adjust their internal water balance. The ticks expand to about 5–10 times their origi-
nal body weight depending on the ability of the cuticle to stretch. Following feed-
ing, often within as little as 30–60 min, the replete ticks drop off to moult or, if
female, to lay eggs. Soft tick females take repeated small blood meals and lay small
batches of eggs (less than 500 eggs in a batch) after each feeding multiple gonotro-
phic cycles. The interval between feedings is typically several months but may be
up to several years, depending on host availability. Mating usually occurs off the
host. Because of the multiple nymphal instars – six or even seven in some species –
argasid ticks live for many years. In addition, these ticks are highly resistant to
starvation. As a result, the entire life cycle may take 10–20 years (Sonenshine and
Roe 2014).
7.2.2 Phytophagous Mites
The gnathosomal structures display different modifications among the phytopha-

gous Acari, i.e. Tetranychidae, Tenuipalpidae, Eriophyidae and Tarsonemidae. Most
of the acarines are members of the order Trombidiformes and possess mouthparts
specialized for piercing and sucking the juices of the plants. Spider mites
(Tetranychidae) and false spider mites or flat mites (Tenuipalpidae) are the major
groups containing phytophagous mites. Mouthparts in all spider mites are piercing
and sucking type and consist of a stylophore at the gnathosomal base from the
underside of which hang a pair of cheliceral or mouth stylets, a pair of palps and the
rostrum situated between and directly under the palps. The stylophore is protrusible,
moving back and forth during feeding. The stylophore pushes the stylets down to
the plant tissue and upon retraction probably allows an opening through which plant
tissues may be forced out of the cells by the plants’ turgor pressure.
In the false spider mites, adults are smaller than spider mites and are more slug-
gish, and their damage is felt after a long feeding time. However, some false spider
mites like Larvacarus transitans (Ewing) have been reported to make tiny closed
packed galls for feeding purposes. These mites depend on keeping the plant tissues
alive for a longer time. Movable cheliceral digits are modified into long, whip-like
stylets that are strongly recurved proximally, arising in an eversible stylophore.
In worm or eriophyid mites, stylophore is absent or a rudimentary one. The che-
licerae are modified into short, needle-like stylets which lie in a groove in the ante-
rior surface of the rostrum. Stylets are short and are capable of only shallow
penetration into the host. Eriophyid mites depend for the most part of their life upon
keeping the plant tissue alive so that they can continue to feed on them (Lindquist
et al. 1996).
In case of broad mites (Tarsonemidae), oral appendages are contained in a dis-
tinct capsular head known as capitulum. Mouthparts consist of stout paired palpi
inserted on that apical portion of the capitulum and slender styliform paired chelic-
erae. The palps are quite reduced in these mites and the apical snout is formed by
the terminal palpal segments. The chelicerae are very short and needle-like and are
incapable of penetrating the tough growth. That is why many tiny mites are found
on both surfaces of young succulent growth of leaves.
7.3 Reproduction
7.3.1 Mites
Mites exhibit a variety of reproductive strategies and modes of transfer. Ancestrally,

mites appear to produce indirect sperm transfer, with males producing and deposit-
ing a package of sperm (e.g. spermatophore) on the substrate. Females then take the
spermatophore into their reproductive tract. This type of reproduction is found in
most acariform subgroups, and individuals of the two sexes may or may not be close
in contact at the moment of insemination. In many parisitiform groups, males typi-
cally use their chelicerae to assist in directly inserting a spermatophore into their
females’ primary genital opening (as in primitive Gamasidae), or the male chelic-
erae bear an organ termed the spermatodactyl which is used to transfer sperm from
the males’ genital opening into secondarily developed sperm induction pores near
the bases of the females’ legs. These paired openings lead to a median spermatheca
which is connected directly to the ovary, where fertilization takes place (Evans
1992).
In several groups of acariform mites, direct insemination is done through an intro-
mittent organ, the aedeagus in the male. Secondary sexual dimorphism typically
accompanies direct mating with males having modified appendages for holding the
female during mating. In many cases, precopulatory guarding of immature females,
either waiting near a juvenile female about to moult or actively attaching to her is
observed. Sex determination mechanisms and reproductive modes also vary widely
throughout the Acari. Some mites are diploid in both sexes, with males having either
a Y chromosome or no sex chromosome. Other mites are arrhenotokous, in that
females are diploid and males haploid. Such males develop from unfertilized eggs.
An unusual reproductive mode, termed parahaploidy, is found in some Gamasida. In
these mites, fertilization is necessary for egg development, but in males the paternal
genome is inactivated shortly after the first embryonic cell divisions, and adult males
are functionally haploid. Thelytoky or all female parthenogenesis is found in many
groups of mites. Such mites reproduce clonally, with diploid eggs developing directly
into females without fertilization. But the chief mode of reproduction in many acari-
form mites (Eriophyidae and Tetranychidae, Gamasida, Dermanyssidae, Laelapidae,
Macrochelidae and Phytoseiidae) is arrhenotoky. But arrhenotoky is not found in
7.4 Tick Ecology 169
Ixodida and Oribatida. Thelytoky is known in all orders of Acari (excepting

Opilioacaridae and Holothyrida). Thelytokous parthenogenesis is rarely found in
Gamasida, but is commonly found in some ticks and prostigmatid and oribatid mites
(Jeppson et al. 1975; Krantz and Walter 2009).
7.3.2 Ticks
Reproduction of ticks is closely associated with feeding. This is of parasitological

significance since many pathogens of veterinary and medical importance are trans-
mitted transovarially to the progeny of female ticks which have taken up the patho-
gens with their blood meal (ER 7). Because many species can lay a very large
number of eggs, this mode of transmission can become a most efficient means of
multiplying the pathogens (viruses, bacteria, rickettsia or protozoans). Reproduction
in ticks is not only of direct interest for the maintenance of population but also has
a serious economic significance in relation to tick-transmitted diseases (Sonenshine
and Roe 2014).
In argasid females, feeding and oviposition are cyclical activities which can be
repeated several times (seven or more times). Mating can also be repeated in asso-
ciation with each female feeding. Mating can take before or after feeding. Mated
females can digest the blood meal and oviposit, after which they are ready to repeat
the process. Virgin soft ticks, on the other hand, interrupt vitellogenesis until mating
takes place, or even if mating does not occur for a long time.
In ixodid ticks, larvae, nymphs and females all take a single complete meal after
which they moult to the next instar. Thus feeding and oviposition are each single
events in the life time of a female. Mating is only possible up to the completion of
feeding and is usually not possible prior to feeding. In most ixodid species, mating
takes place on the host only after attachment (except in Ixodes spp.).
Generally argasid females lay fewer eggs than ixodid females. The argasid or
soft tick female produces a small batch of eggs (100–500); females repeatedly feed
and then lay eggs. Up to six feedings and egg layings have been reported (Vial
2009).
7.4 Tick Ecology
A tick’s habitat is composed of a variety of living and nonliving things in the space
in which it lives. Ticks are adapted to two contrasting components of their habitat:
the physical environment and their host. When ticks are moulting and then questing
in the physical habitat, they are in danger of drying out and starving. The larvae are
most susceptible because they have a high surface area relative to their small vol-
ume. They are also exposed to predators such as rodents, birds, reptiles and ants and
to pathogens such as fungi. These adverse factors limit type of habitats that a spe-
cies will be found in and knowledge of the typical physical habitat of a species aids
in their identification. The most important component of the physical habitat of tick
is the climate that is defined by temperature and humidity. When the same tick is on
the host, it is no longer in danger of being removed by the hosts’ growing or having
its feeding reduced by host immunity. Most ticks have adaptations in their behav-
iour and physiology of feeding to reduce host reactions. The survival of a population
of ticks depends on the presence of hosts suitable for reproduction by the adults.
Before oviposition can begin, the fully engorged females usually become geotropic
and negatively phototactic seeking sheltered places with a suitable microclimate.
Oviposition is almost similar in both tick families. Gene’s organ – the egg-waxing
organ, located dorsally behind or above the capitulum – coats thoroughly each
oocyte with waxy waterproof secretions. In ixodid ticks, each oocyte is simultane-
ously exposed to a secretion from the porose areas. The eggs of argasid ticks are
usually not sticky, while ixodid ticks’ eggs adhere to the surface they rest upon
(Gray 1998; Sonenshine and Roe 2014; Vial 2009).
7.5 Water Balance
To survive, Acari must be able to withstand considerable variations/fluctuations of

temperature, relative humidity and different other potential hindrances in their
milieu which affect loss of water from their body. Most Acari have a relatively
impermeable cuticle and have some control of diffusion of water from their tracheal
system through the movement of stylophore. Usually on leaf surfaces, relative
humidity is comparatively quite high than the surrounding arid atmosphere and
helps in development of spider mites. Silken threads secreted by spider mites also
help in maintaining the humidity adjacent to the leaf surfaces. Diapausing females
(nonfeeding) also help in conservation of water. In spider mites, openings of stig-
mata and their positional adjustment of the stylophore also control the area available
for diffusion from the tracheal system. It seems control of diffusion from the tra-
cheal system is the most important factor in maintaining water balance of mites
particularly at low relative humidities. Under severe water stress, the mites can
reduce the oxygen consumption and seal off their tracheal system for long periods
of time (Jeppson et al. 1975).
Cuticle normally serves as an effective evaporation barrier and hence is effective
in controlling water loss if temperature exceeds critical range of 45–50 °C. In spider
mites, strial lobes and microtubercules on skin of eriophyid mites are also helpful in
regulating water loss from the cuticle. However, depending on the prevailing tem-
perature conditions, water can be absorbed from the air. Nonfeeding diapausing
spider mite females lack these lobes over much or all of the body surfaces. Absence
of strial lobes may help them in conserving body water. Diapausing females may
obtain water during hibernation by metabolism of accumulated body stores.
However, some tarsonemid mites move down into inner recesses on their hosts to
avoid adverse effects of dry periods. In some spider mites, eggs have evolved a
peculiar respiratory system to satisfy their oxygen requirements without losing too
much water under extremes of temperature conditions. Argasid ticks are able to
concentrate their blood meals and thus adjust their internal water balance. Argasid
7.6 Diapause in Acari 171
ticks are able to concentrate their blood meals and thus adjust their internal water
balance.
7.6 Diapause in Acari
7.6.1 Mites
Diapause is a form of dormancy which is characterized by developmental arrest. It

confers survival advantages during unfavourable seasonal conditions such as during
winter. Incidence of diapause in Oribatida, Astigmata and Prostigmata has been
reported. Regular occurrence of diapause is observed in many spider mite species
living in climates with an adverse season. Regular occurrence of diapause is
observed in many spider mite species living in climates with an adverse season.
Both aestivation- and hibernation-type diapauses are found in Bryobiinae and
Tetranychinae. Dispausing eggs are reported in members of Bryobiinae (Bryobia
spp. and Petrobia spp.). However, diapausing females have been reported for only
Tetranychinae (Tetranychus spp. and Eotetranychus spp.), but egg diapause has
been reported in genera Eurytetranychus, Panonychus, Oligonychus and
Schizotetranychus. But grass-inhabiting mite species, Oligonychus pratensis
(Banks), diapauses as an adult female. The females of two-spotted spider mite
Tetranychus urticae Koch undergo adult diapause in response to short day length
and low temperature during their immature development. In regions with a mild
winter climate, these mites do not enter diapause, whereas those inhabiting regions
with a cooler climate enter diapause in winter. Reproductive diapause combined
with the post-diapause quiescence may be found after developmental and behav-
ioural diapause in family Phytoseiidae (Gamasida). Only the usual and post-
diapause quiescence are known in Acari (Jeppson et al. 1975; Veerman 1992).
7.6.2 Ticks
Diapause in ixodid ticks can occur at several different life stages and may be mani-
fested as behavioural diapause, involving a form of quiescence of the unfed ticks at
a time when environmental conditions are unsuitable for host seeking or develop-
mental diapause involving arrested development of the engorged stages or of eggs.
The diapause mechanisms seem to enable the tick to avoid entering host-seeking
phases at unfavourable time of the year, such as high summer and mid-winter. The
proportion of the population that exhibits them will vary according to local condi-
tions, areas where a good cover of vegetation and a mat of decaying vegetation are
present. The most important environment stimulus seems to be day length, though
temperature may have modifying influence (Belozerov 2008, 2009).
The peculiar nature of oil skin and cement components of argasid ticks’ cuticle
makes it highly resistant and reduces water evaporation, allowing soft ticks to sur-
vive at high temperatures and under relatively dry conditions. For example,
Ornithodoros moubata (Murray) and O. savignyi Audouin tolerate air temperatures

up to 63 and 75 °C, respectively. This phenomenon results in the common presence
of soft ticks in tropical and subtropical zones and their spread into arid areas of
Central Asia and Africa (Sahara). However, the success of their developmental cycle
depends on cooler conditions, which are provided by the sheltered microhabitats
they colonize. In addition to intrinsic microclimatic conditions of shelters, body
heat and moisture from the respiration and excrement of the host within the micro-
habitat help establish the atmospheric conditions required for the development of
these ticks (Sonenshine and Roe 2014; Belozerov 2009).
7.7 Conclusions
In literature, there have been many studies on both mites and ticks; more studies
have been reported on economically important Acari under different situations.
Through the Internet, many detailed studies are available online mostly on ticks.
Several studies on development, feeding and reproduction of both mites and ticks
have been reported for different species which have facilitated in different situations
the proper management of these creatures. But here in this chapter only salient
information available is considered. For comparison of studies, Internet search will
be more useful for better understanding of behaviour, life cycle and multiplication
of these tiny creatures.
Further Reading
Belozerov VN (2008) Diapause and quiescence as two main kinds of dormancy and their signifi-
cance in life cycles of mites and ticks (Chelicerata: Arachnida: Acari). Acarina 16:79–130
Belozerov VN (2009) Diapause and quiescence as the two main kinds of dormancy and their sig-
nificance in life cycles of mites and ticks (Chelicerata: Arachnida: Acari). Acarina 17:3–32
Dhooria MS (1981) Studies on ovipositional preference, host-range and seasonal availability of
mite, Eutetranychus orientalis (Klein) in Delhi. Indian J Acarol 6:77–83
Dhooria MS (1986) Observations on the biology of spider mite, Eotetranychus uncatus -a serious
pest of Kachnar (Bauhinia variegata) in Punjab. Acarol Newslett 16:4–5
Dhooria MS (1991) Preliminary studies on the biology of spider mite, Bryobia ehari Pritchard and
Keifer (Acari: Tetranychidae) – pest of Chrysanthemum in Punjab. In: Mukherjee AB,
Somchaudhry AK, Sarkar PK (ed) Contributions to Acarological Researches in India. Kalyani,
India
Dhooria MS, Sagar P (1989) Preliminary studies on the biology of carmine spider mite Tetranychus
cinnabarinus, on four species of Japanese mint at Ludhiana, India. In: ChannaBasavanna GP,
Viraktamath CA (eds) Progress in acarology, vol II. Oxford/IBH, Delhi, pp 39–41
Evans GO (1992) Principles of acarology. CAB Internaitonal, Walling, p 563
Fan Y, Petit FL (1998) Dispersal of the broad mite, Polyphagotarsonemus latus (Acari:
Tarsonemidae) on Bemisia argenticola (Homoptera: Aleurodidae). Exp Appl Acarol
22:411–415
Gray JS (1998) The ecology of ticks transmitting lyme borreliosis. Review. Exp Appl Acarol
22:249–258
Further Reading 173
Gupta SK, Dhooria MS, Sidhu AS (1972) Effect of food and temperature on the rate of develop-
ment, fecundity and longevity of Tetranychus cucurbitae Rahman and Sapra. Indian J Agric Sci
42:980–983
Gupta SK, Dhooria MS, Sidhu AS (1973) Effect of food and temperature on the development,
longevity and fecundity of sugarcane red spider mite, Oligonychus indicus (Hirst). Acarologia
16:436–440
Jeppson LR, Keifer HH, Baker EW (1975) Mites injurious to economic plants. University of
California Press, Berkeley
Krantz GW, Walter DE (eds) (2009) A manual of acarology, 3rd edn. Texas Tech University Press,
Lubbock
Kurban R, Loes GMB (1988) Effects of herbivory and plant conditioning on the population
dynamics of spider mites. Exp Appl Acarol 4:225–246
Lindquist EE, Sabelis MW, Bruin J (1996) Eriophyoid mites: their biology, natural enemies, and
control, vol VI. Elsevier, Amsterdam
McMurtry JA, Croft BA (1997) Life-styles of phytoseiid mites and their role in biological control.
Annu Rev Entomol 42:291–321
Sonenshine DE, Roe RM (ed) (2014) Biology of ticks, vol I, 2nd edn. Oxford University Press,
Oxford, pp 560
Van de Vrie M, McMurtry JA, Huffaker CB (1972) III. Biology, ecology, pest status and plant host-
relations of tetranychids. Hilgardia 41:343–432
Veerman A (1992) Diapause in phytoseiid mites: a review. Exp Appl Acarol 19:1–60
Vial L (2009) Biological and ecological characterization of soft ticks (Ixodida: Argasidae) and
their impact for predicting tick and associated disease distribution. Parasite 16:191–202
1. https://www.youtube.com/watch?v=kAQNoEp8P0o Ticks up close

2. https://www.youtube.com/watch?v=2rz36ufAfEQ (Life cycle of spider mites and damage done)
3. https://www.youtube.com/watch?v=wPbIot9tQ6U The process of attachment of a nymphal
Ixodes ricinus tick to host skin
4. https://www.youtube.com/watch?v=PUuC42_h1jQ Use of a Table Top Scanning Electron
Microscope
5. https://www.youtube.com/watch?v=0g_lt0FcQag Tick Lifecycle.mp4
6. https://www.youtube.com/watch?v=nsnGkGJzlyw Mites and ticks (#486) in motion
7. https://www.youtube.com/watch?v=WpOyWesQtgc&list=PLlRvaTnNBxia9NlhGfdI5-
zgytIv75qbG&index=36 Tick Animation
8. https://www.youtube.com/watch?v=m9H3lnRaiXA&index=180&list=PLlRvaTnNBxia9NlhG
fdI5-zgytIv75qbG Reasons Why Ticks Suck
Molecular Biology and Acarology
8
Abstract
Molecular methods broadly refer to techniques used for the recognition of groups
of individuals whether they are population, species or higher taxonomic groups
and ultimately understanding of relationship between these different groups. The
information provided by molecular biology and genomics has direct application
towards the improvement of pest management tools and strategies. Sequence
data along or in combination with morphology or behavioural information can be
used to develop better phylogenies. Within the Acari, the nuclear ribosomal
genes 18 S and 28 S rDNA are equally powerful tools for phylogenetics in the
deepest levels. Molecular tools have been successfully employed in some cases
for the proper identification of spider mites and phytoseiid mites. DNA sequences
obtained from nuclear ribosomal RNA genes (18 S, 5.8 S and 28 S rRNA) and
their spaces (ITS-1 and ITS-2) are being used routinely. The ITS-2 sequence has
been used to identify species from families Eriophyidae, Ixodidae and
Tetranychidae. Through molecular markers some species of spider mites, phyto-
seiid mites and worm mites have been studied at individual, population and spe-
cies level. Population variations have also been studied in the important bee
parasitic mites. Basic information about molecular techniques like RFLP, RAPD,
AFLP, microsatellites and DNA barcoding is provided. Molecular techniques are
proving to be very powerful in issues like tracking invasive species. Molecular
techniques also help in locating successfully host races and species link and for
correct identification of house dust and storage mites. Genome sequencing of
two-spotted spider mite (TSSM) and some ticks is reported.
Keywords
Genomics • Molecular markers • Genome sequencing • Invasive species

DOI 10.1007/978-981-10-1594-6_8
176 8 Molecular Biology and Acarology
Molecular biology is defined as the study of biology at the molecular level; it is an

interdisciplinary approach that integrates biochemistry, physiology and genetics.
Molecular biology emphasizes the central dogma, which is the relationship of DNA,
RNA and protein, and thus by studying DNA, we can learn about protein and cell
biology. Molecular methods broadly refer to techniques used for the recognition of
groups of individuals whether they are populations, species or higher taxonomic
groups and ultimately the understanding of relationship between these different
groups. With the molecular biology and genomics, the biology of pest organisms
can be better known. The information provided by molecular biology and genomics
has direct application towards the improvement of pest management tools and strat-
egies (Guerra-Garcia et al. 2008; Oliver 1977).
8.1 Phylogenetic Relationship
For proposing relationships, the possession of derived features shared with a com-
mon ancestor (synapomorphies) is important. In either case the mutation events that
result in a nucleotide change in the common ancestor of a group are subsequently
fixed in a lineage and are applicable to either recognition or relationships. Alone or
in combination with morphology or behavioural information, sequence data can be
used to develop better phylogenies, classifications and identification keys which are
fundamental to all biological control programmes. The interpretation of environ-
mental or behavioural change on a given phylogeny can as well improve our knowl-
edge about the rate and means of acquiring novel host associations or other
adaptations that might improve our evaluation of new control agents (Dabert 2006:
Dabert et al. 2009; Xie et al. 2008).
The use of molecular information such as ribosomal DNA (mtDNA) sequence
information is now extensively done in phylogenetic studies in case of mites. The
second internal transcribed spacer of the nuclear ribosomal gene cluster (ITS-2) and
the mitochondrial protein-coding gene cytochrome oxidase (COI) together provide
tool for phylogenetics at low taxonomic levels. Within the Acari, the nuclear ribo-
somal genes 18 S and 28 S rDNA are equally powerful tools for phylogenetics in the
deepest levels. The 18 S rRNA gene is considered more appropriate for resolving
relationships among phyla and superphyla, with the 28 S rRNA gene providing
more signal at slightly lower taxonomic levels. Nuclear rRNA genes have great
advantages: they are generally easy to amplify, and they appear to contain more
signals than other genes used for higher-order questions in animal phylogeny.
However, nuclear rRNA genes often exhibit strikingly different rates of evolution,
with attendant problems in alignment, and probably analysis. Markers that are use-
ful at intermediate levels appear to be lacking. Phylogenetic relationships found
using the molecular approach agree well with family, subfamily and genus subdivi-
sions previously defined by classical taxonomy. By comparing the variation pattern
of DNA sequences, information about species relatedness can be obtained. The util-
ity of various genes for reconstructing phylogenetic relationships in parasitiform
mites is not well established (Fenton et al. 2000).
8.2 Species Diversity 177
8.2 Species Diversity
Species identification is the basis for understanding species diversity, phylogenetic

patterns and evolutionary process (Black and Piesman 1994; Verra and Breeuwer
2007). Only correct identifications allow for comparisons between studies and the
repetition or expansion of earlier experiments. In pest species, species identification
is also important in the development of (biological) pest control strategies.
Identification and delimitation of species within the family Tetranychidae have been
an issue of debate for the past few decades. More than 1200 species have been
described under family Tetranychidae, many of which are of great economic impor-
tance. The genus Tetranychus is well studied and includes the common major agri-
cultural pest species having worldwide distribution, T. urticae Koch and Tetranychus
kanzawai Kishida. These mite species constitute a problematic group for systemat-
ics, and considerable experience is required to identify them correctly (Li et al.
2010).
Phytoseiid mites are excellent control agents for suppressing pest mites on a
variety of crops and preventing yield losses. Several commercial businesses mass
rear phytoseiid mite species, then package and sell them to distributors and growers
for augmentative releases into crops. Often the identity of mite species is not veri-
fied before sale; there is always the possibility that different species could contami-
nate the phytoseiid colony during mass rearing leading to the unintended release of
unwanted species into the field and resulting in a failure to apply the correct dose of
the intended species. The phytoseiid mites are small (less than 0.5 mm long) and are
tedious to identify without the help of trained taxonomist. Identification moreover
requires both adult males and females. But identification of single specimen can be
problematic, and even the identification through immature eggs cannot be done.
Hence, a molecular tool to identify these phytoseiid mites at the species level is
urgently needed, at least for some commercially important species. DNA sequences
obtained from nuclear ribosomal RNA genes (18 S, 5.8 S and 28 S rRNA) and their
spaces (ITS-1 and ITS-2) are used routinely. The ITS-2 sequence has also been used
to identify acarine species from the families Eriophyidae, Ixodidae and Tetranychidae.
The predaceous mite, Galendromus (=Metaseiulus, Typhlodromus) occidentalis
(Nesbitt) contains multiple genomes including the genomes of several microbial
symbionts, as well as its own mitochondrial and nuclear genomes. This predaceous
mite has been genetically improved for use in agriculture by developing strains that
lacked the ability to overwinter in diapause or were resistant to multiple pesticides.
From the gene sequences of phytoseiid mite, Phytoseiulus persimilis Athias-
Henriot, scientists selected yolk protein genes for further study and found that these
genes are not completely female specific; this is important because yolk protein
genes are present in both sexes and all life stages suggesting that molecular pesti-
cides based on RNAI sequencing of yolk patterns could be developed for this spe-
cies. In case of P. persimilis, a 12,356 contiguous sequences (contigs) were
assembled with an average size of 935 bp. From these sequences, the putative trans-
lated peptides of 11 contigs were similar in amino acid sequences to other arthropod
Vgs, while 6 were similar to VgRs.
Efficient control of agronomic pests relies on a clear understanding of the taxo-

nomic status of the target species. Since mites are very small in size and their tax-
onomy may sometimes be questionable. Advances in molecular biology provide
data on nucleotide variation that added to more traditional morphological features
that help in establishing reliable criteria to determine pest systematics. Besides pro-
viding solid taxonomic criteria, DNA-based data may help in understanding the
partition of the variability inside a species. In the case of pests, this is a critical
component of any control effort, because the amount of gene flow existing between
populations determines the diffusion risk of some agronomic important genetic
traits (e.g. virulence, acaricide resistance, etc.). Moreover, an increased number of
pest species are being transported to new biogeographic variation of the species, can
also be useful in tracing molecular biology techniques in acarology related to pest
management and have been reported. Different biological situations are approached:
the accurate identification of species with emphasis in distinction of sibling species,
establishment of phylogenies, appraisal of the genetic variability of the target spe-
cies and tracing the biogeographic origin and diffusion of exotic pest problems.
8.3 Molecular Markers
The application of molecular markers to the study of ticks and mites has yielded
new insights into their population. Some species of ticks and some species of spider
mites (Tetranychidae), predatory mites (Phytoseiidae) and worm mites (Eriophyidae)
have been studied at individual, population and species level. Population variations
have also been studied in the important bee parasitic mite, Varroa jacobsoni
Oudemans (Navajas and Fenton 2000; JeyaPrakash and Hoy 2009).
At the individual levels, methods used to study different mites and ticks include
amplified polymorphic length polymorphisms (AFLP), random amplified polymor-
phic DNA (RAPD), restriction fragment length polymorphism (RFLP), to highly
specific microsatellite analysis. These markers work at the population and species
level; additional analyses of specific nuclear or mitochondrial genes have been done
by RFLP or sequencing (Mozer–Koch and Gerson 1998).
In population studies, molecular tools are being used to identify whether two
individuals are from the mating of specific parent and estimating the degree of relat-
edness among individuals and are helpful for the determination of social behaviour,
reproductive success and mating choice. A number of different techniques are avail-
able for identifying genetic differences between organisms (Cruickshank 2002).
Molecular techniques differ in the way they sample within the genome and in the
type of data they generate. A brief information about molecular techniques is given
below.
(a) Restriction fragment length polymorphism (RFLP). All organisms have differ-
ences in their genomic DNA sequence and, therefore, are genotypically distinct.
This difference results in a restriction fragment length polymorphism. The
8.3 Molecular Markers 179
RFLP is used as a major tool to identify the genetic diversity within and between
species.
(b) Random amplified polymorphic DNA (RAPD). The invention of PCR (poly-
merase chain reaction) is a milestone in the development of molecular tech-
niques. PCR results in the selective amplification of a chosen region of a DNA
molecule. Random amplification of DNA with short primer by PCR is a useful
technique in phylogenetics. The important point is the banding pattern seen.
The difference between the genomes of two organisms can be measured with
RAPD. Two closely related organisms would be expected to yield more similar
banding patterns than two organisms that are distant in evolutionary terms.
Moreover, this technique requires only small piece of animal tissue or as the
extracted DNA can be amplified million times using PCR.
(c) Amplified polymorphic length polymorphism (AFLP). AFLP analysis is able to
direct high levels of polymorphism and has high repeatability and speed of
analysis. These markers have a very high diversity index resulting in a limited
number of primer combinations required to screen a whole genome and have
been applied to develop a system for fingerprinting of an organism and for map
expression. The major advantage of AFLP technique is the large number of
polymorphisms that the method generates. Its ability to differentiate individuals
in a population makes the technique useful for paternity analyses, and gene flow
experiments, and also for ‘plant variety registration’. Other advantages of the
AFLP technique are as follows: no sequence information is required, PCR tech-
nique is fast and a high multiplex ratio is possible.
(d) Microsatellites. Simple sequence repeats (SSRs) or short tandem repeats (STRs)
are all different names for the same type of marker. Microsatellites consist of
only 1–6 bp and the whole repetitive regions span less than 150 bp. These mark-
ers appear to be hypervariable, in addition to which their co-dominance and
reproducibility make them ideal for genome mappings as well as for population
genetic studies. Microsatellites owe their variability to an increased rate of
mutation compared to other neutral regions of the DNA. Microsatellites are
used to answer such questions as: From which population does this individual
originate? What are the genetic relationships between individuals? What is the
mating structure of a population?
(e) DNA barcoding. Until now biological specimens were indentified using mor-
phological features, but in most cases an experienced taxonomist is needed. If a
specimen is damaged or is in an immature stage of development, even special-
ists are unable to make identifications. Barcoding solves these problems,
because non-specialists can obtain barcodes from tiny amounts of tissue. DNA
barcoding can serve a dual purpose as a new tool in the taxonomists’ tool box
supplementing knowledge as well as being an innovative device for non-expert
who needs to take a quick identification.
DNA barcoding is based on the premise that a short standardized sequence can
distinguish individuals of a species, because genetic variation between species
exceeds that within species. DNA barcoding datasets are essentially composed of
short DNA sequences from several individuals of a large number of species. Species
identification through barcoding is usually achieved by the retrieval of a short DNA
sequence – the ‘barcode’ – from a standard part of the genome (i.e. a specific gene
region) from the specimen under investigation. The barcode sequence from each
unknown specimen is then compared with a library of reference barcode sequences
derived from individuals of known identity. A specimen is identified as its sequence
closely matches one in the barcode library; otherwise, a new record can lead to a
novel barcode sequence for a given species (i.e. a new holotype or geographical
variant), or it can suggest the existence of a new encountered species (Anderson and
Truemann 2000).
However, with the increasing number of sequences being deposited in public
DNA sequence databases, concerns about the taxonomic misidentification of the
specimens used to obtain these sequences are also growing which threaten the util-
ity of databases themselves. Challenges in the barcoding approach which is increas-
ingly used to identify taxa will be of a taxonomical rather than technical nature.
Among the Acari, misidentification in DNA sequences databases has been reported
for Tetranychidae taxa (Mendonica et al. 2011; Navajas et al. 1992; Navajas et al.
1996).
8.4 Dispersal of Populations and Species
Microsatellites have been used to establish dispersal patterns and gene flow in a
wide range of mite taxa including the mesostigmatid mite, Varroa destructor
(Anderson and Trueman); eriophyid mite, Colomerus vitis (Pagenstecher); and spi-
der mite, Tetranychus turkestani (Ugarov and Nikolskii). In the near future, these
techniques may be applied to an even wider range of taxa like Bdelloidea and
Raphignathoidea – taxa of potential use in biocontrol, but their dispersal abilities
are quite poorly known. These techniques may also help elucidating different dis-
persal patterns and population structure in oribatid mites.
Molecular techniques are proving to be very powerful in issues like tracking
invasive species like Aceria guerreronis Keifer – a worldwide pest of coconuts.
Such studies allow focussed search for potential predators. Similarly through track-
ing invasions of Varroa destructor – parasite of bees, and dispersal studies of palm
mite, Raoiella indica Hirst, around the world might lead to their improved manage-
ment strategies.
8.5 Development of Host Races and Species Link
In literature, many claims and counterclaims of host specificity and host races are
found. Testing such claims is often difficult and very laborious by using standard
methods. In such studies, molecular techniques allow nuclear and often more accu-
rate assessments of separation between ‘host races’ by measuring actual gene flow.
8.7 Molecular Identification of House Dust and Storage Mites 181
Such approaches have shown cryptic species in genus Varroa and significant indica-
tors of host race formation in the parasitic mites, Sarcoptes scabiei DeGeer com-
plex, and in the tick Ixodes uriae White. Some other molecular studies showed a
lack of host specificity in Psoroptes and some Tetranychus spp. (Carew et al. 2009).
8.6 Mite Associates
Many of associations of mites with bacteria, fungi or insects might be accidental

and have little biological significance, but others may provide some real insights.
Another area of interest is the interaction of various microorganisms in the mite
host, for example, the potential for facilitation or competition. The general area of
microorganisms associated with mites should see lots of exciting developments in
the coming years.
8.7 Molecular Identification of House Dust and Storage Mites
Mites are known as causes of allergic diseases. Currently, identification of mites

based on morphology is difficult if only one mite is isolated from a (dust) sample or
when only one gender is found or when the specimen is not intact especially with
the loss of legs. The use of polymerase chain reaction – restriction frequent length
polymorphism (PCR–RFLP) of the ITS-2 gene to complement the morphological
studies for the identification of mites to the species level – has been reported for six
species, namely, Dermatophagoides pteronyssinus, D. farinae, Blomia tropicalis,
Tyrophagus putrescentiae, Aleuroglyphus ovatus and Glycycometus malaysiensis
Fain and Nadchatram. The genomic DNA of the mites was extracted, quantified,
amplified and digested individually with restriction enzymes. Hinf-1 and Ple-1 dif-
ferentiated the restriction patterns of D. pteronyssinus and D. farina. Bfa-1 is
reported useful for the differentiation of G. malaysiensis. PLe-enzyme is reported
useful for the differentiation between T. putrescentiae and A. ovatus. Bfa-1 was use-
ful for the differentiation of G. malaysiensis from the rest of the species. In conclu-
sion, different species of mites can be differentiated using PCR–RFLP of ITS-2
region. PCR–RFLP methods have been found quite useful in identification of house
dust and grain mites, even if complete and intact adult specimens of both sexes of
mites are not available (Wong et al. 2011).
The mitochondrial genome of D. pteronyssinus is a circular DNA molecule of
14, 203 bp; it contains the complete list of 37 genes (13 protein-coding genes, 2
rRNA genes and 22 tRNA genes) usually present in metazoan mitochondrial
genomes. The mitochondrial gene order differs considerably from that of other
Acari mitochondrial genomes.
8.8 enome Sequencing of Two-Spotted Spider Mite

G
(Tetranychus urticae Koch, TSSM)
TSSM has rapid development rates, has relatively small genomes and is easily mul-
tiplied and maintained under laboratory conditions. These characters make TSSM
as an excellent candidate for developing into a chelicerate model. Spider mites are
of special importance as regards their biology, evolution, ecology and genomics and
are of substantial economic importance and have great significance for the biotech-
nology of pest control and energy conservations. TSSM is rated as most resistant
(among 20 pests) to different pesticides, and its chemical control has become very
difficult. TSSM has one of the smallest genomes in arthropods determined so far (90
Mb) – 60 % of the size of the Drosophila genome (Grbic et al. 2011). The genomes
of other chelicerates are much larger (565–7100 Mb), with the unfinished genome
of the tick, Ixodes scapularis, that is estimated as 2100 Mb (Palmer et al. 1994).
Multiple characteristics of the TSSM genome correlate with its smallest size: small
transposable element content and microsatellite density, increased gene density and
holocentric chromosomes. Transposable elements totalled 9.09 Mb, putting T. urti-
cae together with Drosophila pulex and Apis mellifera as arthropods with 10 % or
less of their genomics comprised of transposable elements. The whole genome
sequencing of T. urticae provides the opportunity for a detailed phylogenetic analy-
sis of arthropods – the most diverse group of animals on earth. TSSM genome illus-
trates the specialized life history of the polyphagous herbivorous pest.
8.9 Tick Genomics
Genomes of ticks, Ixodes scapularis Neumann and Rhipicephalus microplus

Canestrini, are reported 2.15 pq (2.1. Gbp) and 7.5 pq (7.1. Gbp), respectively. The
I. scapularis and R. microplus genomes represent a tremendous resource for tick
and tick-borne disease research. Amblyomma americanum Linnaeus genome is esti-
mated to be approximately 1.04 Gbp. The average haploid genome size of the
Argasidae is estimated to be 1.28 Gbp and that of the Ixodidae as 2.67 Gbp. In addi-
tion to providing information on many aspects of tick biology, sequence data may
permit the identification of new targets for vaccines and acaricide research. The
genomes will also facilitate comparative genomic analysis with a plethora of
sequenced organisms, revealing fundamental differences in the genomes of ticks
and insect-borne vectors as well as between the pro- and metastriate ticks (Klompen
et al. 1996; Klompen et al. 2007; Ginsberg and Stafford 2007; Palmer et al. 1994;
Ullmann et al. 2008).
8.10 Conclusions 183
8.10 Conclusions
Molecular characterization can play a role in uncovering the history and estimating
the diversity, distinctiveness and population structure of mites and ticks of eco-
nomic importance. Awareness of the level of genetic diversity and the proper man-
agement of genetic resources are important issues in modern scenario. New markers
deriving from DNA technologies are variable tools to study genetic variability for
conservation purposes. In the near future, the advent of genomes will give an
impressive tool for genetic resources evaluation. The advent of DNA amplification
and methods of automated DNA sequencing has greatly influenced the availability
of applying nucleotide sequences in phylogenetic studies. Sequences from both
nuclear and mitochondrial genomes are used for inferring phylogenetic history of
various taxonomic levels. The 18 S rDNA is a standard marker especially for inter-
generic levels and also detects intraspecific relationships. Species identifications are
possible by using mtDNA, but the infest evolving rDNA spacers, ITS-1 or ITS-2,
are the most frequently used for validation of a species.
Many new molecular tools have been developed in the last 30 years that have
practical application for biological control for species identification, determining
pest areas of origin, studying the efficacy of natural enemy biotypes and determin-
ing the magnitude of nontarget insects and habitat infiltration. These tools can help
improve the efficacy of biological control, reduce risks from nontarget impacts and
enhance understanding of how genetic structures of pests and natural enemy popu-
lations affect pest regulation and stability.
If we can identify the biological pathways mites use to feed on plants, we can
potentially identify chemical and biological methods to disrupt those pathways and
stop the mites from feeding. The genome of T. urticae has been deciphered recently
revealing the presence of families of genes involved in breaking down toxic com-
pounds either in plants poisonous to spider mites or in pesticides (Jamroz et al.
2000; Mitchella 1996). These mites seem to have many genes which may be
involved in breaking down toxic compounds. Scientists have found about three
times more number of genes in TSSM than that are found in other arthropods. These
studies may bring about new insights into the evolution of arthropods, but also offer
opportunities to develop means of crop protection against the spider mite pest.
Spider mites are known to show resistance to different kinds of pesticides, and the
current study of the genome will shed light on the mechanisms present in the mite
to develop fast adaptation and resistance. New genes have also been identified that
are responsible for the production of silk threads in TSSM.
Molecular systematics clearly can make a contribution in the area of the several
aspects of classification of the group, including around monophyly of the four sub-
orders Opilioacarida, Holothyrida, Ixodida and Mesostigmata. But such analysis
has so far been applied almost exclusively to relationships in ticks. Molecular data
certainly contribute, but when nothing is known about organisms except their
DNA. There is no credible reason to give DNA characters greater stature than any
other character. DNA sequence information may serve as a supplement rather than
a required replacement for morphological data.
Further Reading
Anderson DL, Truemann JWH (2000) Varroa jacobsoni (Acari: Varroidae) is more than one spe-
cies. Exp Appl Acarol 24:165–189
Black WC, Piesman J (1994) Phylogeny of hard-, and soft-ticks taxa (Acari: Ixodida) based on
mitochondrial 16 S rDNA sequences. Proc Antn Acad Sci (USA) 91:1034–1038
Carew M, Schifer M, Umina P, Hofmann A (2009) Molecular markers indicate that the wheat curl
mite, Aceria tosichella Keifer may represent a species complex in Australia. Bull Entomol Res
99:479–486
Cruickshank RH (2002) Molecular markers for the phylogenetics of mites and ticks. Syst Appl
Acarol 7:3–14
Dabert M (2006) DNA markers in the phylogenetics of Acari. Bio Lett 43:97–107
Dabert M, Witalinski W, Kazmierski A, Olszanowski Z, Dabert J (2009) Molecular phylogeny of
acariform mites (Acari: Arachnida): strong conflicts between phylogenetic signal and long-
branch attraction artifacts. Mol Phylogenet Evol 56:222–241
Fenton B, Birch ANE, Malloch G, Lanham PG, Breumann RM (2000) Gall mite molecular phy-
logeny and its relationship to the evolution of plant host specificity. Exp Appl Acarol
24:831–861
Ginsberg HS, Stafford K (2007) Management of ticks and tick borne diseases. In: Goodman JL,
Denius DT, Sonenshine DE (eds) Tick borne disease of humans. ASM Press, Washington, DC,
pp 65–85
Grbic M et al (2011) The genome of Tetranychus urticae reveals herbivorous pest populations.
Nature 479:487–495
Guerra–Garcia JM, Espinosa F, Garcia JC (2008) Trends in taxonomy today: an overview about
the main topics in taxonomy. Zool Balt 19:15–49
Jamroz RC, Guerrero FD, Pruet JH, Oehler DD, Miller R (2000) Molecular and biochemical sur-
vey of acaricide resistance mechanisms in larvae from Mexican strains of the southern cattle
tick, Boophilus microplus. J Insect Physiol 46:685–695
JeyaParkash A, Hoy MA (2009) The nuclear genome of the phytoseiid Metaseiulus occidentalis
(Acari: Phytoseiidae) is among the smallest known in arthropods. Exp Appl Acarol
47:263–273
Klompen JS, Black WC, Keirans JE, Oliver JH (1996) Evolution of ticks. Annu Rev Entomol
41:141–161
Klompen H, Lekvishivili M, Black WC (2007) Phylogeny of parasitiform mites (Acari) based on
rRNA. Mol Phylogenet Evol 43:936–951
Li GQ, Xua XF, Zhang KJ, Hong XY (2010) Identification and molecular phylogeny of agricultur-
ally important spider mites (Acari: Tetranychidae) based on mitochondrial and nuclear ribo-
somal DNA sequences, with an emphasis on Tetranychus. Zootaxa 2647:1–10
Mendonica RS, Navia D, Diniz TR, Auger P, Navajas M (2011) A critical review on some closely
related species of Tetranychus sensu stricto (Acari: Tetranychus) in the public DNA sequences
databases. Exp Appl Acarol 55:1–23
Mitchella M (1996) Acaricide resistance: back to basics. Trop Anim Health Prod 28:535–585
Mozer–Koch R, Gerson U (1998) Separation of acarine species by enzymatic, immunological and
molecular methods. Syst Appl Acarol 3:9–18
Further Reading 185
Navajas N, Fenton B (2000) The application of molecular markers in the study of diversity in
acarology: a review. Exp Appl Acarol 24:751–774
Navajas M, Cotton D, Kreiter S, Guttierrez J (1992) Molecular approach in spider mites (Acari:
Tetranychidae): preliminary data ribosomal DNA sequences. Exp Appl Acarol 15:211–218
Navajas M, Guitierrez J, Lagnel J (1996) Mitochondrial cytochrome oxidase I in tetranychid mites:
a comparison between molecular phylogeny and changes of morphological and life history
traits. Bull Entomol Res 86:407–417
Oliver JH (1977) Cytogenetics of mites and ticks. Annu Rev Entomol 22:407–429
Palmer MJ, Bantle JA, Guo X, Fargo WS (1994) Genome size and organization in the ixodid ticks.
Amblyomma americanum. Insect Mol Biol 3:57–62
Ullmann AJ, Stuart JJ, Hill CA (2008) Public Health Resources University of Nebraska, Lincoln
(2008) Public Health Resources
Verra IDR, Breeuwer AJ (2007) Spider mite (Acari: Tetranychidae) mitochondrial COI Phylogeny
reviewed, host plant relationships, phylogeography, reproductive parasites and barcoding. Exp
Wong SF, Chong AL, Mak JW, Tan J, Ho JM (2011) Molecular identification of house-dust and
storage mites. Exp Appl Acarol 55:123–133
Xie L, Eie RR, Zhang KJ, Hong XY (2008) Genetic relationship between the carmine spider mite,
Tetranychus cinnabarinus (Boisduval) and the two-spotted spider mite, Tetranychus urticae
Koch in China based on the DNA COI and rDNA ITS 2 sequences. Zootaxa 726:18–32
Water Mites
9
Abstract
Water mites are brightly coloured, red or orange acarines. They are also known
as Hydracarina, Hydrachnidia or Hydrachnellae. More than 6000 species of
water mites belonging to 40 different families from prostigmatan and
Parasitengonina groups have been reported. Water mites are more thoroughly
studied in European countries and North America, but relatively poorly studied
in Asia, Africa and much of South Africa. Feeding habits and habitats of water
mites and Hydrachnidia diversity are discussed. Parasitic mites and their juve-
niles generally feed on aquatic insects. Water mites are generally exceptionally
sensitive indicators of habitat conditions and the impact of environmental
changes in the freshwater communities. Considering the diversity and great eco-
nomic importance of water mites, some websites covering different types of
information from global as well as regional level are found. Aquatic mites play
an important role in regulating the population of other invertebrates. Larval water
mites of families Erythraeidae, Trombellidae, Trombiculidae, Trombidiidae,
Arrenuridae, Hydryphantidae, Limnesidae and Pionidae are parasitic on adult
stages of different mosquito species. Larvae of many water mite species are pred-
ators of chironomid midges.
Keywords
Hydracarina • Diversity • Predators
Water mites are brightly coloured, red or orange acarines. They are also known as
the Hydracarina, Hydrachnidia or Hydrachnellae and are a group of nine superfami-
lies: Hygrobatoidea, Leberioidea, Arrenuroidea, Sperchontoidea, Hydrachnoidea,
Eylaoidea, Hydryphantoidea, Hydrovolzioidea and Stygthrombidioidea, compris-
ing more than 40 families from the prostigmatan cohort Parasitengonina. The name
‘Parasitengonina’ is in reference to larval parasitism and the typical life cycle that is
the cohesive characteristic defining this taxon (Walter and Proctor 1999). There are

DOI 10.1007/978-981-10-1594-6_9
188 9 Water Mites
more than 6000 species of water mites described from the world. This number
might be quite low as water mites have not been well studied in Asia, Africa and
much of South Africa. These creatures have been more thoroughly studied in differ-
ent European countries and North America.
9.1 Hydrachnidia Diversity
Galumnidae, Hydrachnidae, Limnocharidae, Hydryphantidae(Protzia),

Hydrodromidae(Hydrodroma), Rhynchohydracaridae (Clathrosperchon,
Rhynchohydracarus and Santiagocarina), Oxidae (Frontipoda), Torrenticolidae
(Monoatractides, Neotractides, Torrenticola), Limnesidae (Limnesia,
Neotorrenticola, Protolimnesia), Hygrobatidae (Atractides, Hygrobates),
Unioncolidae (Neumannia, Unioncola), Aturidae (Aturus, Axonopsis,
Frontipodopsis), Medeopsidae (Medeopsis) and Arrenuridae (Arrenurus) are impor-
tant families containing water mites (Smith and Cook 2001).
The Halacaridae is a group of entirely aquatic mites from the suborder
Prostigmata, of the order Trombidiformes, and the superorder Acariformes. Majority
of the members of this family are marine, but there are many brackish and freshwa-
ter taxa as well (Sabatino et al. 2002).
9.2 Feeding Habits
As adults, most aquatic mites are predatory feeding on zooplankton and insect lar-
vae. During feeding, mites grasp their prey and use specialized piercing mouthparts
to puncture their food and suck juices out. Some of the predatory mite species,
however, feed on detritus and plants; others are algivorous or parasitic. Larval mites
are parasitic but do not usually cause the mortality of their hosts. However, when
they are found in large numbers feeding on their hosts, then they can cause some
damage to their health (Lanciam and Boyd 1980).
Upon hatching from the eggs, six-legged larvae swim around looking for an
aquatic insect host. Water mite larvae are parasitic, so they need a host to live on.
Dragonfly naiads, damselfly naiads, fly larvae (including mosquitoes), true bugs,
chironomids and stonefly nymphs are their hosts depending upon the species
involved. In some cases, one insect may carry up to 20 mite larvae (Smith 1988).
Water mite larvae munch on their hosts while they carry them. When sufficient
growth of mite larva occurs, then its exoskeleton (outer shell) becomes bag-like.
The larva stays inside the bag and transforms to a nymph which then leaves their
host. Water mite nymphs eat the same food as the adults do including small insects,
aquatic worms and other mite species (even of their own kind). Nymphs then con-
tinue to eat and grow until they are ready to go into their next nymphal stage. Each
nymph finds some algae, an aquatic plant or another object to attach to; thereafter,
the nymph shrinks inside another ‘bag’ for about a week while it changes to an
adult. In the larval stage when attached to its host, the mite may face sometimes a
9.4 Effect of Water Pollution 189
problem if the host leaves the water. If this happens, then the mite can pop out of its
bag, or hold on, hoping its host to return to the water. If the host does not return to
the water, then the larva drops off the land and dies (Efford 1963).
Typically the larvae are ectoparasitic on adult insect hosts, and the nymphs and
adults are free-living voracious predators of immature aquatic insects and micro-
crustaceans. Larval water mites of the genus Arrenurus are parasitic on adult mos-
quitoes. When larval mites contact a mosquito pupa, they cling to it and wait for
emergence of the adult host. The mites then transfer to the adult, attach and begin
engorgement. As the mosquito returns to water to oviposit, the mites detach (Smith
1998).
9.3 Habitat
Both the adult and young stages of the water mites live beneath the water surface.
They are found in a wide range of freshwater bodies and free tree holes to hot
springs but prefer to live in still waters such as ponds and slow-moving rivers. Water
mites are most abundant among aquatic plants and in shallow areas of ponds, lakes,
swamps, marshes and bogs. Some water mites can even thrive in the riffles of fast-
flowing streams. They live in deep lakes and in torrential waterfalls, and some water
mite species also invade oceans. As far as known, nearly all species of the true
freshwater mites (Hydrachnidia) are bound to host insect in a parasitic/phoretic
relation during their larval stage. From a qualitative point of view, the most impor-
tant host insect group are the Diptera (chironomids and mosquitoes), but insects of
nearly all orders with aquatic instars are parasitized with the obvious exception of
Ephemeroptera.
9.4 Effect of Water Pollution
Parasitic mites and their juveniles generally feed on aquatic insects, although some
feed on mussels. Parasitism by mites is usually non-lethal, although in high num-
bers they can affect the development of their hosts. However, salinity of water
changes dramatically as a pond dries, and unpredictable rainfall patterns often dic-
tate stochastic variation in pond duration. Water mites breathe by absorbing dis-
solved oxygen all over the body surface; they can survive in dissolved concentrations
as low as 1 part per million (ppm), making them potentially well suited to live in
waters polluted by nutrients. As a group, water mites can exist in a wide range of
pollution levels although individual species range from sensitive to somewhat toler-
ant of pollution. Water mites are generally exceptionally sensitive indicators of
habitat conditions and the impact of environmental changes in the freshwater com-
munities (Sabatino et al. 2002; Proctor and Pritchard 1989).
In different studies from the European continent based on physico-chemical pol-
lution ecology, composition of water mite communities has been found as an excel-
lent indicator of habitat quality. Water mite diversity is dramatically reduced in
190 9 Water Mites
habitats that have been degraded by chemical pollution or physical disturbance.

Some studies regarding the tolerances of water mites to environmental variables
revealed different mechanisms responsible for observed changes in species popula-
tions in stressed habitats.
9.5 Potential as Indicators of Environmental Safety
Species of water mites are specialized to exploit narrow ranges of physical and
chemical regimes as well as the particular biological attributes (including physico-
chemical constraints) of the organisms they parasitize and prey upon. Water mites
are the exceptionally sensitive indicators of habitat conditions and the impact of
environmental changes in the freshwater communities. They have highly specific
habitat and host associations and that many of them have limited distribution.
Mussels, crayfishes, stoneflies, amphibians and fish are among the other most
important inhabitants of freshwater. As one of the dominant groups of arthropods in
many freshwater habitats, water mites have integral roles in the organization and
regulation of freshwater communities. Their life cycle including active parasitic and
predatory phases tightly links them to other aquatic invertebrates. Water mite larvae
are ectoparasitic on adult insect hosts, and the nymphs and adults are free-living
voracious predators of immature aquatic mites and some micro-crustaceans. Larval
mites regularly parasitize adults in natural populations of aquatic insects with sub-
stantial impact on the size and structure of host populations in many habitats.
Deutonymphs and adults of free-living species prey on a wide range of aquatic
organisms (Cook 1974, 1986).
9.6 Websites on Water Mites
Considering the importance of water mites in managing several parasitic aquatic

insects, some organizations have created special websites for exchanging and shar-
ing scientific and popular information on water mites.
9.6.1 Water Mites of North America
(http://watermites.uark.edu/watermite_aboutproject.html)
This project has been promoted jointly by the University of Arkansas, USA, and
the Canadian National Collection of Insects and Arachnids (CNC), to promote edu-
cation and research of water mites and other freshwater arachnids. Studies on mites
of North America are, however, the principal focus. This project fosters collabora-
tive international education and research as well. CNC keeps historical record of
knowledge of Hydrachnidiae of this continent and presently has over two million
curated water mites. A georeferenced database of over 10, 000 line drawings
and SEM photographs of many of the species has been digitized. A library, an
9.7 Life Cycle 191
excellent photo gallery and bibliography of publications (more than 4000), is also
maintained. The CNC even provides training to new generation of research
professionals.
9.6.2 Hydracarina.Org
(http://hydracarina.org/)
This website provides links and visual key of water mites based on their size,
photomicrography and localities found, and salient references for each are given
nicely. In this website, option ‘Search’, is also included for finding out useful ‘infor-
mations about these mites’.
9.6.3 K
ey to Families and Subfamilies of Water Mites
(Hydracarina) in Australia
(http://keys.lucidcentral.org/keys/lwrrdc/public/Aquatics/aghydra/html/about.htm)
Key to families and subfamilies of water mites (Hydracarina) in Australia is
given along with the information about habitats of their members (Harvey 1998).
9.6.4 Acari1: European Water Mites Research
(www.watermite.org/misc/book_acari_1.html)
This website focuses information on water mites of the European region. This
site provides information regarding Pro Acarologia Basiliensis (PAB) foundation
(Stiftung PAB), its members, European fauna, publications, new taxa, collections,
photo gallery and many useful links.
9.6.4.1 Hydracarina
(http://www.tolweb.org/Hydracarina)
This site also provides good information about introductory account, character-
istics, life history and ecological information and phylogenetic relationships of
water mites.
9.7 Life Cycle
Eggs are laid under water either on the surface of substrates such as stores or vegeta-
tion or inside living plant tissues (Hydrachnidae, Hydrachna) or in the tissues of
freshwater sponges and mussels (Unioncolidae, Unioncola; Pionidae, Najadicola).
In the less derived groups, the larvae swim or crawl to the waters’ surface after
hatching from eggs and break through the surface film. They then run about on the
upper surface of the water or on emergent aquatic plants, seeking insect hosts to
192 9 Water Mites
parasitize. In more derived taxa, the larvae remain under water where they search
for pre-imaginal larvae (e.g. of odonates) or pupae (e.g. of dipterans). Larvae of
‘aerial’ group typically begin to parasitize a host immediately upon encounter; in
contrast, ‘aquatic’ larvae merely attend the pre-adult host and only begin parasitism
after the adult has emerged from the last larval or pupal skin (Smith and Cook
2001). The larval period varies from a few days to several months, depending on the
taxon. Engorged larvae drop from the host back into the water. Because hosts of
water mites larvae are invariably winged adult insects (exceptions are in the
Hemiptera, where nymphal hosts may be parasitized), mites are often carried far
from their natal waterbodies.
Parasitism serves not only a nutritional function for hydracarines, but it is the
major means of dispersal and colonization of new habitats. Dispersal via insects is
a highly influential life history characteristic for the pond mites. Once back in the
water, the engorged larva transforms into a calyptostatic protonymph (also called a
nymphochrysalis) that is a pupa-like stage inside the integument of the larva. Within
the protonymphal cuticle, the next stage – the deutonymphs – develops. The
deutonymph encloses as an active swimming or crawling predator of all stages of
aquatic insects, crustaceans and other mites. A well-flattened deutonymph trans-
forms into a calyptostatic tritonymph (also called a teleiochrysalis). The adult that
emerges is also predatory and typically feeds on prey similar to that chosen by the
deutonymph. The adults are also often extremely sexually dimorphic, particularly of
those taxa in which the males transfer sperms directly to the female.
One of the most striking things about water mites is their often brilliant coloura-
tion. Scarlet, orange, red and yellow mites are common and are extremely visible
against a background of green water plants. These bright mites are often very unpal-
atable to fish, presumably due to noxious secretions from their skin glands. This
aposematic connection between redness and distastefulness might have evolved as
an anti-fish predation mechanism. In terrestrial parasitengonines both the bright
colour and foul taste are also present.
9.8 Role of Water Mites in Food Chain
Aquatic mites play an important role in regulating the population of other inverte-
brates. Since hosts they parasitize are often brilliantly coloured, it makes them very
much visible among green and brown aquatic plants. Brightly coloured mites have
been reported to be distasteful to the fishes. Many mites are able to colonize new
bodies of water; they do this by becoming parasitic on other aquatic insects that are
in their larval or pupal stages and waiting until they become terrestrial adults. They
then travel around to new aquatic habitats with their unaware winged hosts (Smith
1988).
9.9 Biological Control of Insect Pests 193
9.9 Biological Control of Insect Pests
Larvae of water mites have been found to play an important role in regulating the
population of different aquatic insects (including mosquitoes). Although very
detailed studies on the utility of water mites are not available in literature, however,
some information found on their role both as parasites and predators of different
insects is given briefly (Proctor and Pritchard 1989).
9.9.1 Parasites
Nearly all species of the true freshwater mites (Hydrachnidia) are bound to host
insects. Important insect groups are the chironomids and mosquitoes (Diptera), but
insects of nearly all orders in a parasitic/phoretic relation during their larval stage.
From a qualitative point of view, the most having aquatic instars are parasitized by
water mites. Hydrachnidia larvae parasitized members of Trichoptera families:
Phylopotamidae, Psychomiidae, Polycentropodidae, Hydropsychidae,
Glossosomatidae, Hydroptilidae, Limnephilidae, Lepidostomatidae and
Leptoceridae (Smith 1988). In most cases, mite larvae were found attached to adults,
but in Limnephilidae, mites were also reported attached to the larvae. Parasitism by
mites is usually non-lethal, but mite infestation can result in adverse effects on the
health of host insects. Larvae of water mite, Hydrachna conjecta, are parasitic on
corixid bugs (Sigara striata). Hygrobates species are commonly found as parasites
of chironomid midges (Proctor and Pritchard 1989).
9.9.2 Parasites of Mosquitoes
Mosquitoes belong to family Culicidae (order Diptera). Some species of genera

Aedes and Anopheles are of good importance in public health since they serve as
vectors of several diseases like malaria, dengue and yellow fever. Larval water mites
of families Erythraeidae, Johnstonianidae, Trombellidae, Trombiculidae and
Trombidiidae (all of Trombidia) and Arrenuridae, Hydrodromidae, Hydryphantidae,
Limnesidae and Pionidae (all of Hydrachnidia) are parasitic on adult stages of dif-
ferent mosquito species (Smith 1988).
Members of family Ascidae (Parasitiformes) feeds on eggs of mosquitoes.
Trombiculid and limnesid mites also feed on eggs of mosquitoes. Some species of
Hydrophantidae and Pionidae feeds on larvae of Aedes spp., but limnesid mites,
Limnesia jamurensis Oudemans, have been reported feeding on larvae of both
Anopheles and Culex species of mosquitoes.
Arrenurus spp. (Arrenuridae); Hydroma spp. (Hydrodromidae); Euthyas,
Hydrophantes and Thyas species (Hydrophantidae); and Limnesia spp., (Limnesidae)
are commonly found parasitic on different mosquito species. Piona spp. are known
as voracious predators of mosquito larvae. Parasitism by different Arrenurus spp.
seems to have substantial effect in reducing populations of Mansonia and Anopheles.
194 9 Water Mites
However, detailed studies on importance of water mites as biological control of

mosquitoes are urgently needed.
9.9.3 Predators
Larvae of many water mite species prey on eggs or larvae of chironomids (midges).
Adults of water mite, Thyas barbigera Viets, feed on the eggs of Aedes mosquitoes,
while their larvae are parasitic on adults. Adults of water mite, Hydrachna conjecta
Koenike, are egg predators of corixid bugs (Sigara striata). Predation by adults of
water mites, Hygrobates nigromaculatus Lebert and H. trigonicus Koenike, on dif-
ferent larval stages of chironomids resulted significantly in reduction of their popu-
lation. For adults and nymphs of water mite, Hydrodroma despiciens, chironomid
eggs are found as primary food. One female mite has been reported to consume 235
eggs in just 12 h (Proctor and Pritchard 1989).
9.10 Predators of Water Mites
Several aquatic insects, crustaceans (such as water fleas and scuds), hydras and
small fishes are sometimes found as significant predators of water mite adults.
9.11 Conclusions
In many freshwater habitats, water mites (Hydracarina) are one of the most domi-
nant groups of arthropods and play integral role in the regulation of freshwater com-
munities. Water mites are the most species-rich group of arachnids occurring in
standing and running freshwater habitats; more than 6000 species of water mites are
reported around the world. But their ecology has been neglected as compared to
quantum of studies made regarding other groups of freshwater arthropods. Water
mites are generally exceptionally sensitive indicators of habitat conditions and the
impact of environmental changes in the freshwater communities. Nearly all species
of the freshwater mites (Hydrachnidia) are found on host insects in a parasitic/pho-
retic relation during their larval stage. Larval water mites parasitize hosts from a
number of insect orders including Diptera, Trichoptera, Plecoptera, Coleoptera,
Hemiptera and Odonata. Some species of water mites have been reported to feed on
eggs of mosquitoes (Aedes spp.) and as parasites or predaceous on mosquitoes,
corixid bugs and chironomid midges. However, parasitism by mites is usually non-
lethal but mite infestation can result in the adverse effect on the health of parasitized
insects.
Further Reading 195
Further Reading
Cook DR (1974) Water mite genera and subgenera. Mem Am Entomol Inst 21:1–860
CooK DR (1986) Water mites from Australia. Mem Am Entomol Inst 40:1–558
Efford I (1963) The parasitic ecology of some water mites. J Anim Ecol 32:141–156
Harvey MS (1998) The Australian water mites: a guide to families and genera. CSIRO, Melbourne.
ISBN 0-643-06301-3
Lanciam CA, Boyd JM (1980) Demonstrative parasitic water mite-induced mortality in natural
host populations. Parasitology 81:465–475
Proctor H, Pritchard G (1989) Neglected predators: water mites (Acari: Parasitengona:
Hydrachnellae). J NAm Benthol Soc 8:100–111
Sabatino AD, Martin P, Gerecke R, Cicolani B (2002) Hydrachnidia (Water Mites). In: Rundle SD,
Robertson AL (eds) Freshwater meiofauna-biology and ecology. Schmidaraya, Backhuys
Publishers, Leiden, pp 105–133
Smith BP (1988) Host-parasite interaction and impact of larval mites on insects. Annu Rev
Entomol 33:487–507
Smith BP (1998) Loss of larval parasitism in parasitengona mites. Exp Appl Acarol 22:187–199
Smith IM, Cook DR (2001) Water mites. In: Thorp JH, Couch, AP (eds) Ecology and classification
of North American fresh water invertebrates, 2nd edn. Academic Press, San Diego, pp 523–
592, 950 pp (Hardcover ISBN: 0126906475)
Walter DE, Proctor HC (1999) Mites: ecology, evolution and behaviour. Univ. New South Wales
Press, Sydney
1. https://www.youtube.com/watch?v=2iYVilhnNck Big Red Water Mites, Trombidiformes?

2. https://www.youtube.com/watch?v=Bjn8jXfGimw Water Mite
3. https://www.youtube.com/watch?v=IOW4TcftfN4 Living water mite
4. http://www.mdfrc.org.au/bugguide/display.asp?type=2&class=16 Water Mites
5. https://www.youtube.com/watch?v=ufHLww1eglM Water mite – Hydrachna
Soil Mites
10
Abstract
A soil is not a pile of dirt but is a transformer – a body that organizes raw materi-
als into tissues that become the mother to all organic life. In the soil ecosystem,
different species of viruses, bacteria, fungi, algae, protozoa, mites, nematodes
and worms are found. Both Parasitiformes and Acariformes mites are found in
different types of soils and cropping conditions. Parasitiformes mites found in
soil include both Ixodida (ticks) and Mesostigmata; Uropodida mites, Gamasina
mites; and Acariformes mites belong to Astigmata, Oribatida, Endeostigmata and
Prostigmata. Gamasina mites. They are important predators of nematodes and are
inhabitants of ground litter in top layer of soil. The presence of several genera of
these mites is considered as good indicators of habitat and soil condition. More
than 10,300 species of oribatid mites in 177 different families are common inhab-
itants of soil. These mites are the world’s most numerous arthropods living in
soil, and their diversity in forest soil can reach hundreds of thousands of individu-
als per square metre. Oribatid mites are considered as important component of
soil decomposers; and their abundance, species composition and diversity in a
particular habitat serve as good indicators of soil health. Some species of oribatid
mite family Oribatulidae are quantitatively the most significant as intermediate
hosts. Oribatid mites, Scheloribates laevigatus, are the most frequently found
with natural cysticercoid infestation of Moniezia (M. benedeni and M. expansa).
Keywords
Ecosystem • Oribatids • Decomposers • Intermediate hosts
A soil is not a pile of dirt but is a transformer – a body that organizes raw materials
into tissues that become the mother to all organic life. When we are standing on the
ground, we are really standing on the rooftop of another world. Living in the soil are
plant roots, viruses, bacteria, fungi, algae, protozoa, mites, nematodes, worms, ants,
maggots and larger animals. Indeed the volume of living organisms below ground is

DOI 10.1007/978-981-10-1594-6_10
198 10 Soil Mites
often far greater than that above ground. Together with climate these organisms are
responsible for the decay of organic matter and cycling of both macro- and micro-
nutrients back into forms that plants can use. Agricultural practices that include crop
rotations and tillage affect the establishment, growth and nutrient content of the
crops that grow (Brown 1978; Dindal 1990; Emmanuel et al. 1985).
In the soil ecosystem, both Parasitiformes and Acariformes mites are found in
different types of soils and cropping conditions. Parasitiformes mites include both
Ixodida (ticks) and Mesostigmata which are comparatively larger and often with
discrete shields. Uropodida mites (slow-moving fungus feeders) and Gamasina
mites (actively moving predaceous) are commonly found in soil (Marshall et al.
1987; Ghilyarov and Bregetova 1997; Evans 1992).
Acariformes mites belonging to Astigmata, Oribatida, Endeostigmata and
Prostigmata are commonly found with ephemeral habitats and have three nymphal
instars. The second nymphal instar also known as deutonymph has no mouth, but is
sclerotized and is highly modified for dispersal. Among the Parasitiformes, mostly
the members are brown in shade except Rhodacaridae mites that are bright pink in
colour. Gamasina mites are common inhabitants of the ground litter in top layers of
soil and are negatively affected by high ant activity. Almost all Gamasina are impor-
tant predators of nematodes. Several genera are considered good indicators of habi-
tat and soil condition. In the coastal meadows, rich diversity of gamasid mites is
found. Members of families Zerconidae, Veigaiidae and Parasitidae are commonly
found feeding on nematodes. Macrocheles species feed on small worms and putre-
fying soil organisms. Many gamasid mites are facultative carrion eaters; some are
found as ectoparasites on beetles. Rhodacarus and Zircon species are found in
deeper soil layers. Pergamasus, Parasitus, and Hypoaspis are important genera of
gamasid mites found in different soil types. Uropodina mites are more abundant in
high-density mounds compared to low-density mounds or meadow soil (Diaz et al.
2000; Gulati and Mathur 1995; Evans 1992; Kuhnelt 1961).
10.1 Oribatid Diversity
Oribatid mites commonly known as beetle mites, armoured mites or moss mites
comprise more than 10,300 described species in 177 different families belonging to
superorder Acariformes, order Sarcoptiformes and suborder Oribatida
(=Cryptostigmata). They are the worlds’ most numerous arthropods living in soil,
and their diversity in forest soil can reach hundreds of thousands of individuals per
square metre (ER6). Oribatids comprise an important component of soil decompos-
ers; their abundance, species composition and diversity in a particular habitat serve
as good indicators of soil health. They are colourless or may have yellow to red
pigment because of melanization, measure from 300 to 700 μm in length and are
mostly saprophagous or mycophagous. In majority of oribatid mites, cuticle is hard-
ened and bears several dorsal or ventral shields. They contribute to soil organic
matter decomposition, nutrient recycling and soil formation (Dindal 1990; Marshall
et al 1987; Evans 1992; Marshall et al. 1987). Detailed account of oribatid mites and
10.2 Distribution 199
their role in soil formation are briefly given below. Important oribatid mite families
along with important genera/species reported under each in soil ecosystem are given
below:
–– Brachychthoniidae (Brachychthonius bimaculatus Willmann)

–– Cosmochthoniidae (Cosmochthonius sp.)
–– Camisiidae (Camisia biurus (Koch) and Camisia horrida (Hermann)
–– Trhypochthoniidae (Mucronothrus nasalis Willmann)
–– Gymnodamaeidae (Gymnodamaeus sp.)
–– Damaeidae (Belba sp.)
–– Eremaeidae (Eremaeus occidentalis Behan-Pelletier)
–– Liacaridae (Dorycranosus spp.)
–– Carabodidae (Carabodes spp.)
–– Oppiidae (Oppia spp. and Oppiella spp.)
–– Scutoverticidae (Exochocepheus eremitus)
–– Scheloribatidae (Scheloribates pallidulus Koch)
–– Oribatulidae (Oribatula spp. and Zygoribatula spp.)
–– Ceratozetidae (Ceratozetes cuspidatus Jacot)
–– Mycobatidae (Mycobates altus)
–– Oribatellidae (Oribatella sp.)
10.2 Distribution
Oribatid mites occur throughout the world and are even active in the arctic/antarctic
regions. They occur in salt and freshwater, caves, rodent burrows, bird nests, trees
and even in human dwellings. Mite species richness and densities decrease in soil
under significant grazing pressure. Heavy grazing and poor pasture conditions result
in substantial reductions in the abundance of mites and the richness of species
(Evans 1992; St. John et al. 2001; Thomas and MacLean 1988; Tomlin and Miller
1987).
The oribatid mite fauna of temperate and arctic regions differs. Oribatid fauna of
temperate and arctic regions is found throughout the soil profile, in surface litter or
low-growing herbs and shrubs on the trunk and in branches and foliage of the trees.
In contrast, life forms at northern latitudes are primarily associated with the soil
surface; litter and moss cover, deep soil forms and arboreal species are absent.
Members of superfamilies Phthiracaroidea and Euphthiracaroidea are primarily
associated with decaying wood. Mites are the most abundant group of arthropods
within moist soil and litter habitats including deserts and arctic areas. Normally
from 10,000 to 500,000, mites may be found per square metre of soil in temperate
and tropical sites. In temperate regions, Uropodina are generally lower than
Gamasina, but in the tropics Uropodina abundance can rival that of oribatid mites.
In general Uropodina are strongly associated with soils of high organic matter.
Oribatid mites are generally very common in relatively stable habitats and usu-
ally form the bulk of mite biomass in forest ecosystems. In forests, Mesostigmata
200 10 Soil Mites
forms 20 % of total soil mite fauna found, but in open habitats it is less abundant.
However, many oribatid mites are generally very common in stable habitats, but
they are quite sensitive to soil disturbances in the conventionally tilled agricultural
fields. Prostigmata and Endeostigmata mites may form about 40–50 % of the total
number of mites found in soil in deserts and grassland ecosystems, but this ratio
drops sharply in temperate forests (less than 10 %). As in temperate areas, however,
soils low in available moisture and/or organic matter support a greater diversity and
density of actinedid than oribatid mites.
10.3 Life History and Development
Oribatid mites have six embryonic instars: egg, an inactive prelarva, active larvae,
protonymph, deutonymph, tritonymph and adult. In cold climates, their life cycle
can stretch up to 7 years (usually 1–2 years). Oribatid mites are usually character-
ized by low fecundity, long immature and adult life spans and a low capacity for
increase despite the fact that population sizes are very high and generally rather
stable (as compared to insect populations). These characteristics should also be con-
sidered as parameters to be used in the context of ecological indicators. Oribatid
mites in temperate and cold habitats exhibit extensive supercooling ability; imma-
ture instars are at least as cold hardy as adults and can overwinter successfully in
mixed populations. Females lay a few eggs; many species are obligatorily partheno-
genetic, with no males recorded. In contrast to many other micro-arthropods, oriba-
tid mites reproduce slowly and are not capable of fast population growth. Oribatida
are usually restricted to relatively stable environments, in contrast to more opportu-
nistic groups such as collembolans. Oribatids in general have a trait similar to so-
called K-selected species (Brown 1978: Dindal 1976; Kuhnelt 1961).
10.4 Feeding Habits
Oribatid mites feed on fungi and algae and consume decomposing plant matter and
occasionally dead collembolans. Some oribatids feed on live nematodes. Feeding
habits of many groups of oribatid mites are still unknown (Haq 1994; Labrun and
Streelan 1995). Oribatid mites in general are classified into six feeding groups.
(i) Macrophytophagous. Feeding on higher plant material

(ii) Microphytophagous. Feeding on microflora
(iii) Panphytophagous. Unspecialized plant feeders
(iv) Zoophagous. Feeding on living animal material
(v) Necrophagous. Feeding on carrion
(vi) Coprophagous. Feeding on faecal material
Feeding habits of different oribatid mites seem to have relationships with the size
of their chelicerae. In general, macrophytophagous and panphytophagous mites
10.6 Nematophagous Mites 201
have larger chelicerae in comparison with those of microphytophagous and frag-

ment feeders. Some species are also recognized pollen feeders. In general, different
species of the same family tend to have similar feeding habits. Because of their dif-
ferent feeding habits, oribatids have successfully invaded all compartments of the
biosphere. In addition to the upper layer of the soil, several other habitats are colo-
nized by oribatid mites: mineral soil (with endemic species living in caves), aquatic
ecosystems (freshwater and seashores) and terrestrial habitats (canopy foliage,
barks, feeding epiphytic and epilithic vegetation in rocks and cliffs). Small habitats
including dung, bird nests, mosses, fungal mycelia, mushrooms, the inside of conif-
erous needles, food products, etc. are among the variety of microhabitats in temper-
ate forest ecosystems found favourites for oribatid mites. The oribatid group thus is
ubiquitous, showing adaptive radiation throughout all niches containing enough
organic matter content. They are often the dominant arthropod group, particularly in
ecological systems where decaying organic matter is predominantly present (Norton
1985; Thomas and MacLean 1988).
10.5 Population Dynamics
Most marked fluctuations in population of oribatid mites are found depending upon
prevailing climatic and edaphic conditions and a number of other factors. However,
there is little fluctuation throughout the year in gamasids and other acarines which
are found in very low numbers in soil. In temperate conditions, peak population of
oribatid mites is found during autumn and winter months, but their population show
a marked decline during mid-summer. However, in tropical conditions peak popula-
tion of oribatid mites is found during rainy season, and a minimum population is
found during early summer. In general, moisture content and temperature of the soil
are the most important limiting factors, but these factors act in conjunction with
rainfall, vegetational growth and extent of litter decomposition (Winchester et al.
2000).
10.6 Nematophagous Mites
Acari belonging to both the superorders Parasitiformes and Acariformes contains

mite families having many nematophagous mite species. Some studies mainly lim-
ited to pot experiments have reported some mite species as potential biocontrol
agents of root-knot and fungus-feeding nematodes. But the soil mites because of
their cryptic habitat and different culturing techniques have received less attention
of researchers. Representatives of oribatid genera Fuscozetes, Galumna, Northrus
and Pargalumna have been reported to feed on free-living nematodes (Chiloplacus
spp. and Rhabditis spp.). In general nematode-feeding mites are abundantly avail-
able and act as potential biocontrol agents of plant parasitic nematodes (Rockett
1980).
202 10 Soil Mites
10.7 Exposure to Predators
Oribatid mites usually have strong exoskeleton, hardened by sclerotization as in

other mites and also by mineralization. Adult oribatid mites experience relatively
little predation because of their strong exoskeleton, but certain ant species, beetles
and small salamanders, have been reported feeding on these mites specifically. For
example, myrmecine ants are specialized predators of oribatid mites.
10.8 Effect of Drought/Burning
Climate change resulting in frequent summer droughts will probably decrease the
abundance and diversity of oribatids and other soil mite fauna of forests. Oribatida
communities seem to be slow, and disturbances on a large spatial scale may affect
the resistance of soil ecosystems, as disturbances may cause permanent changes in
community composition and moisture conditions may primarily result in shifts in
community structure. These mites, however, protect themselves against desiccation
or flooding of the soil as well as against predators such as staphylinid beetles and
myrmecine ants. Long-lived protonymphs in most species are the overwintering
stage with deuto- and tritonymphs developing in spring. High mortality of these
nymphal stages is probably a factor contributing to the decline in their numbers dur-
ing summer. Maximum population of adults has been found during summer
season.
10.9 Intermediate Hosts of Tapeworms
Members of the oribatid mite family Oribatulidae are quantitatively the most sig-
nificant as intermediate hosts with 35 species. The Galumnidae with 22 species and
Ceratozetidae with 15 species are of secondary and tertiary importance only.
Scheloribates laevigatus (Koch) of family Oribatulidae is the most frequent species
acting as intermediate host of 14 species of anoplocephalids. This species is most
frequently found with natural cysticercoid infestation of Moniezia (M. benedeni and
M. expansa). Liacaridae, Damaeidae, Haplozetidae, Oribatelidae, Epilohmaniidae,
Carabodidae, Opiidae and Nohridae are other oribatid families found with minor
frequency. For tapeworm infestations in ruminants, the vertical migration of oriba-
tids appears to be of great importance. The most significant factors inducing vertical
migration are temperature and soil moisture.
10.10 Effect of Pesticides and Industrial Wastes
However, detailed studies on the effect of pesticides and industrial effluents/pollut-

ants on oribatid mite population are not available. But in literature, chlordane, DDT
and diazinon insecticides have often been reported to reduce the oribatid mite
10.12 Economic Importance of Oribatid Mites 203
populations in different ecosystems. Higher soil applications of carbaryl, disulfo-

ton, fenitrothion and methoxychlor also resulted in reduced mite populations. Many
other modern insecticides when applied through soil applications have also been
reported adversely affecting oribatid mite populations. Soil applications of fungi-
cides like micronized sulphur, D-D mixture and methyl bromide used for fungus
control proved very toxic to soil mites and other soil arthropods (Banerjee 1984;
Kuhnelt 1961).
Waste oils strongly reduced oribatid mites and Collembola populations. Effluents
discharged from fertilizer factories have also been reported in many studies to
decrease oribatid mites and other soil arthropods. In some studies, adverse effect of
radioactive wastes against oribatid mites is also reported. Adverse effect of applica-
tion of municipal wastes in the fields against soil mites has also been reported
(Veeresh et al. 1991).
10.11 Effect of Fertilizers
In many studies, abundance of soil mites and other soil arthropods is reported as a
result of various nitrogenous, phosphatic and potash applications, but some con-
trasting studies regarding negative effects of fertilizers against oribatid mites are
also found. Application of lime is also reported in many studies to decrease the
oribatid mite populations. But application of farmyard manure (FYM) is reported to
increase population of oribatid and other soil mites (Diaz et al. 2000; Tomlin and
Miller 1987).
10.12 Economic Importance of Oribatid Mites
Oribatid mites influence decomposition and soil structure by comminution of

organic matter and producing faecal pellets, which provide a large surface area for
decomposition, and are in turn an integral component of soil structure. In northern
soils of Canada, nutrient recycling and decomposition are particularly important as
large invertebrates such as earthworms, myriapods and some insects are absent or
less frequently found. The soil-dwelling mites contribute to litter decomposition,
soil formation and nutrient recycling. Oribatid mites disperse bacteria and fungi,
both externally on their body surfaces and by feeding on spores that survive passage
through their alimentary tracts. Many oribatid mite species sequester calcium and
other minerals in their thickened cuticle. Their bodies may form important ‘sinks’
for nutrients, especially in nutrient-rich environments such as peatlands (Dindal
1976; Haq 1994; Brown 1978).
Presence of oribatid mites is a useful indicator of good soil health, so the abun-
dance and diversity of these mites is greater in soils with perennial grass cover than
in soils with bare ground and annual grass cover (Banerjee 1984; Brown 1978). Soil
mites feed on fungi and nematodes and are extremely important in maintaining soil
health and fertility. Mites are among the most important predators in the soil
204 10 Soil Mites
ecosystem. Several oribatids have been reported to feed on parasitic nematodes and
play a role in managing their population (ER7).
Comminution The breaking up of larger organic units such as dead leaves and
wood into smaller pieces is largely done by oribatid and astigmatid mites. The other
groups of soil mites are mainly predaceous in nature. Comminution greatly increases
the surface areas on which bacteria can complete the actual process of decomposi-
tion. Through their feeding activity and production of faecal pellets, oribatid mites
can alter the structure of the soil. So oribatid mites are considered the most impor-
tant group of arachnids from the standpoint of direct or indirect effects on the for-
mation and maintenance of soil structure (Haq 1994; St. John et al. 2001).
All active instars of oribatid mites are involved in the decomposition of organic
materials as part of humification and nutrient cycling processes in soils. Oribatids
feed on a range of materials including living and dead tissues of higher plants,
microflora, carrion eggs of fish and living microfauna. Many oribatid mites are asso-
ciated with various parts of higher plants (including pollen). Oribatid mite
Minunthozetes semirufus lays its eggs on grasses; larvae and nymphs feed on stems
through burrowing. Oribatid mite Orthogalumna terebrantis mines the leaf of the
water hyacinth, Eichhornia crassipes, an important aquatic weed, and Parlohmannia
dissimilis damages root systems of potato, strawberry and tulip. Oribatid mite
Galumna allatum has been reported to attack the seeds of winter wheat after they
are sown (Haq 1992, 1994). Some species of oribatid mites are of great veterinary
importance because of their role as intermediate hosts of animal parasitic tape-
worms as given earlier (Denegri 1993).
Oribatid mites are reported as potentially useful in bioassay work as ecological
indicators of biotypes and of the effects of human activities of terrestrial ecosystems
including air pollution, acid rain, use of fertilizers, controlled and accidental burn-
ing, forest harvesting, radioactive pollution, renovation of mine spoils, sewage
water irrigation and trampling (Labrun and Streelan 1995).
10.13 Conclusions
The role of mites in soil ecosystem is only partially understood. They contribute in
fragmentation of litter (comminution) and soil formation, nutrient recycling, disper-
sal of microbial spores and stimulation of the microflora (bacteria and fungi) by
grazing. Soil mites also help in predation on both micro-arthropods and nematodes.
Comminution – the breaking up of larger organic units such as dead leaves and
wood into smaller pieces – is a process that is largely limited to oribatid mites and
Astigmata. The other group of mites are either predaceous or feed by puncturing,
not breaking up fungal hyphae, etc. Comminution greatly increases the surface
areas on which bacteria can complete the actual process of decomposition. Through
feeding and production of their faecal matter, oribatid mites can alter the structure
of the soil. So they are considered as the most important among arachnids from the
Further Reading 205
standpoint of direct or indirect effects on the formation and maintenance of soil

structure. Many oribatid mites also sequester calcium and other minerals in their
cuticle, and thus their bodies may form important ‘sinks’ for nutrients in some
nutrient-limited environments. Some species of oribatid mites are of great veteri-
nary importance, because of their role as intermediate hosts of animal parasitic
tapeworms.
Further Reading
Brown A (1978) Ecology of soil organisms. Heineman Educational Books, London (ISBN-
10:0435606204,128 pp; ISBN-13: 978–0425606206)
Banerjee S (1984) Qualitative and quantitative composition of oribatid mites (Acarina) in relation
to certain soil factors. In: Griffiths DA, Bowmann CE (eds) Acarology VI. Ellis Harwood,
Chichester, pp 878–885
Denegri GM (1993) Review of oribatid mites as intermediate hosts of tapeworms of the
Anoplocephalidae. Exp Appl Acarol 17:567–580
Diaz A, Okabe K, Eckenrode CJ, Villani MG, Oconnor BM (2000) Review: biology, ecology, and
management of the bulb mite of the genus Rhizoglyphus (Acari: Acaridae). Exp Appl Acarol
24:85–113
Dindal DL (ed) (1976) Biology of oribatid mites. Suny, Syracuse/New York
Dindal DL (ed) (1990) Soil biology guide. Wiley, New York, p 1349
Emmanuel N, Curry JP, Evans GO (1985) The soil Acari of barley plots with different cultural
treatments. Exp Appl Acarol 1:101–113
Evans GO (1992) Principles of acarology. Cambridge University Press, Wallingford, pp 563
Ghilyarov MS, Bregetova NG, eds (1997) A key to the soil inhabiting mites: mesostigmata.
Zoologicheskogo Institute Academi Nauk SSSR, Leningrad, 1973; pp 718
Gulati R, Mathur S (1995) Effect of Eucalyptus and Mentha leaves and Curcuma rhizomes on
Tyrophagus putrescentiae (Schrank) (Acarina: Acaridae) in wheat. Exp Appl Acarol
19:511–518
Haq MA (1992) Beneficial aspects of oribatid mites. In: Haq MA, Ramani N (eds) Man, mites and
environment. Anjengo Publications, Calicut, pp 128–138
Haq MA (1994) Role of oribatid mites in soil ecosystems. In: Ecology and biology of soil organ-
isms. Agrotech Publishing Academy, Udaipur, pp 143–177
Kuhnelt W (1961) Soil biology (with special reference to the United Kingdom). Faber and Fabar,
London
Labrun P, Streelan NMV (1995) Oribatid mites: prospects for their use in ecotoxicology. Exp Appl
Acarol 19:361–379
Marshall VG, Reeves RM, Norton RA (1987) Catalogue of the oribatida (Acari) of continental
United States and Canada. Mem Ent Soc Can 119(139):1–418
Norton RA (1985) Aspects of biology and systematics of soil arachnids particularly saprophagous
and mycophagous mites. Quest Entomol 21:523–541
Rockett CL (1980) Nematode predation by oribatid mites (Acari: Oribatida). Int J Acarol
6:219–224
St. John MG, Behan-Pelletier M, Bagatto G, Lindquist EE, Shorthouse JD, Smith IM (2001) Soil
formation and mite colonization of rehabilitated mine tailings. Plant Soil 245:295–305
Thomas RH, MacLean SP (1988) Community structure in soil Acari along a longitudinal transect
of tundra sites in Northern Alaska. Pedobiologia 31:113–138
Tomlin AD, Miller JJ (1987) Composition of the soil fauna in forested and grassy plots at Delhi.
Ont Can J Zool 65:3048–3055
Veeresh GK, Rajagopal D, Viraktamath CA (eds) (1991) Advances in management and conserva-
tion of soil fauna. Oxford and IBH Publishing Company, New Delhi
206 10 Soil Mites
Winchester NW, Behan-Pelletier VM, Pang RA (2000) Arboreal specificity, diversity and abun-
dance of canopy-dwelling oribatid mites (Acari: Oribatida). Pedobiologia 43:391–400
1. https://www.youtube.com/watch?v=wMNdnQZGoqo Soil Mites Feasting

2. https://www.youtube.com/watch?v=G3in74EBalQ Red Velvet Mite
3. https://www.youtube.com/watch?v=vKhdIITsTjw Soil mites
4. https://www.youtube.com/watch?v=AMBgEi0yaV8 Soil moss mites – Oribatida
5. https://www.youtube.com/watch?v=XqRLVUtAAV8 Selected Invertebrates of the Soil Food
Web
6. http://tombio.myspecies.info/files/MitesKeyTest-2014-04-01.pdf A Key to Soil Mites in the UK
7. http://www.fcps.edu/islandcreekes/ecology/soil_mite.htm Soil mites(Oribatidae)
Quarantine Acarology
11
Abstract
Many arthropods including insects and mite species have been transported
around the globe and successfully invaded new regions. An invasive species is
alternatively known as an alien or exotic and is introduced into an environment
where it is not native. The introduction of invasive species in any country whether
deliberate or unintentional from around the globe can pose a significant threat to
native animals or plants. Adventive (non-invasive) and invasive Acari share the
impact with small arthropods such as scale insects, whiteflies, thrips and psyl-
lids. All of these organisms are closely associated with plants and have minute
size, cryptic colouration, tolerance for extreme low or high temperatures and the
ability to survive dry or wet conditions. Mites are the most diverse and different
group of arthropods encountered in quarantines. Infestation of mites,
Proctolaelaps sp., is the single biggest threat to Drosophila stocks. Many spider
mite species intercepted at the port of entry belong to the genus Tetranychus (T.
evansi, T. fijiensis, T. kanzawai and T. pacificus), tenuipalpid mites, eriophyid
mites (gall, erinose and rust mites) and tarsonemid mites (broad mites and cycla-
men mites) because of the intercontinental movement of fruits, flowers and orna-
mental plants. The exchange of honeybees across the globe has accelerated the
spread of bee parasitic mites, Varroa destructor, in European honeybees, Apis
mellifera. Many animal parasitic mites and ticks have been deliberately intro-
duced through the imports of food such as sheep, cattle, horses, goats, pigs and
camels and imports of wheat, cotton and clovers. To deal with the problem of
invasive alien species (IAS), various international conventions and organizations
have been initiated around the world.
Keywords
Invasive pests • Quarantines • Fact sheets • International trade

DOI 10.1007/978-981-10-1594-6_11
208 11 Quarantine Acarology
Many arthropod species have been found transported around the globe and success-
fully invaded new regions. An invasive species (alternatively known as an alien,
exotic, injurious species) refers to an animal or plant that is introduced into an envi-
ronment where it is not native. The introduction of invasive species in any country
whether deliberate or unintentional from around the globe can pose a significant
threat to native animal and plant communities and may result in extinctions of native
animals or plants. Invasive arthropods can have severe impacts on animal and human
health, agriculture and forestry and the biodiversity of natural habitats as well as
those modified by humans. The economic and environmental effects of invasion can
be both direct through feeding and competition and indirect such as the transmis-
sion of plant or animal pathogens (Pimental 2002; Sanders et al. 2010; Sharma and
Thakur 2007). Hundreds of billions of dollars are lost annually as a result of direct
and indirect damage caused by invasive arthropods. In the USA, it is estimated that
50,000 non-native species have been introduced. The potential economic costs asso-
ciated with nonindigenous plant and animal species are estimated at $129 billion
annually, and in 2012, the US government spent an estimated $2.2 billion across a
range of federal agencies and activities in an effort to prevent, control and eradicate
invasive species domestically (ER1).
Adventive (non-invasive) and invasive Acari share the impact with the small
arthropods such as scale insects, whiteflies, thrips and psyllids. All of these organ-
isms are closely associated with plants and have in common their tiny size, cryptic
colouration, tolerance for extreme low or high temperatures and ability to survive in
dry or wet conditions, and even some of them have the ability to go without food for
months and perhaps years, making the mites as one of the most successful adventive
and often invasive groups of arthropods. Although strict quarantine and inspection
requirements are now enforced in many areas (Corn 2013; Dent 1981), invasive
arthropods are often inconspicuous or live in cryptic habitats and can be transported
undetected (ER2).
11.1 Acari Dispersal
Mites and ticks disperse by attaching themselves to other species. For example,
adult ticks may be transported to long distances while attached to the migratory
birds. The bee mite, Varroa destructor (Anderson and Trueman), spreads through
attachment to adult bees when colonies split and through the queen bee trade.
Dispersal may also be inadvertent by human activities such as international trans-
portation of people, foodstuffs and other materials when some species have been
intentionally introduced, either by cultivation or to control other invasive species
(Ashburner 1989; Estoup and Guillem 2010).
The increasing levels of trade and transport between continents and countries
have meant an increased risk of introduction and spread of pests (insects and mites).
Pest mites which were earlier capable of dispersal only over short distances by natu-
ral processes can now be transferred inadvertently from country to country or from
one geographical region to another by means of fast international transport links.
11.3 Invasive Mites and Ticks 209
Acari have become one of the fastest, unwanted pest travellers since the beginning
of twenty-first century because of their small size and tolerance to many of the pes-
ticides and other procedures on a wide variety of commodities (Boubou et al. 2011;
Griffiths and Bowman 1981; Hunt 1998; Milani 1999).
From the USA, about 100 non-native tick species have been reported in the past
century, and these introductions of invasive ticks have increased markedly in recent
years through the international trade in live animals (particularly reptiles). Some of
these species have been reported as vectors of serious diseases affecting both
humans and animals, besides some other species doing direct damage to their hosts
(Mullen and Oconnor 2009; Sanders et al. 2010).
11.2 Quarantine Pests
Quarantine pests have been defined as pests of potential economic importance to the
area endangered thereby and not yet present there, or present but widely distributed
and being officially controlled. Therefore, quarantine pests are those that are con-
sidered to be potentially very serious for the country concerned (Follett and Neven
2006; Murthaiyan 2009).
Mites are the most diverse and difficult group of arthropods encountered in quar-
antine. The single biggest threat to Drosophila stocks is an infestation of mite,
Proctolaelaps sp. (family Melicharidae). Some mite species eat only the flies’ food;
others eat embryo and pupae and can completely wipe out a fly laboratory. If left
unchecked, mites can increase in number so rapidly (often by parthenogenesis) that
they can destroy fly stocks. To guard against mites, all flies coming into the labora-
tory from other laboratories or stock centres must go directly to the quarantine room
in the attic and remain there for 6 weeks (Ashburner 1989).
Species established in a new environment that proliferate and spread in ways that
are destructive to human interests are considered ‘invasive alien species’ (IAS).
Nowadays, such species are recognized as some of the greatest threats to the well-
being of the planet; they can cause a serious damage to the biosecurity and to agri-
cultural ecosystems. Phytophagous mites are very prone to becoming IASs. A single
surviving female of many of the phytophagous mite species can start a new popula-
tion in an invaded environment because of their characteristic parthenogenetic type
of reproduction. In addition, mite populations have been observed to quickly
develop resistance to pesticides and supposedly resistance to other detrimental con-
straints to its biology (Anonymous 2004; Walter 2006).
11.3 Invasive Mites and Ticks
The ecological impact of invasive species includes direct and indirect effects on
native plants, animals and whole ecosystems. Direct predation on and competition
with native organisms may lead to the displacement or extinction of the native popu-
lations. Their impact on biosecurity is seen particularly within inland habitats
wherein the past such competition has been minimal due to the isolation of the
indigenous populations. Indirect and ecosystem level effects are also important, and
complex ecosystem processes may be disrupted by the presence of alien invasive
mite species. Invasive species threaten biodiversity by causing disease, acting as
predators or parasites, acting as competitors in altering habitat or hybridizing with
local species (Burridge 2011).
11.4 Phytophagous Mites
Most of the plant feeding mites belong to families Tetranychidae (spider mites),
Tenuipalpidae (false spider mites) and Eriophyidae (worm mites). Reports of most
of the spider mites intercepted at port of the entry belong to genus Tetranychus, and
prominent examples are T. evansi, T. fijiensis, T. kanzawai and T. pacificus (Hummel
et al. 2009; Yaninek and Harren 1988). The increasing number of species inter-
cepted indicates that intercontinental movement of fruits, flowers and ornamental
plants provided adventive species with the opportunity to expand their range into
new areas (ER3 & ER4).
False spider mites intercepted at port of entry include mainly Brevipalpus phoe-
nicis (Geijskes), Raoiella indica Hirst and other related species. B. phoenicis is a
known vector of citrus leprosis, and several related viruses of ornamental plants that
occur in the Central and South America. So dilemma exists because many of the
diseases including citrus leprosis do not occur in the USA, but its potential vectors
are already present. So legislation is needed regarding thorough treatment/examina-
tion of plant material received at port of entry and mandatory risk mitigation in
nurseries abroad wherefrom shipments originate.
Several horticultural plants have been found invaded by worm or rust mites.
Eriophyid mites are of great quarantine concern as they live in secluded places, and
it is almost impossible to quarantine that a plant is free of eriophyid mites without
destroying it. So transportation of vegetative parts, whole plants, branches with
buds for grafting, etc., as well as fruit seeds from palm trees, for example, as coco-
nut, unless the bracts are carefully removed and the remaining nut washed in an
acaricide solution, can be done only very carefully. These mites cause rusting of
flowers and buds or cause leaf edge rolling. Coconut palm mite, Aceria guerreronis
Keifer, is a threat on queen palm and many indoor ornamental plants. In coconut
palms, this mite sometimes affects more than 60 % plants and causes losses from 20
to 60 % of the production. These new introductions are a serious threat for the
booming trade of ornamental plants in different countries. Tarsonemid mite
Steneotarsonemus spinki Smiley, commonly known as rice white mite, is a great
threat for rice cultivation as its infestations cause sterility of rice seeds by transmit-
ting the rice sheath rot fungus and a mycoplasma-like organism. Other mite species
like Polyphagotarsonemus latus Banks and Tarsonemus confusus Ewing (both
Tarsonemidae) and Rhizoglyphus spp. (Acaridae) are also potentially serious inva-
sive pests (Fan and Petit 1998; Grbic et al. 2011; Hummel et al. 2009; Navajas and
Ochoa 2012).
11.6 Animal Parasitic Mites and Ticks 211
11.5 Bee Mites
European honey bee, Apis mellifera (Hymenoptera: Apidae), has been used by
humans as a crop pollinator and producer of honey. The exchange of bees across the
globe has accelerated the spread of bee parasitic mites, V. destructor and V. jacob-
soni Oudemans. A. mellifera is very sensitive to varroa mite infestations and serious
colony losses are reported. In addition to direct losses caused by mite feeding, V.
destructor is a vector of bee viruses and bacteria. Untreated bee colonies generally
die within 2–3 years, a consequence which has resulted in the complete disappear-
ance of wild bee colonies in many regions. V. destructor and Acarapis woodi
(Rennie) mites have resulted in the near elimination of feral honey bees (A. mel-
lifera) in the USA. Significant economic losses have resulted in reduced availability
of managed bees that are essential for pollinating many crops within the USA
(Navajas and Ochoa 2012; Oldroyd 1999).
In Australia, A. mellifera is reported as a major pollinating agent for crops like
almonds, avocado, blueberry, cucumber, watermelon and sunflower. In case of
canola, cotton and pasture clovers, yield is increased by 10–20 % with honey bee
pollination. Devastating entry into Australia of V. destructor, which has spread
around the world in recent decades, but yet with a few of the places like Australia
still left for it to colonise, may prove disastrous for beekeeping. In Europe, the USA
and New Zealand, varroa mite has virtually eliminated feral colonies and also is
responsible for a major decline in managed hives. Australia is now bracing itself for
the entry of varroa mite in an undetected swarm of honey bees hitching a ride by
ship from New Zealand or other nearby country (Hunt 1998).
11.6 Animal Parasitic Mites and Ticks
In countries that are dependent on exotic species across the globe, many species of
ticks and mites have been deliberately introduced from imports of food and fibre
such as sheep, cattle, horses, goats, pigs, camels, wheat, cotton and clovers.
Numerous other foreign species obtained entry permits for cultural or aesthetic rea-
sons, including rabbits, cats dogs and foxes (Burridge 2011; Mullen and Oconnor
2009).
More than 250 species of mites that cause problems for humans and domestic
animals have been reported. These mites can cause problems such as temporary
itching of the skin due to bites, persistent ‘dermatitis’ due to mites invading skin or
hair follicles, mite-induced allergies, transmission of pathogenic microbial agents,
intermediate agents of tapeworms and invading respiratory passages of internal tis-
sues and organs. Mites can infest skin, scales, feathers or fur of their host and some-
times enter the respiratory passages or internal tissues and organs. The chicken
mite, Dermanyssus gallinae (De Geer), causes dermatitis in avian hosts; follicle
mite (Demodex bovis Stiles) infests the hair follicles of cattle; itch mite (Psoroptes
ovis Hering) causes itch in sheep; and psoroptic ear mite (Psoroptes cuniculi
Delafond) causes inflammation of the ears in sheep; they are the most important
invasive mites encountered in quarantine laboratories (both regional and interna-

tional). So strict quarantine regulations should be followed at all levels. About one
hundred species of ticks have been reported from various reptiles in various imports
from numerous countries at various ports of entry and destroyed (Mullen and
Oconnor 2009). But in some other cases, however, the ticks have gone unnoticed
and in a few cases have become established at least temporarily. Cattle tick,
Rhipicephalus microplus (Canestrini), is an important pest that feeds on cattle and
deer but can be carried by other large animals. Native to India, Indonesia and Japan,
it has also been found in the tropical and subtropical regions of Africa, Australia and
the USA. R. microplus causes heavy losses to the cattle industry through its feeding
activities. It also acts as the major vector for the protozoan parasites, Babesia bige-
mina and B. bovis, which cause babesiosis or cattle fever. Unable to move on its
own, R. microplus is spread by movement of host while it is feeding. The practice
of moving cattle by driving and, despite animal movement restrictions, R. microplus
is now endemic in many areas of the world (Goka et al. 2013).
11.7 Quarantine Organizations
To deal with the problem of IAS, various international convections and organiza-
tions have been developed around the world. Quarantine requirements for organ-
isms in general are based on the list of quarantine species of each Regional Plant
Protection Organizations (RPPOs) and of each country. RPPOs are required to
review their list of quarantine mites for not including some knowingly important
species and including species of known minor importance. Phytosanitary require-
ments in international trade to prevent and control pests have to be specifically justi-
fied through pest risk assessment (PRA) processes. Phytophagous mites as IAS
have received great attention particularly regarding international trade of fresh
products like fruits and cut flowers and vegetative propagation material. In PRA,
phytophagous mites have been considered of medium/low risk of establishment
when associated with products for immediate consumption, due to their low mobil-
ity. However, when associated with vegetative parts, the risk has been assumed to be
medium/high.
Evaluation of pest risk analysis (PRA) needs salient information like host range,
geographic distribution, bioecological characteristics and the potential to cause
social and economic problems. To proceed for a detailed PRA, information regard-
ing updated catalogues and data with the required information about the invasive
quarantine mites are needed (Anonymous 2004; Corn 2013).
Phytosanitary measures as mite interception in entry points and eradication pro-
grammes are hampered because of the difficulty to detect mite infestations and are
often considered not viable procedures. Quarantine officials should be trained to
proceed for acarological inspections; rather a permanent collaboration of taxono-
mists and quarantine reference collections are fundamental points to interception of
quarantine mites (Oceuli et al. 2014).
11.8 Internet Resources for Information on Pest 213
The ideal method of controlling mites is to avoid all exposure to the pest mites,
since many important mite pests have local or regional, rather than global, distribu-
tion often because of the limitations on quality of the pest to spread great distance
without human assistance. So regulatory or legal steps should be taken to exclude
pests that might be inadvertently imported on nursery stock, seeds, etc. In the USA,
the responsibility for interpreting and implementing laws at the international level
is assigned to the US Department of Agriculture, Animal and Plant Health Inspection
Service (USDA-APHIS). Even each state also retains authority to implement analo-
gous regulations directed at interstate movement of plant pests (Corn 2013; Dent
1981).
FAO has provided pest risk analysis (PRA) details to determine if pests are quar-
antine pests. In these details, the integrated processes to be used for risk assignment,
as well as the selection of risk assignment options for analysis of risks of plant pests
to the environment and biological diversity, including where affecting uncultivated/
unmanaged plants, wild flora, habitats and ecosystems contained in the PRA area
(Dent 1981; Follett and Neven 2006). Pest risk analysis for quarantine pests follows
a three-stage process:
Stage I (Initiation of the process). Identifying pests and pathways that are of quar-
antine concern should be considered for risk analysis in relation to the identified
PRA area.
Stage 2 (Risk assessment). It begins with the categorization of individual pests to
determine whether the criteria for a quarantine pest are satisfied.
Stage 3 (Risk management). It involves identifying management options for reduc-
ing the risk identified at stage 2. These are evaluated for efficacy, feasibility and
impact in order to select those that are appropriate. Irradiation-controlled atmo-
spheres and temperature treatments have been developed for mite disinfestation
of the fresh products. In this way the probability of introduction and establish-
ment of the mites will be reduced to insignificant levels.
11.8 Internet Resources for Information on Pest
Mites (Acari) are considered to be the most diverse group of arthropods encoun-
tered in quarantine and plant protection activities and are of great importance in
import and export activities concerned with crops, fresh fruits, vegetables and live-
stocks and pose major challenges for quarantine officials. So a resource for informa-
tion about invasive mite identification and tools concerning quarantine and plant
protection has been developed (http://keys.lucidcentral.org/keyserver/player.
jsp?keyId=51) which provides information regarding taxonomy and identification
(preliminary/advanced). The tools range in complexity from ‘Is it a mite? A basic
introductory level tool’, to species level treatments (such as Tetranychus and
Phytoseiulus). This resource also includes an acarine glossary and an up-to-date
taxonomic classification of the Acari.
Acarological Society of America and the US Department of Agriculture, Animal

and Plant Health Inspection Service (USDA-APHIS) have published a report enti-
tled ‘A Ranking Model and List to identify foreign Tetranychoidea pests of
Agricultural and Environmental Significance to the USA'. Ranking model included
criteria like value of impacted crops, regarding potential high risks of countries,
host plants and damage potential of intercepted species. Report also includes infor-
mation regarding reproductive potential and vector of any plant disease. Details
about quarantine information are available at this site: http://www.aphis.usda.gov/
wps/portal/aphis/ourfocus/planthealth/sa_manuals/!ut/p/a1/
pZHLUoMwFIafxYVLJoeU6xLayqV.
11.9 O
ther Internet Resources Providing Information
on Mites and Insect Pests
–– International Standard for Phytosanitary Measures (ISPM)

–– (http://www.ippc.in/default.jsp)
–– American Phytopathological Society (APS)
–– (www.apsnet.org).
–– CAB International (CABI)
–– (www.cabi.org)
–– CABI Plant Protection Compendium
–– (www.cabicompendium.org/CPC)
–– European and Mediterranean Plant Protection Organization (EPPO)
–– (www.eppo.in)
–– Global Invasive Species Programme (GISP)
–– ( h t t p : / / w w w. d ive r s i t a s - i n t e r n a t i o n a l . o rg / a c t iv i t i e s / p a s t - p r o j e c t s /
global-invasive- species-programme-gisp)
–– International Plant Protection convention (IPPC)
–– (www.ippc.net/IIP/En/default.htms)
–– Invasive Species Specialist Group (ISSG)
–– (www.issg.org)
–– Pestnet
–– (www.pestnet.org)
11.10 Pest Information/Fact Sheets/Field Guides
Pest information sheets provide identifying details of target pests that the survey
team can refer to in the field (ER4). These sheets should be written in a manner that
it is simple and easy to read. A pest information sheet includes information regard-
ing the pests’ common and scientific name, host range, symptoms, morphology and
colour photographs or diagrams of the pest showing the typical morphology and
characteristic stages and on multiple hosts and preferred habitats. Such fact sheets
have been made available by USDA-APHIS, EPPO, FAO, CABI and some other
organizations of different countries (Corn 2013; Dent 1981).
11.11 Conclusions
At present we have entered in an era of unprecedented levels of travel and interna-

tional trade, and with this there have been too much increases in the movement of
pests which were once considered exotic. With the increase in imports of ornamen-
tal plants, cut flowers, fresh vegetables and fruits, there are high chances of invasive
pests entering the countries/continents and establishing in new locations where their
natural enemies are not existing. Mite-vectored diseases such as ‘citrus leprosis’ and
other related virus-like pathogens pose additional threats resulting in more eco-
nomic or environmental harm. The single biggest threat to Drosophila stocks is an
infestation of parasitic mite, Proctolaelaps sp. (family Melicharidae), which mainly
eat the flies’ food, but also eat embryos and pupae affecting behaviour of flies and
sometimes completely wipe out a fly laboratory. So to prevent mite outbreaks,
always quarantine coming stocks. Fact sheets about these mites should be available
and proper quarantine operations should be followed before keeping stocks. Keep
the fly stock for two or more mite free generations.
Invasive bee parasitic mite, Varroa destructor, has resulted in the near elimina-
tion of feral honey bees (Apis mellifera) and even managed bees that are essential
for pollinating seed production of many crops like apple, avocado, cucurbits, canola
and sunflower resulting in reduced yields. In Australia, varroa mites have been con-
sidered as major threat for beekeeping and honey bee crop pollination industries.
Quarantine in northern Australia is very important because of its proximity to
Southeast Asia and the Pacific which have many pests not present in Australia. In
Europe, New Zealand and the USA, varroa mite has virtually eliminated feral colo-
nies and resulted in a major decline of managed hives. Productive hives are aban-
doned and the workers just vanish and V. destructor implicated in colony collapse
disorder resulting in formation of strict quarantine measures at ports of entry. The
potential cost of Varroa infection to the Australian Economy has been estimated at
US $ 16.4 million to US $ 38.8 million per year. Control of V. destructor has been
made more difficult to as resistance to acaricides has evolved in the mite population,
and no eradication strategies are currently considered feasible. CSIRO estimated
that preventing the varroa mite from entering Australia over the next 30 years would
lead to savings up to AUS $ 21.3–50.5 million per year.
Australia, Canada and the USA have strict quarantine standards for dealing with
agricultural materials and animals. Fact sheets have been prepared and made avail-
able in quarantine laboratories (both international and regional movement of mate-
rial). These fact sheets provide very useful aid for quarantine staff regarding colour
photographs and their scanning electron microscope (SEMs) prepared photographs
of different pests.
Considering great importance of mites and ticks, which are tiny and sometimes
difficult to find by quarantine staff, scientific collaboration of entomologists and
veterinarians with quarantine staff should be active. There is a need of integrating
information on molecular, morphological and ecological data information and mak-
ing it available particularly for invasive species at quarantine stations. Outdated
facilities in laboratories at ports of entry of material should be replaced by modern
facilities like differential interphase contrast (DIC) and confocal microscopes

(LT-SEM) which provide a detailed view of characters for more clear observation
and quickly help in discriminating invasive species. LT-SEM high-resolution digital
images of intercepted specimens after freezing and then extracting their DNA will
substantially help by keeping in a voucher collection. Already some selected inter-
national efforts for addressing invasive species of insect, mites and other arthropods
have been made. Convention on International Trade in Endangered Species of
World Fauna and Flora (CITES), 1973; South Pacific Regional Environment
Programme (SPREP) Convention, 1990; Convention on Biological Diversity
(CBD), 1993; World Trade Organization (WTO); and Agreement on the Application
of Sanitary and Phytosanitary (SPS) Measures, 1995, are already doing useful work
regarding regulation of trade in some wildlife and plant species, taking measures to
protect rare or threatened species and also taking measures to protect the health and
lives of humans, plants and animals (Seeman and Beard 2011).
There is a strong need for developing additional national and international stan-
dards for phytosanitary measures to improve uniformity and transparency of infor-
mation exchanged between countries when mitigating trade in new commodities.
Further Reading
Anonymous (2004) Pest risk analysis for quarantine pests, including analysis of environmental
risks and living modified organisms. ISPM no. 11. pp 117–138
Ashburner M (1989) Drosophila: a laboratory handbook. Cold Spring Harbor Laboratory Press,
Cold Spring Harbor
Boubou A, Migeon A, Roderick CK, Navajas M (2011) Recent emergence and worldwide spread
of the red tomato spider mite, Tetranychus urticae: generic variation and multiple cryptic inva-
sions. Biol Invasions 13:81–92
Burridge MJ (2011) Non-native and invasive ticks: threats to human and animal health in the
United States. University Press, Florida, p 320
Corn ML (2013) Invasive species: major laws and the role of selected federal agencies.
Congressional Research Service Report 7–5700 (www.crs.gov/) R 43258. Congressional
Research Service, Washington, D.C
Dent D (1981) Quarantine, legislation and policies. In: Insect pest management. CAB International,
Wallingford, pp 421–437
Estoup A, Guillem T (2010) Reconstructing routes of invasion using genetic date: why, how and
what? Mol Ecol 19:4113–4130
Fan Y, Petit FL (1998) Dispersal of the broad mite, Polyphagotarsonemus latus (Acari:
Tarsonemidae) on Bemisia argentifolii (Homoptera: Aleyrodidae). Exp Appl Acarol
22:411–415
Follett PA, Neven LG (2006) Current trends in quarantine entomology. Annu Rev Entomol
51:359–385
Goka K, Okabe K, Takerno A (2013) Recent cases of invasive mites and ticks in Japan: why is a
regulatory frame work needed? Exp Appl Acarol 59:245–261
Grbic M et al (2011) The genome of Tetranychus urticae reveals herbivorous pest adaptations.
Nature 479:487–492
Griffiths DA, Bowman CE (1981) World distribution of the mite, Varroa jacobsoni – a parasite of
honey bee. Bee World l62:154–163
Further Reading 217
Hummel NA, Castro BA, McDonald EM, Pellerano MA, Ochoa R (2009) The panicle rice mite,
Steneotarsonemus spinki Smiley, a pest of rice in the United States. Crop Prot 28:547–560
Hunt J (1998) The war against Varroa: how are we doing? Am Bee J 138:372–374
Milani N (1999) The resistance of Varroa jacobsoni Oud. to acaricides. Apidologie 30:229–234
Mullen GR, Oconnor BM (2009) Mites (Acari). In: Mullen GR, Durden LA (eds) Medical and
veterinary entomology, 2nd edn. Academic, New York, pp 433–492
Murthaiyan MC (2009) Principles and practices of plant quarantine. Allied Publishers Pvt Ltd.,
New Delhi
Navajas M, Ochoa R (2012) Editorial: integrating ecology and genetics to address Acari invasion.
Exp Appl Acarol 59:1–10
Oceuli S, Nejad KHI, Ziaie F, Moghuddam M (2014) Gamma irradiation used on adult Tetranychus
urticae Koch as a quarantine treatment. J Pl Prot Res 54:150–155
Oldroyd BP (1999) Coevolution while you wait: Varroa jacobsoni, a new parasite of western hon-
eybees. Trends Ecol Evol 14:312–315
Pimental D (2002) Environmental and economic costs associated with alien invasive species in the
United States. In: Biological invasions: economic and environmental costs of alien plant, ani-
mal and microbe species. CRC Press LLC, Boca Raton. ISBN 0-8493-0836-1
Sanders CJ, Miller PS, Wilson AJ (2010) Invasive arthropods. Rev Sci Tech Int Epiz 29:273–286
Seeman OD, Beard JJ (2011) Identification of exotic pest and Australian native and naturalized
species of Tetranychus (Acari: Tetranychidae). Zootaxa 2961:1–72
Sharma S, Thakur M (2007) Roles of plant quarantine in the management of pest organism – a
review. Agric Rev 28:235–244
Walter DE (2006) Invasive mite identification: tools for quarantine and plant protection. Interactive
Keys. Lucid V. 3.3. (www.Itp.lucidcentral.org/keys/v3/mites)
Yaninek JS, Harren HR (1988) Introduction and spread of the cassava green mite, Mononychellus
tanajoa (Bonder) (Acari: Tetranychidae), an exotic pest in Africa and search for appropriate
control methods: a review. Bull Entomol Res 78:1–13
1. https://www.youtube.com/watch?v=4pT0ylaRYY8 Plant Protection and Quarantine

Recruitment Video
2. https://www.youtube.com/watch?v=7aVpGpf8zRQ MIAMI PLANT INSPECTION STATION.
avi
3. http://itp.lucidcentral.org/id/mites/invasive_mite/Invasive_Mite_Identification/key/Whole_site/
Home_whole_key.html Invasive Mite Identification: Tools for Quarantine and Plant Protection
4. http://www.planthealthaustralia.com.au/wp-content/uploads/2013/09/Exotic-spider-mites-FS.
pdf Plant Health Australia: Factsheet Exotic Spider Mites
Mite Pests of Horticultural Crops
12
Abstract
Horticulture is defined as that branch of agriculture which is concerned with
intensive cultivation of plants that are used by people for food, for medicinal
purposes and for aesthetic diversification. Pomology (fruit production), olericul-
ture (production of vegetables) and floriculture (production of plants for orna-
mental use) are branches of horticulture. Mites are encountered as pests regularly
or as sporadic pests under all situations. Fruits (both temperate and tropical
fruits) are infested by different groups of mites at all growing stages and appear
under favourable conditions of climatic conditions. Spider mites, false spider
mites and eriophyid mites are commonly observed damaging different horticul-
tural crops in different regions. Vegetable production mainly during hot and dry
conditions and in protected cultivation is commonly infested by different mite
species. Floricultural plants grown in open field conditions as well as under pro-
tected conditions are comparatively highly infested by mites causing significant
reduction in flower and seed production. Mite pests of important fruit crops,
vegetable crops and flowering plants are elaborately discussed giving nature of
damage caused, their host range and measures to manage them in different situ-
ations so as to decrease damage caused by them.
Keywords
Horticulture • Spider mites • Management • Protected cultivation
Horticulture is defined as that branch of agriculture which is concerned with inten-

sively cultivated plants that are used by people for food, for medicinal purposes and
for aesthetic gratification. Horticultural plants are commonly divided into those that
are edible, those that are used for culinary or medicinal purposes and those that are
used for ornamental and aesthetic purposes. The scientific study of horticulture is
divided into various subdisciplines. Pomology is defined as that branch of horticul-
ture dealing with fruit and tree nut production. Fruit production includes the

DOI 10.1007/978-981-10-1594-6_12
220 12 Mite Pests of Horticultural Crops
so-called tree fruits such as apples, peaches and oranges and small fruits such as
strawberry, blueberry and raspberry. Olericulture is that branch of horticulture
which deals with the production of vegetables and herbs. Floriculture deals with the
production of field grown or greenhouse grown plants for their flowers or showy
leaves. Environmental horticulture is that branch of horticulture that deals with the
production of plants for ornamental use in constructed environments, both indoors
and outdoors; environmental horticulture also includes nursery production. Mites
are encountered as pests regularly or as sporadic pests under all these situations and
are considered below under separate heads particularly mentioning the mites that
are of economic relevance.
12.1 Fruits
12.1.1 Temperate Fruits
Temperate fruits are adapted to climates in the middle latitudes. Temperate fruits
have two climatic adaptations: they require some cold periods (dormancy) to com-
plete their life cycle which conditions their adaptability in cold climates. They may
be tree fruits (apple, pear, peach), vine fruits (grape, kiwi fruit) and small fruits
(strawberry, raspberry, currents and blackberry).
12.1.1.1 Almonds (Prunus amygdalus, Rosaceae)

Almond is a deciduous tree, originated from the Middle East and South Asia. The
USA, Spain, Iran, Turkey and China are important producers. Almond prefers
Mediterranean climate.
Peach Silver Mite (Aculus cornutus (Banks), Eriophyidae) The body is teardrop
shaped and because of its small size can only be seen with a high-powered hand lens
(15× or more). Peach silver mite usually does not appear in damaging proportions
on almonds but trees under 6 years of age are sometimes highly infested by mites.
Mite feeding results in tiny chlorotic spots that give the leaves a silvery appearance
especially along the midvein region of the upper leaf surface. Treatments against
this mite are needed only if the mite population is very high (hundreds of mites/leaf)
which can cause defoliation.
Spider Mites Studies in California reported that severe spider mite infestation
resulted in about 16 % reduction in yield, a 25 % drop in terminal growth and a 7 %
drop in leaf size. Pacific spider mites, oriental brown mites, two-spotted spider
mites (Tetranychus urticae Koch, TSSM), European red mites (Panonychus ulmi
(Koch), ERM) and citrus red mites (Panonychus citri (McGregor), CRM) sometimes
cause damage in almond orchards (Fig. 12.1). Monitoring for mites should be done
at weekly intervals during May–August and focus particularly on monitoring of hot
spots – areas where mites develop first. When mite populations reach treatment
threshold in these areas, then initiate measures for controlling mites (ER1).
12.1 Fruits 221
Fig. 12.1 Mite-damaged
almond tree
12.1.1.2 Apple (Malus domestica, Rosaceae)

The apple tree has its origin in Kazakhstan but is now found in many regions with a
temperate or subtropical climate. Most apples are produced in China, the USA, Iran,
Turkey, Russia, India and France.
Web-Spinning Spider Mites Four species of spider mites, namely, two-spotted

spider mites, Pacific spider mite (Tetranychus pacificus McGregor), European red
mite and McDaniel spider mite (Tetranychus mcdanielli McGregor), have been
reported feeding on apple leaves and fruits. As a result of mite feeding, fine stip-
plings on leaves are found. Heavy mite infestations result in severe bronzing of
foliage and premature defoliation. Fruits on heavily infested trees fail to achieve
proper size and colour. In heavily mite infested orchards, fruit production for the
following year may be lowered (Hoyt 1969; Gerson and Cohen 1989). Mites are
found in abundance during hot summer months, especially in dusty and water-
stressed areas of the orchard (ER2).
Monitoring of mite population during May–August helps in determining the

need for treatment. In general apple varieties, Red Delicious, Golden Delicious and
Jonathan, are comparatively more susceptible to mite injury. Adequate irrigations
should be provided to avoid water stress. Manage properly the cover crops from
where the mites may move up into the apple trees. During active season, i.e. July–
August, monitor mite population regularly and pesticide treatment may be consid-
ered when an average of ten mites per leaf is reached; it is also reported that if one
predatory mite per ten spider mites is found, then treatment may not be applied.
Keeping the trees well irrigated and reducing dust sources help in minimizing mite
infestation in the apple orchard. Keeping economic threshold 10–30 mites/leaf
proves useful in initiating treatments against these mites (Cranham and Soloman
1981).
Apple Rust Mites (Aculus schlechtendali (Nalepa)) These mites have been
reported both as a pest and a beneficial arthropod. Mites damage both the fruit and
foliage of apple; however, these mites also play a major role as food of predatory
phytoseiid mites. Apple rust mite attacks cultivated apples and several other Malus
spp. like pear. Rust mite overwinters as deutogynes (females) in crevices on twigs
and under bud scales. Sometimes large cluster of mites is found under a single scale.
Mites emerge during spring and feed on undersurfaces of leaves as the buds begin
to open. Development is more rapid in warm temperature conditions. As a result of
rust mite feeding, a silvery cast to the leaf in the early stages appears which tends to
get browner as the season progresses. Damage can be particularly severe on young
trees in nurseries and newly planted material. Rust mite damage sometimes causes
leaves to roll lengthwise. Excessive amounts of damage with peak populations
greater than 300 mites/leaf or 4800 mite days can reduce significantly the fruit
growth (Easterbrook 1996).
Leaf Blister Mites (Phytoptus mali (Burts), LBM) Feeding by LBM causes blis-
ters on leaves and fruits. LBMs are rarely found in commercial orchards, but also
reported to attack quince, mountain ash, snowberry and hawthorn. Eggs are oval
(40 μm long). Female is light to amber yellow, cylindrical and tapering sharply at
the posterior end. It overwinters as mature females at the base of buds or under outer
bud scales. In spring, when buds begin to swell, overwintered females penetrate
deeper into the buds and lay eggs on the live tissue. Mite feeding causes blisters on
developing leaves. Several generations of mites develop within blisters during a
growing season. When blisters became crowded or leaves become heavily damaged,
mites may migrate to growing terminals where their feeding produces new blisters.
Fruit damage is caused by injury to buds before bloom. Severe infestation can
deform apple fruits, but scarring can make the fruit unmarketable. Severe damage to
foliage can cause leaf drop and reduce shoot growth. Well-managed orchards are
less infested by leaf blister mites.
12.1 Fruits 223
12.1.1.3 Apricots (Prunus armeniaca, Rosaceae)

Brown mites and ERMs may also infest and damage apricots and these mite pests
may be controlled as given under peaches.
12.1.1.4 Cherries (Prunus avium and P. carasus, Rosaceae)

Cherries are deciduous, originated from China and mainly produced in Europe
(Italy, France and Spain) and in some other countries around the Mediterranean and
Central Asia. Apricots like continental climate with cold winters and are native to
the temperate region of the Northern Hemisphere. Brown mites, CRM and web-
spinning spider mites also sometimes infest cherries. Control measures as given for
peaches may be applied to manage mites.
12.1.1.5 Currants (Ribes spp., Grossulariaceae)

Currants include black currants, red currants and white currants; currants are native
to temperate regions of the Northern Hemisphere.
Two-Spotted Spider Mites (T. urticae, TSSM) Two-spotted spider mites are light
tan or greenish with a dark spot on each side. Mottling and bronzing of leaves as a
result of mite feeding occur, crop vigour is reduced and leaves turn brown and
sometimes drop. In the fall, adults cease feeding and migrate to overwintering sites.
Female mites overwinter under loose bark or organic debris of host plants. Scouting
for mites and their webbing on the undersurface of leaves should be done for know-
ing the need of spraying acaricides. As the spider mite infestations are favoured by
dry and dusty conditions, avoid creating these conditions and stressing the plants.
Periodic hosing of plants with a forceful jet of water can physically remove and kill
many mites.
Big Bud Mite (Cecidophyopsis ribis (Westwood)) Damages in late winter and the
infested buds become abnormally swollen and rounded, while healthy buds are
pointed and longer than broad. Infested buds often dry up, producing no leaves in
spring, or stunted foliage and a few or no flowers. Always purchase certified stock
plants, after inspection of nursery as being free of big bud mites and reversion.
Dispose of heavily infested plants after the fruit has been picked, and plant in
autumn with clean new stock. The mite can also transmit a virus disease ‘black cur-
rant reversion virus’.
12.1.1.6 Grapevines (Vitis vinifera, Vitaceae)

Grapes are grown as deciduous woody vines in warmer temperate climates with a
lot of sunshine. Important grape-producing countries include China, Italy, the USA,
Spain, France, Turkey, Chile and Argentina.
Spider Mites Mainly Pacific spider mite (T. pacificus), Willamette spider mite
(Eotetranychus willamettei (McGregor)), two-spotted spider mite, T. urticae and
Oligonychus mangiferus (Rahman and Sapra) are among the important pests of
vineyards (Fig. 12.2). Pacific spider mites prefer the warmer upper canopy of the
Fig. 12.2 Mite-damaged grapevine leaf
vine and can cause damage early in the season, but prefer hot and dry part of the
season for its rapid multiplication. The Pacific spider mite is of larger size than the
Willamette mites. In general forelegs of Pacific spider mite are reddish in colour but
are translucent to yellow in case of Willamette mites. The two-spotted spider mite is
almost identical in appearance to the Pacific spider mites but it rarely causes dam-
age on vineyards. Immediate treatment of T. pacificus is advisable whenever more
than 50 % leaves exhibit moderate to heavy injury. Manage spider mites in a vine-
yard by integrating all the available biological, cultural and chemical control
options.
Grape Leaf Bud Mite (Colomerus vitis Pagenstecher) They are creamy white
(200 μm long), worm-like creatures. Adult females lay eggs inside the swelling bud
during spring, and immature mites feed under the bud scale. Bud mites overwinter
as adults under the outer scales of buds. During bud burst some mites move from the
budding shoot to new developing buds. Mite feeding can lead to malformed leaves,
aborted or damaged bunches and even bud mortality. The damage can be unsightly
but does not usually have economic significance.
Grape Leaf Rust Mite (Calepitrimerus vitis (Nalepa)) They are cream to pinkish
worm-like creatures. Rust mites are much more active than bud and blister mites.
Rust mites overwinter under the bark of cordons or the trunk near the crown. The
most visible and easily recognizable symptoms of rust mite occur from January to
March. The leaves start to darken and take on a bronzed appearance because of the
rust mites feeding and damaging on the surface cells of the leaf.
12.1 Fruits 225
False Spider Mites (Brevipalpus californicus (Banks) and Brevipalpus lewisi

McGregor) Adults are 0.3 mm long, are shield-shaped and are reddish brown.
Females deposit oval, bright red eggs throughout the vine. In spring these mites feed
on developing canes and later on the undersurfaces of leaves. Early season damage
is characterized by small dark spots or scars around the base of canes; mites then
move to the bunch stalks and can partly starve berries preventing sugar accumula-
tion. The adults spend the winters under the outer bud scales and under the rough
bark at the base of the canes. In general these mites do not seem to be of great
significance.
12.1.1.7 Mulberry (Morus spp., Moraceae)

Mulberries originated from Southeast Asia, are deciduous in nature and are widely
distributed in warm temperate or subtropical climates.
Broad Mite (Polyphagotarsonemus latus (Banks)) Broad mites have been

reported attacking mulberry leaves and resulting in reduced yield of leaves needed
for silkworm rearing.
Spider Mite (Tetranychus equatorius McGregor) Sometimes they are found as a

minor pest of mulberry in Karnataka (India). Spider mite, Tetranychus ludeni
Zacher, which normally infests bean, brinjal, castor and cucurbits, is also reported
infesting mulberry leaves. Initially white specks on leaves are found which later
coalesce to form big patches when the density of mite increases. Mite-damaged
leaves appear rusty, gradually dry and fall. Another spider mite, Eotetranychus sug-
inamensis (Yokoyama), is also reported attacking the undersurface of mulberry
leaves causing yellowish-white symptoms on mite-infested leaves. The quality of
silk produced is lowered when mite-infested leaves are used as food for rearing
silkworms.
Bud Mite (Aceria mori (Keifer)) Bud mite has been reported infesting mulberry
from India. Due to mite attack, poor sprouting of cuttings is seen. Newly trans-
planted fields show sometimes patches due to the death of planted cuttings/saplings.
Drying of auxiliary buds is found which results in poor leaf yield per plant. The mite
attacks mainly during December to February.
12.1.1.8 Nectarines (Prunus persica, Rosaceae)

Nectarines are also commonly known as ‘shaved peach’, ‘ fuzzy-less peach’ or ‘
Brazilian peach’ because of the lack of fuzz or short hairs.
Brown mite (Bryobia rubrioculus (Scheuten)), ERM (P. ulmi), peach silver mite
(Aculus cornutus (Banks)) and web-spinning spider mites (T. urticae and T. pacifi-
cus) also infest nectarines. These mites need constant monitoring during the sum-
mer months. Control measures as given under peaches may be used for management
of mite pests on nectarines.
12.1.1.9 Peach (Prunus persica, Rosaceae): Deciduous, Originated

from China
Brown Mite (B. rubrioculus) Brown mites have flattened body with long front
legs. Adult mites are brownish green; eggs are red and are similar in appearance to
the eggs of European red mite but lack a stipe. Nymphs are red. Brown mites over-
winter as eggs on spurs and branches, and these eggs hatch in spring. Young mites
move out to leaves where they feed but do not produce webbings. Mites feed only
during the cooler parts of the day and migrate off the leaves during mid-day. As a
result of mite feeding, leaf injury begins as mottling and browning of leaves. In case
of mite infestation, the entire trees will take on a pale-yellow appearance. Trees can
tolerate low to moderate populations of mite. Monitoring for brown mite eggs is
most important during the dormant season. Use dormant sprays with oils at the high
rate to help control the overwintering eggs if 20 % or more of the shoots have eggs
(Croft and Hoyt 1983).
European Red Mite (P. ulmi, ERM) Females of ERM have a brick red, globular
body with long curved hairs that arise from white spots or tubercles on the back.
Nymphs or unfed females may appear greenish. Eggs are red, are slightly flattened
and have a stripe protruding from the top. The mites overwinter in egg stage on
twigs and spurs and hatch in early spring at the time of leaf formation. About eight
to ten generations are produced before fall. Mite-damaged leaves which take on a
finely mottled appearance rarely fall because of mite feeding (ER4).
Peach Silver Mite (A. cornutus) They are worm-like mites having yellowish to
pinkish white colour and are somewhat wedge-shaped and can only be seen with a
hand lens of 10× or more power. These mites overwinter as females in bark crevices,
around buds and under bud scales and move to leaves soon after bud break in spring.
On small fruits they often inhabit the area between the calyx and the fruit. Later in
the season, the mites are found primarily on the lower leaf surface. Female mites
move to overwintering sites before leaf fall occurs. During the season, many genera-
tions of mite are produced. Heavily infested leaves take silvery appearance. If heavy
mite populations (200–300 mites/leaf) are found, consider application of
acaricides.
Spider Mites (T. urticae and T. pacificus) Two-spotted spider mites (TSSM) and
Pacific spider mites overwinter as adult females in protected places on the trees or
in the leaf litter, trash and weeds on the orchard floor. Overwintering forms of both
mite species are reddish orange; mites become active in early spring soon after trees
start producing leaves. Hot and dry conditions are favourable for both the species.
Active summer females are greenish or pale yellow with large dark spots on each
side of the body. The Pacific mite often has a second pair of spots near the posterior
end of the body which helps in distinguishing it from TSSM. Eggs are spherical and
almost translucent when first laid; egg laying is mainly on the underside of leaves,
but during heavy mite infestation, eggs may be deposited on both surfaces. There
may be 8–18 generations depending on temperature.
12.1 Fruits 227
Stone fruits can tolerate some mite damage particularly on water sprouts in the
centre of trees. TSSM generally feeds on lower leaf surface but Pacific mites may
be found on both leaf surfaces. Mite feeding results in mottling of leaves, but in case
of severe mite infestations, leaf fall is quite common. Both mite species produce
heavy webbings. If defoliation happens early in the season, fruits fail to achieve
proper size and are exposed to sunburn. Depending on the per cent leaves infested
by mites, mite infestation on leaves has been rated in different studies as high (80–
100 %), moderate to high (61–79 %), moderate (40–60 %), low (21–39 %) and very
low (1–20 %).For successful management of these mites, regular monitoring is
essential. During April–August, monitor mites at more frequent intervals. Plants
should not be water stressed. Keep orchards well irrigated so that hot, dry and dusty
conditions necessary for mite build-up are not readily available.
12.1.1.10 Plums (Prunus domestica, Rosaceae)

Plums are temperate fruits, probably originated from the Caucasus mountains near
the Caspian Sea. Plum trees are deciduous in nature. Brown mite (B. rubrioculus),
European red mites (ERM) and eriophyid mites Aculus fockuei (Nalepa and
Troussart) and Diptacus gigantorhynchus (Nalepa) have been reported on plums.
Brown mite and ERM have been discussed earlier under peaches. Eriophyid mites
are yellow to pinkish white with wedge-shaped body. Heavily infested leaves take
on a silvery or bronzed appearance; severe infestation may interfere with photosyn-
thesis. Mite population as high as 120 per leaf does not impact yield during the year
of mite infestation. Another eriophyid mite, Eriophyes phloeocoptes Nalepa, some-
times infests the base of the current year’s shoots and scars the affected tissues.
Feeding by nymphs of mites causes formation of small spherical galls, the leaves of
which thicken. Flowering shoots are attacked, growth is interrupted and flowers
develop imperfectly. There is only one generation in a year.
12.1.1.11 Pear (Pyrus spp., Rosaceae)

Pear is native to coastal and mildly temperate regions of the old world. Most pears
are cold hardy deciduous trees.
Pear Leaf Blister Mite (Eriophyes pyri (Nalepa)) The mite body is white, long
and slender and striated and can be seen with a hand lens of 15× or more power.
Eggs of the mite are spherical and pearly white and are laid in blisters. Blister mites
feed under the bud scales during winter and may cause buds to dry and fail to
develop in spring (ER3). Mites feed on emerging leaves from green tip through
bloom and also on developing fruit. Because of the mite feeding, oval russet spots
on fruits are found. Several generations per year may develop within the blisters.
Leaf feeding causes small blisters which are first red and later turn black. Abandoned
and unsprayed young pear trees are subject to the periodic severe blister mite infes-
tations. Loss of foliage through mite feeding weakens trees and reduces tree growth.
The use of pyrethroids and some other pesticides may increase blister mites and
their damage on pear. So regularly monitor the blister mites.
Pear Rust Mite (Epitrimerus pyri (Nalepa)) Pear rust mites are sporadic but are
very serious pests of fresh market pears. Rust mites may appear in pest forms if
pyrethroids are used in pear orchards. These mites look pale brown and females
overwinter in bark crevices or behind loose bud scales usually on 2–4-year-old
woods. As the buds open in spring, the mites move to developing clusters, leaves
and fruits to feed. During the growing season, adults look pale white to cream
coloured. Rust mites damage the surface of the fruit and foliage resulting in bronz-
ing of the tissue. Mites develop on fruits around the calyx or stem end, giving a
localized mite-infested areas if mites are unchecked; russetting may spread over the
entire fruit. Rust mites are not an economic pest of naturally rusted varieties. The
control of pear rust mites is best obtained during the postharvest period. Monitoring
for mite population is a must to determine if acaricide treatments are desired. Rust
mites are a problem particularly in orchards where predaceous mites are destroyed
through the use of pyrethroids and other materials.
European Red Mite (P. ulmi, ERM) Eggs of the mite are laid on leaves during the
season. ERM overwinters as eggs at bases of buds and spurs on smaller branches
and twigs or in wounds. Feeding by ERM causes the leaves to lighten in colour and
become mottled or stippled. Prolonged mite feeding during high mite infestations
eventually causes leaf bronzing, and under high temperatures, portions of the leaf
blade or its entire surface may turn brown and dry. Mite burn can even be noticed
after the mites have been controlled, if high temperature still occurs. Pear trees may
be regularly monitored and controlled if more than 10 % spurs are infested during
dormant to delayed dormant period. Acaricide application against mites depends on
mite damage during different periods of the year.
Bryobia Mites (B. rubrioculus) Bryobia mites attack a range of fruit trees but are
most common on pears and almonds. The adult female mites are quite long (0.7 mm)
and are purplish brown. Heavily mite-infested leaves become pale and may prema-
turely fall. Bryobia mites overwinter in the egg stage on the bark of branches and
twigs; sometimes the number of overwintering eggs is so high that parts of the bark
have a reddish tinge. The young mites hatch in spring and feed on the leaves. Several
generations of mites may develop in a season before the overwintering eggs are laid.
Sprays of winter oils usually control satisfactorily the Bryobia mites.
Two-Spotted Spider Mite (T. urticae, TSSM) Adult female bears dark spot on
either side of the body. Female lays tiny, spherical, colourless to light straw-coloured
eggs. Overwintering females are orange and hibernate under bark scales on the tree
and in trash on the ground, emerge in late March–April and feed on leaves. During
hot and dry weather, rapid multiplication of mites occurs, and peak population is
found in June and continues if the weather remains dry for a longer duration.
Because of severe mite feeding and too much web-spinning activity, characteris-
tic blackening of pear leaves is found. Usually two to three mites feeding on a leaf
during hot and dry weather can produce black areas from the midrib to the margins
12.1 Fruits 229
of the infested leaves. High mite population causes defoliation, and severe defolia-
tion can stunt fruit (Figs. 12.3 and 12.4). TSSM is found most abundantly on pear
orchards during hot and dry periods when too much dusting conditions are found
and trees are water stressed. Orchards with cover crops and sprinkler irrigation are
most suitable for an IPM of mites. Do not allow the cover crop to become dry as this
will cause TSSM to disperse to pear trees. Many low-growing crops and weeds
allow easy dispersal of the mite to the pear trees. Manage the weeds so that mite
population is checked from migrating to the pear trees. Regular monitoring of leaves
and fruits can be done when pears turn brown. During turn down pear to harvest, an
action threshold of 1–50 mites/100 leaves should be followed and treatment with oil
may be considered. But if the population is more than 50 mites/100 leaves, then
include miticide sprays to the oil spray.
Oriental Brown Mite (Eutetranychus orientalis (Klein)) Oriental brown mite is

reported as a major pest of sand pears during April–August in Jalandhar (Punjab),
India (Fig. 12.5).
12.1.1.12 Raspberry (Rubus idaeus, Rosaceae)

Raspberries are perennial deciduous plants grown in temperate climates in Europe
and northern Asia.
Spider Mites On raspberries two species of spider mites, namely, two-spotted spi-
der mites (TSSM) and yellow mite (Eotetranychus carpini (Oudemans)), are com-
Fig. 12.3 Grades of mite damage on pear leaves

Fig. 12.4 Leaf fall by mite feeding on pear trees
Fig. 12.5 E. orientalis-damaged pear twig
monly found. Feeding by TSSM produces small yellowish spots on the upper
surface of leaves but the mite infestation and their webbings are found on the under-
surface of leaves. Margins of high mite-infested leaves appear dried; infested leaves
turn silver or bronze and later yellowish brown before completely drying up and
falling off. TSSM is overwintered as adult females later within the soil and plant
debris. Yellow mite appears as a more concentrated yellowish-brown colouration
along the secondary and tertiary veins. Compared to TSSM, yellow mite produces
very little webbing. Infested leaves exhibit lower photosynthesis, increased transpi-
12.1 Fruits 231
ration, deformed leaves and lower chlorophyll. Mite injury causes a mottling of the
leaves which may later become brown or bronze.
Raspberries can tolerate significantly greater densities of yellow mites compared

to TSSM. A preharvest spray may be necessary if spider mite population approaches
ten or more mites per leaflet and predator prey ratio is less (1:10). Spider mite popu-
lation can rapidly increase after harvest through early September, but treatments are
not necessary unless mite population reaches 25 mites or more per leaflet. Integrated
management practices that reduce dust on foliage and fruit minimize the risks for
spider mite outbreaks. Use only selective miticides for spider mite control to
enhance natural control by predatory mites.
Raspberry Leaf and Bud Mite (Phyllocoptes gracilis (Nalepa)) Raspberry leaf
and bud mite attacks raspberries and some hybrid berries. Mites are more numerous
in sheltered environments (Gordon and Taylor 1976).
12.1.1.13 Strawberry (Fragaria ananassa, Rosaceae)

Strawberries grow best in temperate climates or the cooler hills in the tropics. They
are mainly grown in the USA, Mexico, South America, Spain and other European
countries.
Spider Mites Two species of spider mites, namely, two-spotted spider mites, T.
urticae and T. cinnabarinus, are commonly found on strawberry at different stages
of plant growth (ER5). Mite feeding results in stipplings, scarring and bronzing of
the leaves. T. urticae is particularly damaging during the first 2–5 months following
transplanting in late summer substantially reducing a number of berries per plant.
Plants are less sensitive to mite feeding after initial berry set; however, a substantial
yield loss occurs if 15–20 mites per mid-tier leaflet are observed at this time.
Infestations of more than 75 mites per leaflet may severely weaken the plant; stunt-
ing, drying or even development of reddish colouration may occur. If possible,
avoid early season insecticide applications or apply insecticides that are less disrup-
tive to natural enemies of spider mites. Uncontrolled spider mite infestations can
reduce strawberry yields by 25 %. Most growers use predatory mites in combination
with insecticides for better mite control.
Cyclamen Mite (Phytonemus pallidus (Banks)) They are whitish brown, live in
the crowns of the plant and feed on developing leaves. It is most prevalent in hot and
dry summers. At low population densities, cyclamen mites are usually noticed along
the midvein region of young unfolded leaves. These mites overwinter as eggs,
nymphs or even adults on the underside of the leaf close to the ground. Cyclamen
mites feed mainly on developing leaves. The mites inject toxic saliva in the plant
cell which causes the leaves to become wrinkled and puckered. The first generation
takes 1 month, but later during the summer months, each generation is shorter
(10 days). There can be seven generations in a year. P. pallidus prefers high relative
humidity (80–90 %), and at temperature above 20 °C, these mites avoid taking ref-
uge in the crown of the plant. Cyclamen mites are primarily pests in fall-planted and
second year plantings. Heavily infested leaves become severely stunted and crin-
kled resulting in a compact leaf mass in the centre of the plant. Mite feeding on
flowers can cause them to wither and die. Fruit on infested plants are dwarfed, and
the seeds stand out on the flesh of the berry. Propagating nursery stock free of cycla-
men mites is essential to prevent introducing mite populations to fruit-producing
fields. In general cyclamen mites are difficult to control because the mites live in
closed leaf and in flower clusters where the penetration of pesticide is difficult.
12.1.2 Tropical/Subtropical Fruits
Fruits that are very sensitive to cold and are often injured by low temperatures above
freezing are grouped under tropical fruits. But the fruits that do not specifically
require cold but have slight frost tolerance are grouped under subtropical fruits.
12.1.2.1 Avocado (Persea americana, Lauraceae)

Avocado is an evergreen tree originated from Southern Mexico and Central
Americas and requires tropical or subtropical climate. The trees need a climate
without frost and cannot tolerate strong wind.
Persea Mite (Oligonychus perseae Tuttle, Baker and Abbatiello) Persea mite
has been reported as a key pest of avocado in California. Adult females have an
oval-shaped body and are slightly flattened and elongated. Females and immature
mites are yellowish or greenish with two or more small dark blotches on their abdo-
men. Persea mites feed and reproduce mostly beneath webbed patches. Female lays
round, pale-yellow eggs. Peak mite population is generally found in July and
August, but low humidity and high temperature (more than 37 °C) for several con-
secutive days result in rapid decline of the mite population. Premature leaf drop and
defoliation occur as a result of high infestation of mites. Mite feeding on undersides
of leaves results in circular chlorotic to brown spots which become visible on the
upper surface of leaves. At high population levels (100 mites/leaf) if avocado trees
were not treated, then about 20 % reduction in yield is reported. To manage these
mites on avocado, minimize stress by having appropriate timings and frequency of
irrigations. Persea mite-susceptible ornamental host plants should be reduced or
eliminated. Excess fertilization of avocado trees results in increasing persea mite
populations and their damage. Spraying undersides of leaves with a forceful stream
of water can reduce mites on a few small trees wherever possible.
Avocado Brown Mite (Oligonychus punicae (Hirst)) Females are dark brown
with oval body and lay amber-coloured eggs having a short projecting stalk. Eggs
are laid on the upper surface of leaves along the midrib region. Temperatures above
32 °C usually kill these mites and their eggs. O. punicae is a sporadic pest; mite
feeding during July to September results in partial defoliation. Higher mite infesta-
12.1 Fruits 233
tion causes leaf bronzing along the midrib and small veins, and finally the entire leaf
lamina turns brown. Many times mite infestations are found on border row trees
particularly along dusty roads which leads to mite outbreaks. For proper manage-
ment, controlling dust and avoidance of application of broad-spectrum insecticides
for different other avocado pests proves helpful. If 50–70 adult female mites or
about 100–200 mobile mites are found, then control measures should be initiated.
Six-Spotted Spider Mite (Eotetranychus sexmaculatus (Riley)) Adult mites

have lemon-yellow colour with six dark blotches on the abdomen. Eggs are globu-
lar, pale-greenish yellow to translucent. High mite populations are rarely reported
during spring and early summer. Mite population rarely exceeds an average of 2–3
mites/leaf. Whenever six or more mites per leaf are found, then only treatments may
be initiated. These mites occasionally attain the pest status but can become a prob-
lem if trees are drought stressed or when mite control by natural enemies is dis-
rupted through the use of broad-spectrum pesticides in avocado orchards. Dusts
around avocado orchards should be minimized, and consider using a water trailer to
wet dusty roads. Individual backyard trees can be hosed down in early to midsum-
mer to remove dust and enhance biological control of six-spotted mites.
Avocado Bud Mite (Tegolophus perseaflorae Keifer) During March to May, avo-
cado bud mites sometimes are found on buds and on developing fruits. Bud mite
feeding causes necrotic spots and irregular openings in apical leaves and may cause
deformation and discolouration of fruit.
12.1.2.2 Banana (Musa paradisiaca)

Banana is an evergreen and grown in almost all tropical regions. Banana originated
from Southeast Asia.
False spider mite, Brevipalpus phoenicis (Geijskes), is reported causing some
damage on banana plantation. Banana spider mite, Tetranychus lambi, is among the
most important and widespread mite pests of banana and most active in the dry
spring to summer periods. Outbreaks of these mites are usually localized within a
plantation (ER6). The warm, dry conditions under plastic bunch covers are particu-
larly favourable for build-up of banana spider mites. In general this is a minor and
sporadic pest of bananas. Mite damage is mainly confined to the underside of lower
older leaves; however, in severe outbreaks the mites move onto the bunches and
damage the fruit. Infested leaves turn brown grey and in extreme cases leaves wilt.
If mites are found on young leaves, treatments should be applied to prevent damage.
Reduce dusts on roadways as much as possible.
12.1.2.3 Citrus
Citrus originated from Southeast Asia. Citrus is a general term which refers to a
large number of species of fruit trees in the family Rutaceae. Citrus is an evergreen,
usually spiny shrubs or trees, and is subtropical in origin and cannot tolerate severe
frosts. Brazil, the USA, Mexico, Spain, China and India are the leading
citrus-growing countries. Citrus includes grapefruit, lime, lemon, malta, mosambi,

orange and pummelo.
More than 104 phytophagous mite species have been reported on citrus from dif-
ferent parts of the world, mainly belonging to families Tetranychidae, Tenuipalpidae,
Tuckerellidae, Phytoptidae, Eriophyidae, Diptilomiopidae and Tarsonemidae.
Losses due to mite injury are quite substantial especially during years when climatic
conditions are more favourable for their development (Beettle et al. 1991; Dhooria
et al. 2005; McCoy 1977).
Citrus Brown Mite (Eutetranychus orientalis) Citrus brown mite is also com-
monly known as oriental red mite, citrus mite and citrus brown mite and, in earlier
literature, is also known as Anychus orientalis and Anychus ricini, but both these
names are considered as synonyms of Eutetranychus banksi. E. orientalis has very
broad host range; more than 85 plant species distributed in 28 different plant fami-
lies are reported as host plants from India alone. However, family Rutaceae and
Rosaceae have maximum recorded host plants. Besides found on different citrus
species, Cassia fistula, Ricinus communis, cannabis, oleander, Azadirachta indica,
Melia azadirachta, papaya, almonds, peaches and pear are among the most impor-
tant host plants of this mite (Readshaw 1975). Total developmental period (egg–
adult) of 12–13 days and average fecundity per female of 51 eggs are reported.
Faster development of citrus brown mite has been reported on R. communis, French
bean, lime and mandarin leaves. However, shorter developmental period with higher
longevity and fecundity is reported on young leaves (ER7 and ER8).
Mites suck the sap of leaves predominantly from the upper surface of leaves; the
affected leaves turn chlorotic and finally drop resulting in poor plant growth.
Infestation of mites normally starts along the midrib of leaves and later spreads
along lateral veins. The leaf region on either side of midrib and veins turns pale yel-
low, and in case of severe infestation, the upper surface of the leaf turns pale com-
pletely. The webbing produced by the mite attracts dust particles. However, the
lower surface of leaves remains comparatively free of mites and their damage. Peak
mite populations on citrus species are reported during March–June and again during
September–October. Positive correlation between mite population and temperature
on different citrus species is found, while rainfall and relative humidity are nega-
tively correlated with mite population. Citron (Citrus medica), C. sinensis, key lime
(C. aurantiifolia) and sweet orange are relatively reported less susceptible to mite
attack. C. limettioides, C. karna, grapefruit (C. paradisi) and C. acida are reported
most susceptible for mite attack (Dhooria et al. 2005).
Citrus Red Mite (Panonychus citri, CRM) CRMs are potential pests of citrus in
Europe, Asia, Russia, China, Japan, North and Central America, Africa and
Australia. Roses, almonds, pears and mulberry are other important hosts of
CRM. The adult females are oval and dark red; the dorsal setae are long and carried
on distinct tubercles or knobs that are of the same colour as the integument. The
males are smaller, more pointed posteriorly and with relatively longer legs as in
12.1 Fruits 235
citrus brown mite. Eggs are pale to dark red, spherical and somewhat flattened hav-
ing a stalk from which guy threads extend to the surface of the leaf. Eggs are laid on
the surface of the leaf mainly along the midrib.
At mean temperature of about 24 °C and a relative humidity of about 55 %, eggs
hatch in 7–9 days, and the combined nymphal development takes 5–7 days. The life
cycle from egg to egg takes about 17 days. Each female lays 20–50 eggs at a rate of
2–3 eggs/day. Eggs are laid on both sides of leaves. Net reproduction rate per gen-
eration (Ro) and the mean intrinsic rate of increase (rm) have been estimated to be
19.57 and 0.1708, respectively, at 25 °C. Egg development takes about 17 days.
Temperature of 40 °C or a few warm days (32 °C and 5 % RH) with strong winds
may result in a high mite mortality. A relative humidity below 25 % is unfavourable
and hinders the development of CRM. A temperature of 35 °C is unfavourable for
survival of males, while all stages die at 48 °C. In general both high temperatures
and very low humidity adversely affect the mite population.
Mite population increases in spring, late summer and early fall in response to
plant growth. CRMs prefer to feed on fully expanded young leaves but also infest
fruits. CRMs feed on the upper surface of the leaves, the fruit and the bark of green
branches of all citrus species. Major damage is done on leaves and green fruits. As
a result of mite feeding, the infested leaves initially take on a stippled appearance;
later pale patches appear. As the mite population increases, the leaf looks grey, sil-
ver or yellow (Beettle et al. 1991).
Citrus Flat Mites (Brevipalpus californicus and B. phoenicis) Citrus flat mites
are very small and are difficult to see with the naked eye. The body of adult female
is oval red, with a dark-red pattern on the middle of the body. Mixed populations of
B. californicus and B. phoenicis are frequently found on infested citrus fruits. The
males are more pointed posteriorly and do not have the dark-red pattern in the
middle. The females lay elliptical bright red eggs which are covered with an adhe-
sive substance and are sucked to the leaf surface. Incubation period is about 9 days
at 18–24 °C and a relative humidity of 55 %. Initially the larvae are uniformly red
but after feeding, a black pattern begins to form on the body. At 21–30 °C and 85 %
relative humidity, it takes about 3 weeks to complete the life cycle (Childers et al.
2003).
Citrus flat mite has a very broad host range, and the mite feeding has been
reported to produce a variety of damage symptoms on citrus. From Argentina, B.
californicus has been reported to cause ‘leprosis’-type symptoms on citrus leaves.
During feeding the mite injects toxic substances along with its saliva and results in
a condition known as ‘nail-head rust’ on leaves and fruits. On fruits, mite feeding
results in silvering, and on twigs and branches, mite feeding causes scaly bark
symptoms which are commonly known as ‘Florida scaly bark’. Feeding by B. phoe-
nicis has been reported to cause rind spotting in case of grapefruits. From 10 to 15 %
fruits in case of grapefruits, pummelo and orange are reported damaged by mites,
but kinnow fruits are more severely damaged by the mite. In case of Blood Red,
Jaffa and Valencia Late, negligible rind disorder symptoms are reported. Mites dam-
aged the epidermal region mainly, and infested fruits looked brown to blackish with
irregular corky skin. Mite damage is particularly severe on plants which are stressed.
However, rind-disordered fruits may not adversely affect yield and sweetness of
fruit, but certainly affected the customers’ preference for fresh fruits with healthy
skin (Childers et al. 2011).
Citrus Bud Mite (Eriophyes sheldoni Ewing, CBM) The citrus bud mite is
worm-like pale yellow, is very small and can only be seen with a 10× hand lens
under field conditions. CBMs attack all species of citrus, but lemon is most dam-
aged because of its larger buds which offer good protection for the mites. CBMs are
reported from almost all lemon-growing areas of the world where humidity is high
enough for its development (Bergh and French 2002).
Female mite lays about 50 pearly white, subspherical eggs during its lifetime;
eggs hatch in 3–4 days and egg hatching is most successful at 25 °C and 98 % rela-
tive humidity. Egg hatching is significantly reduced when relative humidity is low
(35–40 %). One generation is completed in 12–33 days. The mites are found under
bud bracts, on petiole bases next to buds, beneath bud scales, on developing flowers
and under fruit buttons. All stages of buds including dormant buds on the old wood
are infested by mite. Mites move to new growth and confine themselves under fruit
buttons. Bud scales are blackened and often the entire bud is killed. As a result of
continuous mite injury extensive multiple budding on infested twigs, cessation of
new growth and blossom development is found. Mite-infested buds are shortened or
thickened and flattened. Sometimes on lemon trees, rosette growth is observed due
to bud proliferation. Leaves from damaged buds assume peculiar shapes; blossoms
are malformed with abnormal or aborted floral parts. Fruits formed from infested
flowers may drop prematurely or assume odd shapes (Walker et al. 1992).
Citrus Rust Mite (Phyllocoptruta oleivora (Ashmead), CRM) Citrus rust mite
(CRM) is very small, fusiform, dorsally flattened and yellow. The eggs are smooth,
spherical and semitranslucent and are laid in groups in indentations on fruits and on
ventral surface of leaves. CRM has been reported as a serious pest of citrus in dif-
ferent parts of the world and reproduces rapidly. Mean incubation period of 3 days
and nymphal development of 2–11 days have been reported (32 °C). The life cycle
takes 7–10 days in summer and 14 days in winter. A female laid 20–30 eggs during
its life span of about 20 days (Allen 1979).
Citrus fruits are attacked by the mite when they are of the size of peas. Mite-
damaged fruits become silvery, reddish brown or purplish black. Sometimes the
whole surface of the fruit is damaged. Affected fruits bear a comparatively thicker
skin and have rusty spots which render them unacceptable as fresh fruits in the mar-
ket. Mite feeding resulted in a silvered or shark skin-like appearance in case of
grapefruits and lemons. The damaged fruit remains small in size and deteriorates
rapidly. High infestation of rust mites causes bronzing of leaves and green twigs.
12.1 Fruits 237
Mite feeding on fruits adversely affects the external appearance of fruits. Mites
avoid direct sunlight and infest mostly the undersurface of leaves and shaded area
on the fruit; infestation is generally more during warm and humid weather condi-
tions. In summer mites prefer fruits than the leaves. Fruits on upper branches are
more preferred. Economic threshold (ET) of ten rust mites/cm2 for fruits is required
for processing purposes, but ET level of only two rust mites/1 cm2 is reported for
sale as fresh fruits. So monitoring of leaves and fruits regularly from early April is
most important.
Broad Mite (Polyphagotarsonemus latus) Broad mite is also mentioned as

Hemitarsonemus latus in earlier literature. Commonly broad mite is also known as
yellow tea mite or tropical mite. Oval elongate eggs are laid by female mites on new
growth. The dorsal surface of egg is studded with whitish tubercles in clumps. Eggs
are mostly laid in depressions on the leaf or fruit surfaces, firmly attached to the
base. Larvae feed actively and nymphs remain enclosed within skins of the quies-
cent larvae until the adults emerge. The adult females are large, oval and broad hav-
ing rich amber or dark-green look; the colour of mites depends on the host on which
they feed and multiply. The adult male is short and broad, tapering at the posterior
end, with large fourth pair of legs (Pena et al. 1999).
P. latus has a very broad host range – citrus, tea, tobacco, potatoes, chillies, ger-
bera, dahlia and jute are comparatively more damaged by the mite. Broad mites are
distributed throughout the tropics and also in the greenhouses on a wide variety of
plants in temperate regions and on a wide variety of agricultural crops. Optimal
environmental conditions include warm temperatures, high humidity and low light
intensity. Adult mites can survive through prolonged exposure to freezing tempera-
tures but are sensitive to temperatures greater than 32 °C.
Broad mite is capable of feeding on very young tender leaf or fruit tissues; toxic
saliva injected by the mite can result in significant damage. On citrus, mites feed on
the lower surface of leaves causing them to become rigid and curled. Feeding injury
is confined to young foliage; nursery plants are more damaged by the mites. On
ageing, damaged leaves split or crack producing a rugged appearance of various
shapes. The ventral surfaces of infested leaves become bronzed; mites also attack
citrus fruits leading to silvery-white symptoms which later turn corky. Under green-
house conditions limes begin to show damage 4–6 days after infestation, and severe
damage to the fruit epidermis appears 12 days after infestation. In coastal areas
when 5 % of the fruits are infested, plant protection measures need to be initiated.
Citrus Leprosis ‘Rhabdovirus’(CiLV) CiLV is also known as leprosis, lepra

explosiva de les citricos (in Spanish), Leprese dos citrus (in Portuguese) and leprous
dos agrumos (in French). Citrus species especially grapefruits and oranges are
found naturally infected by CiLV. Lemons and mandarins are less susceptible. No
other plant species is known to serve as natural host for the agent causing citrus
leprosis. CiLV is reported only in the American continent (Argentina, Brazil,
Uruguay, Paraguay, the USA and Venezuela) but the vectors of leprosis (B. califor-
nicus, B. obovatus and B. phoenicis) are found widely in Europe, Asia, Africa and
American continent. However, citrus leprosis is always associated with infestation
of Brevipalpus spp. of mites (Childers et al. 2011).
The disease is characterized by round lesions on leaves, twigs and fruits which
do not become systemic. Severity of lesions varies with the type of citrus species. If
proper mite control is not done when disease appears, severe losses in yield may
occur. Fruits with lesions have low commercial value especially for fresh market.
But citrus leprosis because of its nonsystemic infection can only be important where
attacks by vector mites are significant. So any citrus plants being imported from
different nurseries should be free from leprosis lesions and come from nurseries
found free from and/or treated against Brevipalpus mites during the growing
season.
Management of Mite Pests on Citrus Spider mites feed primarily on leaves and
are capable of removing cell contents, causing cell destruction and reduction in
photosynthesis resulting in collapse of mesophyll cells and leaf drop. Mite damage
is particularly more when citrus trees are stressed by high spider mite infestations
alone or in combination with sustained dry and windy conditions that may occur in
fall, winter or early spring conditions. Spider mites prefer dry weather and low rela-
tive humidity in the range of 30–60 % and generally do not pose a problem in the
higher humidity conditions (McCoy 1977).
An average of five mobile spider mites/leaf corresponds to 70–80 % infestation

levels and constitutes a treatment threshold for processing fruit. In general the need
for controlling spider mites is based on temperature and humidity conditions, spider
mite levels, trees’ vigour and time of the year. From 5 to 15 mites/leaf have been
reported as economic threshold level for these mites. Petroleum oil provides some
ovicidal activity, but other miticides provide no significant ovicidal action. Pesticides
having comparatively longer residual activity can better kill the emerging larvae.
Monitor orchards actively during active period. Predaceous mites (mainly
Phytoseiidae), predaceous insects (lady beetles and six-spotted thrips) and a virus
specific to CRM are widespread in citrus-growing areas and contribute in biological
control of spider mites. Mites increase their reproduction on water-stressed trees.
Good irrigation practices reduce CRM outbreaks. Water roads limit the dust build-
up which also promotes mites.
Rust mites are important pests of fresh fruit; however, they also cause damage on
stems and foliage but fruit damage is the main concern. Generally mite population
and its damage are more on the north bottom of the tree canopy. The southern top of
the tree canopy is least favourable for mite population increase. The primary effect
of fruit damage caused by rust mite (russetting/bronzing) appears to be reduction in
grade. Other conditions that have been associated with severe fruit injury include
reduced size, increased water loss and increased fruit drop. Fruit growth and abscis-
sion are not affected until 50–75 % of the surface is injured. So there is reduced
justification for miticidal control of rust mites on fruits grown for processing. But
12.1 Fruits 239
for fresh markets, cosmetic appearance of the fruit is more important, and in such
cases 2–3 miticide applications at different intervals can be considered.
12.1.2.4 Coconut (Cocos nucifera)

Coconut is an evergreen tree and requires a full sun and a humid climate. They are
grown in all tropical regions of the world.
Coconut Mite (Aceria guerreronis) Eriophyid mite, A. guerreronis, is considered

as a serious pest of coconuts in Africa, tropical America and India. Feeding of the
mites causes physical damage and results in significant yield reduction. Mite colo-
nies are found under the perianth and over the button surface covered by the peri-
anth. Initially triangular pale-white or yellow patches close to perianth are observed
over the damaged nuts. Increased damage results in formation of brown-coloured
patches (ER9). Longitudinal fissures and splits occur on the outer surface of the
husk as the nuts grow in size. In severe cases, brownish gummy exudates are seen
oozing out from the affected portion, and the nuts are reduced in size or malformed.
Mites occur in large numbers in young buttons (2–6 months old). As the age of nuts
increases, mite population decreases. Mites are usually found under the bracts of the
fertilized female flowers. On queen palm (an ornamental palm), the mean genera-
tion time (egg–adult) of A. guerreronis has been reported as 30, 12 and 8 days when
reared at 15, 25 and 8.3 °C, respectively. Coconut mite appeared to be a serious
problem in nursery on queen palms. From 10 to 30 % loss of copra because of mite
feeding has been reported; losses are greater from early season infestations. In addi-
tion to damaging the fruit, A. guerreronis can kill coconut seedlings by feeding on
their meristematic tissues. Different control measures including cultural and bio-
logical controls (predaceous mites including acarofungus Hirsutella thompsonii)
may provide good control of mites and in reducing yield losses.
Red Palm Mite (Raoiella indica) Red palm mite is also known as the coconut
mite, coconut red mite, red date palm mite, crimson mite or scarlet mite in some
literature. Red palm mite has been reported as serious pest of coconut palm in some
countries, but has also been reported as an important pest on bananas, date palms,
and other palm species. Adult females are red with dark patches on the body. The
eggs are red, oblong and smooth, attached to the abaxial leaf surface by a slender
stalk in patches of 100–300 eggs. Young coconut palms appear to be most severely
damaged. Extensive yellowing of lower leaves because of mite feeding is some-
times noticed. Peak mite populations during June–September have been reported.
Other Mite Pests On coconut eriophyid mite, Colomerus novahebridensis (Keifer)

and Acathrix trymatus (Keifer) are reported from leaves of coconut palms resulting
normally in no noticeable damage. Incidence of spider mites, Oligonychus iseilemae
(Hirst), Tetranychus fijiensis (Hirst) and Tetranychus ludeni (Zacher), has also been
reported on coconuts but in uneconomic proportions.
12.1.2.5 Date Palms (Phoenix dactylifera)

Date palms are evergreen trees originated from the Persian Gulf area or from deserts
of North Africa. These trees do well in hot climates and are often grown in oasis in
desert areas.
Spider Mites Oligonychus afrasiaticus (McGregor) and Eutetranychus orientalis

have been reported infesting lower leaf surfaces of date palm from Egypt. Spider
mite, Eutetranychus palmatus Attiah, is also reported as a minor pest on date palm
and some other ornamental palms in some areas of Israel. These mites are reddish
brown in colour and are found sporadically throughout the year in date fronds and
infesting the fruit during midsummer. As the dates turn yellow, then these mites are
restricted to the fronds. They do not spin webs; mite feeding results in exudates that
become covered with dust as well as with the shed whitish exuviae of the mites.
Phyllotetranychus aegyptiacus Sayed have also been reported in high numbers on
leaves of the date palm, resulting in reddening of the upper surface of infested
leaves. Heavy mite infestations results in too much of webbings. High temperature
and limited rainfall favour mite development (Blimberg 2008).
Eriophyid Mite (Mackiella phoenicis Keifer) Eriophyid mite is also reported in

large numbers in emerging fronds and buds of the date palm causing malformation
of old frond; then the leaves become dry and fall off. False spider mites, B. phoeni-
cis and Raoiella indica, have also been reported on date palm.
12.1.2.6 Fig (Ficus carica)

Figs are deciduous trees grown mainly in Mediterranean countries (Iran, Pakistan
and Australia) and grow best in temperate or Mediterranean climate.
Fig Mite (Aceria fici Cotte) They are pale-yellow, wedge-shaped eriophyid mites.
Infests bud scales and young leaves, causing faint russetting of the leaves generally
in the interior portion of the canopy. Sometimes leaves drop and stunting of twigs
occurs because of mite feeding. Fig mite also transmits fig mosaic virus but virus is
not transmitted through the egg stage. Virus is only acquired during mite feeding
and is retained through moults. Treat only when damage on leaves is found.
Spider Mites (Tetranychus pacificus and T. urticae) Spider mites produce abun-
dant webbings on both sides of the leaves. Females of both the mite species over-
winter under the bark as females which develop orange-red colour. Heavy mite
feeding from spring onwards causes browning of leaves and even their defoliation
resulting in reduced vitality of fig trees. Vigorously growing trees are much more
tolerant to mite attack than trees which are under stress. So maintain trees through
adequate irrigations and reduce dust around fig trees to minimize mite population
development.
12.1 Fruits 241
12.1.2.7 Guava (Psidium guajava)

Guava is a subtropical shrub or tree mainly distributed in many subtropical areas.
The tree is evergreen in nature.
Tenuipalpid Mites B. californicus are reported feeding on bronzed fruits of guava

from India. Another tenuipalpid mite, Tenuipalpus pernicis Pritchard and Baker, has
been reported on guava plants from Kolkata (India). From Central America, B.
phoenicis is also reported causing browning and epidermal cracking of fruits; epi-
dermal cracking is also reported on infested leaves (Gould and Rage 2002).
Spider Mite (Eutetranychus orientalis) Spider mite is also occasionally found in

uneconomic proportions on the upper surface of leaves during April–June. Eriophyid
mite, Tegolophus guavae, is reported on guava from Florida (USA) causing pimple-
like structures on the fruit epidermis followed by fruit bronzing. Mites are also
found feeding on guava leaves. Broad mite, P. latus, is also reported on the terminal
leaf and flower buds which later turn into corky brown areas (Pena et al. 1999).
12.1.2.8 Jujube (Zizyphus mauritiana)

Jujube is a deciduous tropical fruit tree or shrub and seems to have been originated
from southern Asia (China).
Eriophyid Mite (Eriophyes cernuus Massee) Eriophyid mite also known as ber
gall mite is reported as a minor pest of jujube trees from India. Mites are commonly
found on the woody galls. On young plants small galls are found but quite big galls
are found on mature trees. Enormous mite populations are found when a small por-
tion of a fresh gall is disturbed under the microscope. Another tenuipalpid mite
Larvacarus transitans (Ewing) is reported causing blisters on jujube plants, and the
twigs of bushy cultivars were found heavily infested by the mite (Fig. 12.6). From
20 to 80 % of the bushes reported infested from Rajasthan (India). Twigs bearing
galls had less fruit yield than normal healthy twigs. The mite infestation appears on
twigs, in the form of minute galls (1.5–2 cm dia) in which mites enclosed within the
galls suck the cell tissues and affect the plant growth adversely. L. transitans is
found throughout the year in bushes except during the monsoon when the galls
rupture and the mites emerge for initiating new infestation.
Two-Spotted Spider Mite (T. urticae, TSSM) Sometimes TSSM infest jujube
nursery plants in screen houses affecting the growth of young plants. Oriental brown
mite (Eutetranychus orientalis) is also occasionally found in uneconomic propor-
tions on the upper surface of leaves during April–June.
12.1.2.9 Lychee (Litchi chinensis)

Lychee is a tropical and subtropical fruit tree native of Southern China, Taiwan,
Bangladesh and Southeast Asia.
Fig. 12.6 Twig galls made by eriophyid mites (a) Larvacarus transitans (b) Eriophyes cernuus
Litchi Erinose Mite (Eriophyes litchii Keifer) Litchi erinose mite measures
0.13 mm long and can only be seen through a high-powered magnifying lens. As a
result of mite feeding, leaflets become curled and distorted exhibiting a velvety-brown
appearance (Figs. 12.7 and 12.8). Mites begin their attack on new leaves at the onset
of growth flushes. Early indications of mite damage are wartlike swellings (about
1.6 mm dia) on the upper surface of the leaflets and light yellow spots on their lower
surface. The erinose mite damage seldom kills lychee trees, but the type of damage
is certainly unsightly (ER10). Pruning mite-infested foliage from the trees and by
destroying the pruned material, new mite infestations are avoided. The mite is easily
spread from nursery plants especially taken from infested trees. So use only clean
mite-free planting material.
12.1.2.10 Mango (Mangifera indica)

Mango is an evergreen tree that likes lowland tropical and subtropical areas. India,
China, Thailand, Indonesia, Mexico, Pakistan and Brazil are the leading mango-
producing countries.
12.1 Fruits 243
Fig. 12.7 Litchi leaf curl
Spider Mites (Oligonychus mangiferus and Oligonychus yothersi

(McGregor)) Commonly found on mango trees. The upper surface of leaves along with
the midrib areas is mainly infested by spider mites and the areas along the veins look red-
dish brown. Control measures during active season may be initiated if six or more spider
mites/leaf are found.
Mango Bud Mite (Aceria mangiferae Sayed) Mango bud mites are many times
reported on buds and are associated with mango malformation depending on the
presence of fungus, Fusarium moniliforme. When the fungus is absent and the buds
are attacked by the bud mites, a ‘witches’ broom’ develops on the terminal branch.
However, when fungus is present, then galls form on flowers and foliage. Another
eriophyid mite, Cisaberoptus kenyae Keifer, is reported producing ashy-white
membranes on the dorsal surface of mango leaves but seems to play no economic
role as very low mite incidence is reported (Fig. 12.9).
12.1.2.11 Olive (Olea europaea)

Olive is an evergreen tree, mostly grown in countries near the Mediterranean Sea
especially Spain, Turkey, Italy and Greece. Eriophyid mite, Oxycenus maxwelli
(Keifer), is commonly found in olive orchards. They are yellowish to dark tan, are
Fig. 12.8 Algal growth

and litchi leaf curl
Fig. 12.9 Mango leaves showing mite damage

12.1 Fruits 245
slow moving and have a wedge-shaped body. Generally these mites are not a major
problem on olive trees. Succulent stem and bud tissues are infested by mites; some-
times mites are also found on the upper surface of leaves. Mite-damaged leaves
appear sickle-shaped, but vegetative buds sometimes die because of mite injury.
Bud drop, discolouration of flower buds and blossom blasting are also reported as a
result of mite infestation. Treat before blooming if large mite populations are found.
12.1.2.12 Papaya (Carica papaya)

Papaya or pawpaw is an evergreen and originated from Mexico, Central America
and Northern South America but is now grown in most tropical regions of the world.
Cultivation of papaya needs a warm tropical climate.
Spider Mites (Tetranychus spp., E. orientalis, Oligonychus gossypii (Zacher)

and P. citri) Spider mites are found in pest form on papaya (Fig. 12.10). Feeding
by mites causes scarring and discolouration of the fruit and reduced the fruit size
significantly affecting its market value. Infestations usually begin on the older
leaves and then spread to the younger growth. Serious infestation of mites is found
during long hot and dry months. E. orientalis feeds on the upper surface of leaves
particularly between main veins and midribs.
False Spider Mite (Brevipalpus phoenicis) False spider mite is sometimes found
in uneconomic proportions on both leaves and fruits. Damage by mite feeding on
young papaya fruits is manifested by sunken areas.
Broad Mites (Polyphagotarsonemus latus) Broad mites are many times found on
terminal growth during warm and humid period. Mites feed on young leaves as they
Fig. 12.10 Spider mite damage on papaya trees

emerge from their growing points. Mite-affected leaves turn thick and are brittle
with down-curled edges. Severe mite infestations inhibit new stem growth with con-
sequent reduction in fruit production.
12.1.2.13 Passion Fruit (Passiflora edulis)

Passion fruit is a woody evergreen perennial vine, originated from Brazil and
Argentina, and grows in subtropical climate.
Spider Mites (Tetranychus spp.) Sometimes spider mites infest passion fruits.
Infested leaves exhibit tiny yellow or white speckles. Eventually leaves become yel-
lowish and may drop and sometimes may lead to complete defoliation. Heavily
mite-infested plants become stunted and may also result in vine dieback, shrivelling
and dropping of immature fruit.
Broad Mite (P. latus) Broad mite has been reported as an important pest of passion
fruit in Kenya. Mite feeding produces discolouration, necrosis of tissues and defor-
mation. Initial attack occurs on stems of terminal shoots and young terminal leaves
which remain stunted and deformed. Stems of terminal shoots may become slightly
swollen, roughened or rusted. Attacked fruits are deformed and show white to tan or
brown scars on their skin.
12.1.2.14 Pineapple (Ananas comosus)

Pineapple is a herbaceous perennial plant, originated from Brazil and Paraguay, and
likes tropical climate. Pineapple mite (Schizotetranychus asparagi (Oudemans)) is
widely distributed and sometimes causes severe damage to recently established
plants in the field. Plants infested in early stages remain small and fruit production
is either curtailed, and sometimes heavily infested plants may die before producing
fruit. Pineapple can be managed in a better way by planting only mite-free seed
plant material.
Tarsonemid mite, Steneotarsonemus ananas Lindquist, is sometimes found
infesting pineapple later in the plants’ phenological cycle. Red mite, tenuipalpid
mite and Dolichotetranychus floridanus (Banks) are also reported feeding on pine-
apple in large populations and sometimes severely damaged leaves and young
plants. D. floridanus can cause significant problems during establishment and early
growth stages of crop.
12.1.2.15 Pomegranate (Punica granatum)

Pomegranate is a deciduous shrub or small trees, originated from Iran and widely
cultivated throughout the Mediterranean region of Southern Europe, tropical Africa,
Indian subcontinent and other parts of Southeast Asia.
Flat Mite (Brevipalpus lewisi) Mite feeding causes leathering or ‘alligator skin’
damage to pomegranate rinds, similar to their damage in citrus and pistachios.
Occasionally mite damage can be quite serious. Flat mite damage starts at the stem
end, and their brown discolouration extends further than the cracking. Mites and
12.1 Fruits 247
their cast-off skins may also be found in the cracks of the fruit. Flat mites, Brevipalpus
lewisi McGregor, overwinter under the flakes of the bark on large branches and later
move to leaves and fruits in the summer.
Red Spider Mite (Oligonychus punicae) Red spider mite is reported from tropical
Asia and Central and South America. Oligonychus punicae (Hirst) is reported
infesting during flowering and fruiting stages of the crop. Mite feeding is mainly
confined to the upper surface of leaves along the midribs and smaller veins and is
reported as a minor pest doing little damage to pomegranates.
Eriophyid Mite (Aceria granati (Canestrini and Massalongo)) Mite feeding is

confined along edges of leaves resulting in downward curling of the whole length of
the leaf margin. The mites are found wrapped in the thickened turned-over leaf
edges (Fig. 12.11).
False Spider Mites (Tenuipalpus granati Sayed) False spider mites are reported
on pomegranate fruits from Iran and some neighbouring parts. Another false spider
mite (Tenuipalpus punicae Pritchard and Baker) is reported causing defoliation and
development of some spots on fruits resulting in cracking of fruits (Fig.12.12). T.
Fig. 12.11 Spiral curling

of pomegranate leaves
Fig. 12.12 Tenuipalpus

punicae damage on
pomegranate leaves
punicae is also considered as the most serious pest of pomegranates in Iraq. There
are eight generations in a year. Females overwintered under the bark and in crevices
of tree trunks and branches. Peak population is reported during May to July.
12.2 Vegetables
Different vegetable crops are classified based on the edible parts of plants (under-
ground/above ground part), temperature (cool season/warm season) and botanical
basis (families). Knowing the family classification of different vegetable crops
plays an important role in the prevention of mites as many crops in the same family
are often hosts to the same mite species. While planning a rotation scheme for dif-
ferent vegetables, it is important to rotate families but not just species. Here, for the
sake of convenience, discussion of mite problems and their management is being
done under different family groups.
12.2 Vegetables 249
12.2.1 Family Alliaceae
12.2.1.1 Onions (Allium cepa) and Garlic (Allium sativum)

Onion is the most widely used vegetable in the world and is grown mainly in India
and China. Garlic is a perennial plant but usually grown as an annual and can be
grown year round in mild climates. The highest production of garlic and onion is in
China and India. Onions are grown in different climates but prefer cool season.
Bulb Mites (Rhizoglyphus robini Claparede and Tyrophagus spp.) The mite is
shiny creamy white bulb-shaped (0.5–1 mm long) with its legs moved forwards and
a bulbous rear with many long fine hairs. The mouthparts and legs are purplish
brown while the main body is creamy white. Heteromorphic deutonymphs form
when there is overcrowding or the area in which they are feeding deteriorates. There
is high risk of spreading and establishing this mite in new areas. These mites gener-
ally occur in clusters inhabiting damaged areas under the roof plate of onion bulbs
or garlic cloves. Bulb mites have a wide host range and feeds on many kinds of
bulbs, roots and tubers and can also infest bulbs of onion, garlic, lilies, gladiolus,
cereals and many other crops in the storage or in the field. They can survive on
decaying vegetations in the field until it is completely decomposed. Bulb mites
damage bulbs by penetrating the outer layer of the tissue and allowing access to rot-
ting organisms. They are very serious when plant growth is slowed by cool and wet
weather. As a result of their infestation, there may be reduced plant stands, stunted
plant growth and rotting of bulbs in the storage. On seeded onions, mite infestation
can cut off the radicle before the plants are established. A female can live up to
40 days and produce 700 eggs under optimum conditions; bulb mites can increase
rapidly under field conditions and a generation can be completed in about 4 weeks.
The best way to determine whether these mites are present is to carefully dissect
the region where the roots and bulb come together. The mites can also be found
under one or two layers of scales at the lower end of the bulb. These mites survive
in the soil or on organic matter left behind from previous crop. Avoid planting suc-
cessive onion or garlic crops. Flood irrigation or heavy rains during the winter may
reduce mite levels in the soil. Garlic growers must insist on planting clean seed
cloves. Treatment of seed garlic with hot water before planting reduces mite infesta-
tion (Capinara 2011).
Wheat Curl Mite (Eriophyes tulipae Keifer) E. tulipae is primarily a pest of

stored bulbs and causes stored onions and garlic to desiccate. These mites may also
serve as vector of the rot-causing organisms under both the field and storage condi-
tions. Heavy infestations under field conditions may result in streaking and twisting
of garlic leaves and may reduce plant stand and crop yields (ER19). Flood irrigation
or winter rains may reduce mite incidence under field conditions. Avoid planting
successive onion or garlic crops to manage wheat curl mite infestation (Gill et al.
2006).
12.2.2 Family Convolvulaceae
12.2.2.1 Sweet Potato (Ipomoea batatas)

Sweet potato originated from North and South America, but the highest production
is in China. It is grown mainly for the sweet-tasting tuberous roots.
Vine Spider Mite (Tetranychus marianae McGregor) Vine spider mite feeds on
the undersurface of leaves and causes the area around the feeding punctures to
become chlorotic and appear as conspicuous whitish to yellowish stipplings. In case
of heavy infestation, photosynthesis is greatly reduced and chlorotic areas coalesce
to form mottled yellowish interveinnal patches. Infested leaves later turn yellow and
may become brown and scorched and may drop prematurely. Eggs are hyaline and
later turn translucent white. Adults are greenish on emergence and later may look
carmine red with dark lateral markings as they develop. This mite is polyphagous in
nature. Two-spotted spider mite (TSSM) also sometimes appears in high infesta-
tions causing minor damage on sweet potato.
False Spider Mites (Brevipalpus californicus) Infestation occurs during the dry
season and may cause whitish feeding punctures. B. californicus has a broad host
range.
Eriophyid Mite (Aceria sp.) Beginning at the shoot tip, infested vines become
excessively hairy and more prevalent during hot and dry season. Some undescribed
eriophyid mites producing unsightly galls of irregular sizes on vines and leaves are
reported from the Philippines.
12.2.3 Family Cruciferaceae
12.2.3.1 Cauliflower (Brassica oleracea)

Two-spotted spider mite (TSSM) has been reported feeding in sporadic proportions
during the period near harvest. Population of mobile mites ranged from 800 to 4800
per leaf on both the ventral and dorsal surface of leaves. As a result of severe infesta-
tion, smaller heads started drying and the larger heads did not develop properly.
Normally cruciferous vegetables are not preferred by mite for feeding and develop-
ment (Dhooria 1976).
12.2.4 Family Cucurbitaceae
Cucurbits (watermelon, summer and winter squash, cucumber, pumpkin, canta-

loupes, etc.) are one of the most common vegetables.
12.2 Vegetables 251
Spider Mites (Tetranychus urticae, T. pacificus, T. turkestani (Ugarov and

Nikolskii), T. desertorum (Banks) and T. cinnabarinus (Boisd.)) These mites
many times are found infesting cucurbitaceous crops particularly during hot and dry
months. As a result of mite infestation, initially leaves become pale and stippled but
in later stages of infestation, the entire crop may dry up and die (Fig. 12.13). Mites
are most problematic on musk melon, watermelons and cucumbers but lesser on
other cucurbits (Figs. 12.14, 12.15 and 12.16). Chlorophyll loss is quite pronounced
on the undersurface of leaves before it becomes apparent on the upper surface of
leaves. High mite infestation cannot be tolerated and can result in lowered yields
and reduced quality of the fruit. Spider mites have broad host range, so monitoring
on weeds and other hosts is most important for reducing mite infestations on cucur-
bits. Mites preferred watermelon, summer squash, ash gourd and squash melon, but
least feeding preference for wild melon, cucumber and long melon is reported. Do
not use broad-spectrum pesticides for other insect pests as they significantly inter-
fere in natural control of spider mites. Control mite build-up on different broad-
Fig. 12.13 Spider mite-damaged cucurbit leaves (a) watermelon (b) musk melon (c) cucumber
(d) ash gourd
Fig. 12.14 Spider mite-damaged watermelon field
Fig. 12.15 Spider mite-damaged musk melon field
leaved weeds growing along edges of a cucurbit field as they may help in dispersal
of mites to the main crop. Minimize dust around cucurbit fields. Sometimes water
sprays on dusty paths and along margins of the field can help in preventing mite
build-up on cucurbits. Initiate sprays against mites before development of pro-
nounced webbing symptoms or when the runners are about 40 cm in length.
Tarsonemid Mite (Polyphagotarsonemus latus) In the USA, P. latus mites are

also reported as a potential pest of watermelon and musk melon.
12.2 Vegetables 253
Fig. 12.16 Spider mite-damaged cucumber field
12.2.5 Family Leguminosae
12.2.5.1 Beans (Both Phaseolus spp. and Vicia spp.)

Spider Mites (Tetranychus cinnabarinus, T. pacificus, T. turkestani and T. urti-
cae) Spider mites are commonly found during hot and dry periods (April–July). If
the plants are stressed, mite damage is very severe and significant yield reductions
may occur. Heavy rains can eliminate stress on the plant. Scouting for spider mites
and their damage symptoms is very important. However, sometimes spraying for
aphids with pyrethroid pesticides may aggravate the mite infestations. Generally
mite infestations are commonly found in border regions of the field. But if the mite
infestations are found in the interior of the fields, then the entire field should be
sprayed. Mite damage can be rated after seeing stipplings and webbings on lower
leaves. High mite population is found in the middle and lower canopy, but mite
damage during full pod and beginning seed stages determines the per cent yield
loss. So treatment against mites is only recommended in cases when mites and their
damage are detected throughout the field. Check fields every 4–5 days if drought
persists as mite infestations can develop into damaging proportions quickly in such
situations.
12.2.5.2 Peas (Pisum sativum)

Spider Mites (Tetranychus spp.) Mite feeding causes small yellow specks on
leaves; however, if found in large numbers, the leaves may dry and fall off. Infested
plants have poor growth. Monitor regularly the crop to find out the presence and
level of infestation. Spider mites sometimes cause damage in later stages. Conserve
their natural enemies by avoiding the use of broad-spectrum pesticides. The crop
should be irrigated properly. Do not plant a new crop near an infested field, and also
destroy crop residues immediately after harvesting.
12.2.6 Family Solanaceae
12.2.6.1 Brinjal (Solanum melongena)

Brinjals are also commonly known in some countries as aubergines or eggplants.
They are perennial but are often grown as an annual. Cultivation requires a tropical
or subtropical climate and grown in temperate areas during the warmer part of the
year.
Spider Mites (Tetranychus urticae, T. macfarlanei and Eutetranychus orienta-

lis) These mites are commonly reported on brinjal leaves. Both T. urticae and T.
macfarlanei Baker and Pritchard are reported to cause sometimes too much damage
during hot and dry periods (Fig. 12.17).Brinjal crop kept as ratoon is more severely
attacked as compared to the normal crop (Fig. 12.18). T. urticae is also a serious
pest of tomato, French bean, cucurbits and some other field crops. In case of severe
mite infestation, leaves will completely desiccate and drop. Mite injury is aggra-
vated if plants are stressed. Overhead irrigation and heavy rains can significantly
reduce mite infestations. Avoid early season, broad-spectrum insecticide applica-
tions for other insect pests as they interfere with the natural control of spider mites.
Do not overfertilize the fields, and check the weak plants in and around the field to
avoid carry-over of mites in the brinjal crop. Use only selective acaricides whenever
it is necessary (Ravenberg et al. 1983; Srinivasan 2009).
Tarsonemid Mites (Polyphagotarsonemus latus) and eriophyid mite (Aceria

lycopersici) are also sometimes found in high proportions but they seem to have
lesser economic relevance (Fig. 12.19).
12.2.6.2 Capsicums (Capsicum annum)

Capsicums include chillies, red pepper or hot pepper. Bell pepper (C. frutescens) is
a cool climate crop (17–22 °C) and its production is low during summer months.
Broad Mites (P. latus) They are quite small, less than 300 μm and oval in shape;
eggs are clear and oval-shaped with the surface covered in white tubercles making
a spotted pattern and are usually found in the growing tips on the underside of newly
formed leaves and under the calyx of flowers and fruit or in other protected depres-
sions (ER12). The male is smaller and faster moving than the females. The life cycle
from egg to adult takes between 4 and 10 days depending on temperature. From 7
to 12 eggs per female are laid. Adult broad mite lived from 5 to 13 days depending
upon the favourable conditions for growth and reproduction (about 24 °C and high
humidity). Reproduction does not occur below 13 °C and above 34 °C. Under
favourable conditions, 20–30 generations may occur annually. Broad mite is a major
pest and can cause severe damage in warmer areas on capsicum plants, but is a
12.2 Vegetables 255
Fig. 12.17 Grades of spider mite-damaged brinjal leaves
Fig. 12.18 Spider mite-damaged brinjal field
minor pest in temperate areas. Mite attacks mainly young growth of capsicum and
feeds by injecting toxins from their saliva. A few mites can cause a lot of damage;
mite infestation is quite common in greenhouses on many summer-grown vegeta-
bles and ornamentals. Mite damage is quite significant during warm and humid
conditions (80–90 % R.H.) and temperatures above 25 °C (Gerson 1992; Karmaker
1995).
Fig. 12.19 Broad mite-damaged brinjal leaves (healthy vs. mite-damaged)
Mite feeding causes leaves to bronze and thicken, brittle, corky or cupped down-
wards and narrower than normal (Fig. 12.21). Heavy mite feeding causes young
terminal buds to die and drop off. More than 45 host plants have been reported from
different areas of the world. Monitoring broad mite infestation from early stages of
crop helps in careful crop protection initiatives. Avoid introducing infested nursery
plants into the crop; also use a fallow period when possible to clear pest popula-
tions. Use nursery plants that have grown away from infested areas to have a clear
crop. Mite control is simple and less expensive when plants are young and spray
coverage is not an issue.
Two-Spotted Spider Mites (T. urticae, TSSM) Two-spotted spider mites over-
winter as females on crop debris. Eggs are laid on the undersurface of leaves or on
protected plant parts. Nymphs and adults puncture the leaf tissue and extract plant
juices; leaf injury first appears as white stipplings or small white blotches which
fuse together into larger yellow patches (ER11). Under hot and dry conditions when
infestation is very heavy, leaves dry out and turn brown (Fig. 12.20). Mite develop-
ment is quite rapid and severely stunt appearance of the fruit rendering them unmar-
ketable. For monitoring the presence of mites, a minimum of two leaves and two
fruits on five plants in ten locations of the field should be examined. Although no
exact thresholds have been established, a treatment may be needed if 10 % of the
leaves or fruits are infested with mites. Intelligent use of pyrethroids for other pests
should be done as their excessive use can cause spider mite outbreaks (Hofan et al.
2004).
12.2 Vegetables 257
Fig. 12.20 Spider mite-damaged bell pepper field
12.2.6.3 Potato (Solanum tuberosum)

Potatoes are perennial plants grown mainly in China, India and Russia and origi-
nated from the Andes region in South America. The tubers of the potato plant are
used as main staple food.
Two-Spotted Spider Mite (T. urticae, TSSM) Two-spotted spider mite is occa-
sionally reported in outbreaks on potatoes from different potato-growing regions.
The mites injure potato leaves by puncturing the surface cells producing initially
small yellow spots that later darken to reddish brown and may affect most of the
foliage and result in reddish-brown patches of affected plants. Injury is most com-
mon in hot and dry weather and seldom occurs before mid-season. But mite damage
is rarely found in cooler areas. In some cases, it is necessary only to treat a strip
along the edge of the field. Sprinkler irrigation helps limit mite damage by increas-
ing humidity in the plant canopy making conditions less favourable for the mites
(Capinara 2006).
To decide whether to spray, early detection of TSSM should be done. Application

of non-selective pesticides such as pyrethroids, certain carbamates and organophos-
phates which negatively impact mite predators sometimes flares up mite population.
Proximity of potato fields to dusty roads during hot and dry weather promotes mite
outbreaks. Generally mites are an induced problem, brought on by management
practices aimed at other pests. Spider mites overwinter in leaf litter and other debris
found on the soil surface. Application of miticides should begin before mite popula-
tions reached two mites per leaf. Once populations exceed five mites per leaf, con-
trol with miticides becomes difficult. Thorough coverage is essential for good mite
control, and leaves should be particularly dry at application time. If possible,
Fig. 12.21 Broad
mite-damaged capsicum
twig
irrigation should not be applied for 24 h after pesticide application. Mite population
is generally controlled after single application; however, if second application is
desired, then it should be made with a different pesticide having different mode of
action.
Tarsonemid Mite (P. latus) Minor infestation of P. latus on potato is reported dur-
ing June–September period in Dharwad and Punjab (India).
12.2.6.4 Tomato (Lycopersicon esculentum)

Tomato is perennial, but often grown as an annual in a wide range of climates from
tropical to temperate. Central and South America are the regions of origin of tomato.
India, Turkey and Egypt are important tomato-producing countries.
Tomato Russet Mite (Aculops lycopersici (Wolffenstein)) Tomato russet mite is

sometimes found in hundreds on green leaves immediately above the damaged
bronzed leaves and feeds on leaves, stems and fruits. Damage usually starts near the
ground; the colour of infested leaves sometimes changes to greasy bronze or rust-
coloured. If uncontrolled, mite infestation may kill the plants (ER14). So monitor-
ing of mites is important if damage is noticed or bronzing is observed. Russet mites
12.2 Vegetables 259
Fig. 12.22 Spider mite

damage on tomato
may be present year round on the undersurface of leaves but are more abundant
during hot and dry weather (April–June). Petunia, Datura stramonium, field bind-
weed and some other solanaceous plants like brinjal are also found as hosts of russet
mites.
Two-Spotted Spider Mites (T. urticae, TSSM) Infestations of these mites usually
start on the field edges. The upper leaf surface shows speckling or mottling appear-
ance. Mites overwinter on many weed plants. But for their rapid multiplication, hot
and dry weather is most congenial. Dusty conditions also favour mite activity
(Fig. 12.22).
12.2.7 Family Malvaceae
12.2.7.1 Okra (Abelmoschus esculentus)

Annual or perennial, warm and tropical climates are suitable for okra; it is a heat-
and drought-tolerant crop.
Spider mite (T. urticae). Spider mite is a potential pest of okra. Photosynthesis is
reduced as a result of mite feeding, and stressed conditions aggravate mite
Fig. 12.23 Spider mite damage on okra leaves
population (Fig. 12.23). Blooms abort and leaves become discoloured and thick-
ened. Synthetic pyrethroids interfere with natural control and flare up mite infesta-
tion. Broad-spectrum pesticide sprays against other pests also result in mite
outbreaks on okra crop. Broad mite (P. latus) in some regions is also reported as an
important pest of okra.
12.2.8 Family Umbelliferae
12.2.8.1 Carrot (Daucus carota)

Carrots are biennial plants; the highest production is in China, Russia and the USA
and is common in many European countries. Sometimes two-spotted spider mites
(T. urticae) appear as an important pest on carrot during late summer. Chemical
control of leafhoppers on carrot by too much use of pyrethroids seems guilty of flar-
ing up spider mites. Low infestation of brown wheat mite, Petrobia latens, is also
sometimes found on carrots grown near a rain-fed wheat crop.
12.2.8.2 Celery (Apium graveolens) and Parsley (Petroselinum

crispum)
Celery is a biennial plant but often grown as an annual and grows best in temperate
climates with mild or cool weather. Two-spotted spider mite (T. urticae, TSSM)
sometimes develops rapidly during hot and dry weather. As a result of mite infesta-
tion, whitish stipplings may be found on the upper surface of leaves, while on lower
leaf surfaces, too many mites and their webbings are commonly observed. TSSM
occasionally appears as a minor pest, but sometimes TSSM are troublesome during
the later part of the crop season when their presence on harvested product may
12.2 Vegetables 261
prove troublesome for market purposes. Dry and dusty conditions may favour
development of spider mites; water-stressed plants are more susceptible. Overhead
irrigation may help alleviate mite problems. Avoid use of excessive nitrogen.
12.2.9 Other Vegetables
12.2.9.1 Artichoke (Cynara scolymus, Asteraceae)

Artichoke is a perennial plant and prefers a Mediterranean climate or areas that get
no frost in winter and have cool and foggy summer weather. It is grown mainly in
Italy, Spain, Egypt, France and Morocco.
Two-spotted spider mite (T. urticae, TSSM) appears occasionally in pest form on
artichokes and causes economic damage. But because of their small size, they are
observed only when high mite infestations are found on artichoke leaves. As a result
of mite feeding, yellow stipplings are found on the upper surface of leaves along the
large veins of leaflets. As the damage increases, the interveinnal leaf area begins to
yellow, later turns brown and dries up. In case of serious infestations, mites spread
to buds. Infested buds have white spots on the surface and are covered with web-
bings of mite. TSSM appears in the form of serious infestations when crops are
water stressed. Mite management is aided by good crop hygiene.
12.2.9.2 Amaranth (Amaranthus spp., Amaranthaceae)

Amaranth is a plant grown all over the world especially in warmer climates.
Spider Mites (Tetranychus spp., Mononychellus sp., Oligonychus spp.) Spider

mites are reported feeding on amaranth plants resulting in lowered plant growth,
flowering and seed yield. Damage is most severe when mite attacks young plants;
mite damage may be particularly severe during the dry season. Avoid use of
broad-spectrum pesticides particularly pyrethroids, and also avoid planting the new
crop near the infested fields. Use overhead irrigation or wash plants with a strong jet
of water to knock down mites and destroy their webs.
12.2.9.3 Mentha spp. (Mentha spp., Lamiaceae)

Mint is commonly used in Indian cuisine; its fresh leaves are mainly used as a culi-
nary source. Mint grows quickly as a network of runners. Spearmint (Mentha spi-
cata, M. cardiaca) and peppermint (Mentha piperita) are grown commercially for
mint oil which is sometimes used in flavouring chewing gum, candy, toothpaste and
medicines.
Spider Mites (Tetranychus urticae) Mite feeding causes leaves to yellow, bronze
and fall under heavy infestations, thereby reducing oil yields. Mite population at
five or more mites per leaf can significantly result in oil reduction if uncontrolled.
Patchy distribution of this mite is reported from Punjab (India) during March–June
when the Mentha crop was near harvest. M. piperita suffered a maximum damage;
M. arvensis was also reported as a favourable host, but M. citrata was reported as an
unfavourable host. Fall ploughing and fall and spring flaming tend to delay spider
mite build-up by 3–4 weeks in the early season. The use of some organophosphate
insecticides may stimulate or increase spider mite populations by killing predatory
mites or even stimulating spider mite reproduction. Sporadic incidence of false spi-
der mites (Brevipalpus spp.) on Mentha is reported from Punjab (India) but mites
did not cause any significant damage. Sometimes the damage done by mites
remained unnoticeable.
Floridatarsonemus spp. have been found as a pest of peppermint in some areas

of the USA; mite feeding may result in a condition known as ‘squirrely mite’. The
mite infestation is sometimes severe on muck soils on old stands. It also infests the
mint buds but produces no symptoms until late in the season. The upper two or three
internodes are usually shortened and the new terminal leaves are reduced in size.
Hay weight is not affected but oil yield may be reduced by up to 80 % in mite-
infested fields. This mite may occasionally be found on spearmint but does not
develop in damaging proportions on this crop.
12.2.9.4 Spinach (Basella alba)

Spinach is perennial and grown in warm tropical climates, but in cooler areas it can
be grown as an annual during the warmer part of the year.
Spinach crown mite (Rhizoglyphus spp.). Spinach crown mites are almost trans-
parent mites with prominent long hairs/setae. They are found on soil and above
ground deep in the crown of spinach plant. Eggs are spherical and transparent and
laid on the innermost part of the plant. Crown mites damage sprouting seeds, seed-
lings before or after emergence, sometimes found on older plants. Feed primarily on
newly expanding leaves at the heart of the plant. The ability of the mite to damage
grown up plant is reduced. Because of mite damage, deformed leaves or small holes
in the expanding leaves may be found. Mite damage is comparatively more during
cool and wet conditions when soils are too high in organic matter. Good sanitation
of fields including prompt destruction of harvested fields may reduce mite
populations.
12.2.9.5 Sweet Corn (Zea mays, Poaceae)

Sweet corn also known as sugar corn is a variety of maize with a high sugar content.
Unlike field corn varieties which are harvested when the kernels are dry and mature
(dent stage), sweet corn is picked when they are immature (milk stage). They are
prepared and eaten as a vegetable rather than as grain.
Banks grass mite (Oligonychus pratensis (Banks), BGM) suck sap from the
leaves producing light fleckling wounds at the feeding site. In high populations,
BGM will cause leaves to prematurely die, although it is uncommon in sweet corn
that is picked for fresh market. The greatest injury is to the appearance of husks,
which can be seriously bleached and discoloured by BGM feeding. Sweet corn
grown for seed is more commonly damaged by BGM since mites have more time to
develop on maturing plants. Reduction in yield (seed weight) from leaf damage can
be caused by BGM. Problems with BGM often result from the use of certain
12.3 Mite Pests of Floricultural Plants 263
pesticides that destroy their natural enemies but do not properly affect BGM. Sweet
corn fields should not be located adjacent to small grains particularly winter wheat.
Large grassy areas adjacent to sweet corn fields will also increase risks of BGM
problems in sweet corn. In fields where BGM problems are common, mites need to
be controlled prior to tasseling stage. Control is best achieved when pesticide appli-
cations are made to control small colonies. However, outbreaks of mite can be dif-
ficult to control.
12.2.9.6 Taro (Colocasia esculenta, Araceae)

Taro is the most important food throughout the Hawaii Island. Two-spotted spider
mites (T. urticae, TSSM) are important pests of taro, especially during dry condi-
tions. Initially tiny spots and mite webbings on the upper leaf surface are signs of
the presence of mites. When mite infestations are severe, the speckling is seen all
over the leaf. Mite webbings occur on the undersurface of leaves, as the mite infes-
tation advances the leaves turn yellow and die prematurely. Okra, papaya, sweet
potato, cassava, beans and cucurbits are other important plant species infested by
the same spider mite. Managing spider mites on taro requires intelligent use of pes-
ticides without killing their predators. Proper irrigations should be applied to avoid
water-stressed plants which stimulate mite outbreaks. Sometimes Rhizoglyphus
spp. are also found on corms of taro but no significant information about them is
available.
12.3 Mite Pests of Floricultural Plants
Several species of ornamental plants are grown as annuals under both the field and
protected conditions, as perennial in houses and along roads for their shade and
beautiful flowers, and under different other landscapes. Many species of spider
mites, eriophyid mites and other mites are reported on different plant species in dif-
ferent regions. As most of the mites infesting different ornamental plants have wide
host range, so a brief account of these mites is given below, and the information may
be utilized for different management strategies on different ornamental plants
depending on the climatic conditions and method of cultivation. Some species of
different eriophyid mites are of great economic importance on many ornamentals as
they cause several deformities like bud blisters and galls. The leaf shape may be
abnormal, distorted, russeted, stunted and destroyed, and mite-damaged bulbs may
be destroyed. The health of plants emerging from infested bulbs is affected. New
growth is deformed and distorted. Affected leaves may be thickened, strap-type and
reduced in size and may exhibit signs of scarring or bronzing (Dreistadt 2001;
Naegele and Jefferson 1964).
12.3.1 Bulb Mites (Rhizoglyphus spp.)
Bulb mites are slow moving and relatively large in size (0.5–0.9 mm long) and are
pearly white and smooth with short reddish legs. From shipments of infested bulbs,
these mites have spread to different parts of the world. Bulb mites occur as large
colonies, not as individuals. A female produces about 100 eggs that are deposited
near the injured or decaying tissue. At 25 °C, egg–adult development takes about
12 days. The development of mite is mainly affected by temperature, humidity and
hosts. In addition to feeding damage, mites provide access to fungal root pathogens
(such as Pythium, Rhizoctonia and Fusarium). Mites are usually secondary pests of
bulbs, but can be responsible for serious losses. Dahlia, freesia, gladiolus, hyacinth,
iris, narcissus, orchids and tulips are infested by bulb mites. Steam pasteurization
has been reported to eliminate bulb mites from the soil. In the storage, low relative
humidity should be maintained to reduce potential for mite infestations. Predaceous
mites Cosmolaelaps claviger and Hypoaspis miles have been reported feeding
actively on bulb mites (Gerson and Weintraub 2007; Dhooria 1999a, b).
12.3.2 Cyclamen Mites (Steneotarsonemus pallidus (Banks))
They are tiny (0.13 mm), colourless mites found during cooler period when relative
humidity is high. Egg–adult development is completed in 10–14 days. Mites feed on
all parts of the plant but young foliage is most affected. Mites avoid light and feed
on unopened leaflets and buds, resulting in wrinkled deformed leaves; buds may or
may not open and produce distorted blooms. African violets, azalea, begonia, chry-
santhemum, cyclamen, dahlia, delphinium, fuchsia, gerbera, geranium, impatiens
and snapdragon are the favoured hosts among floricultural plants. Mite-affected
plants cease blooming and have shortened internodes, distorted leaves and stems
with blasted buds. Mite infestations are mostly localized and can be examined
through dissecting a tissue under a microscope. If only a few plants are affected by
mites, then discard them so that mites may not disperse to healthy plants. Mite
infestation levels can also be reduced by decreasing humidity, and on non-blooming
plants, mites can be killed through immersion of potted plants for 15 min in water
heated to 43 °C. For chemical control, use sufficient spray volume and pressure of
recommended pesticides covering thoroughly both surfaces of the leaves.
12.3.3 False Spider Mites (Brevipalpus spp.)
These mites are flattened, very small (0.3 mm long), reddish and slow moving. Eggs
are reddish and somewhat flattened, with an incubation period of about 3 weeks.
Egg-to-adult development takes about 5–6 weeks; the life cycle is about four to five
times longer than two-spotted spider mite (T. urticae, TSSM). Mite-infested plants
initially develop mottling symptoms and later develop a silvery appearance. In case
of severe infestation, the plant tissue turns brown and dies; sometimes mites may
also attack the stem of the plants. They are commonly found on orchids and on
many other ornamental plants (Childers and Rodrigues 2005).
12.3.4 Lewis Mite (Eotetranychus lewisi (McGregor))
Lewis mite is primarily a pest of poinsettias. They are smaller than TSSM but have
several small spots on their body as against two large spots in case of TSSM. The
developmental time from egg–adult is 12–14 days at 21 °C. Females lived for about
25–30 days and laid eggs at the rate of two to three per day. Mite damage symptoms
are nearly similar to the damage caused by TSSM. Mites feed on the undersurface
of leaves by piercing the epidermis and sucking cell sap. Initially infested leaves
show stippled appearance on the upper surface of leaves; eventually the entire leaf
becomes bleached and fall off. These mites also spin webbings, but not as extensive
webbings as spun by TSSM; no hibernation stage or diapause has been reported.
Scouting of Lewis mite is essential when plants appear to lose green colour or have
bleached foliage. In general, management and control methods are similar to those
of TSSM.
12.3.5 Two-Spotted Spider Mite (Tetranychus urticae, TSSM)
TSSM is an important pest of cut roses and orchids under polyhouse/greenhouse

conditions (ER18). They are probably the worst overall mite pests of ornamental
plants and have very broad host range. TSSM thrives in hot dry weather and seems
to have a particular affinity for water-stressed plants. Predacious mites, Phytoseiulus
persimilis (Athias-Henriot) and P. macropilis (Banks), are very effective against
TSSM; predaceous thrips are also effective against these mites. Broad-spectrum
insecticides like carbaryl and pyrethroids kill natural enemies and enhance mite
outbreaks. Most of the pesticide applications in polyhouses growing cut roses are
directed against TSSM (Kaur et al. 2006a, b, c). Continuous use of synthetic pesti-
cides to control mites leads to resistance to most of the pesticide compounds. Mites
extract the cell contents from the leaf parenchyma tissue, causing foliar stippling
and disruption of the plants’ photosynthetic activity and water balance mechanisms
(Dhooria 1999a, b). TSSM also produces extensive webbings on foliage and high
densities of mites can move onto flowers rendering them unmarketable (Figs. 12.24
and 12.25). High densities of mites cause sometimes significant damage to flowers;
however, at much lower densities, visible injury and physiological effects on plants
are only observed. Carnations, marigolds, orchids and many other annuals and
perennial ornamental plants are sometimes infested by TSSM resulting in great
losses (Figs.12.26, 12.27 and 12.28).
Removal of mite favourite weed plants and other susceptible hosts surrounding
greenhouses provides a good cultural control of TSSM. If one is selling whole
plants as opposed to cut flowers, it is better to discard heavily infested plants but
make sure incoming plants are free of mites before placing them near other crops
Fig. 12.24 Grades of mite-damaged rose leaves
Fig. 12.25 Webbings of

spider mite on rose flower
and leaf
Fig. 12.26 Spider
mite-damaged dahlia plant
Fig. 12.27 Spider mite-damaged marigold plant

Fig. 12.28 Spider
mite-damaged celosia plant
(Naegele and Jefferson 1964; Hamlen 1978; deMoraes and Tamai 1999). TSSM
thrives well under dusty conditions, so by directing a forceful spray of water at the
plants is a good practice to remove dust and in reducing mite populations. But this
practice should be done enough before nightfall so that plants can dry and resist
fungal/bacterial diseases. Chemical control applications should be done only after
considering economic threshold values. Try to select a chemical that does not affect
natural enemies of mites, but may control different stages of TSSM. Be sure to have
spray cover on all parts of plants mainly the underside of leaves which are most
preferred by mites.
12.3.6 Broad Mite (Polyphagotarsonemus latus)
Broad mite is commonly known as tropical mites and belongs to the same family of
mites as the cyclamen mites. Broad mites cause same type of damage as do cycla-
men mites. Puckering, curling or wilting symptoms on leaves because of mite feed-
ing are commonly observed. A female in her lifetime may lay up to 20 eggs, and the
life cycle is completed in 4–6 days (Dhooria and Bindra 1977; Ahmed et al. 2000).
Broad mites attack growing points or terminal ends of flowers causing distortions
and death of floral buds. P. latus is often a major problem on begonia, azalea, dahlia,
geranium and African violets particularly in autumn season (Fig. 12.29). Once
broad mites are detected on ornamental crops, ongoing vigilance is required (ER19).
12.3.7 Erinose Mite (Aceria hibisci (Nalepa))
A. hibisci damage is most clearly seen on the young leaves and the developing veg-
etative buds of hibiscus shrubs. Mites are very small (160–185 μm) and are difficult
to detect with 10× lens. Damage symptoms include crumpling of the leaves, auxil-
iary buds, petioles and calyces. A common characteristic is the production of pock-
ets of the galled tissue which are most obvious on the lower surface of the leaf as
yellow velvety overgrowth. Leaf axils and proximal margins of the leaves are par-
ticularly distorted. These mites inject salivary compounds into the expanding green
tissue which stimulates plant reaction whose presence is indicated by the velvet-
textured galls. Transmission of the mites mainly occurs through cutting of the plants
used for producing new plants. Pruning of affected plants may help in some control
of mites. Badly affected shrubs and prunings should be removed or destroyed
through burning or burying to the dump in an enclosed plastic bag. Other eriophyid
mites may produce colour patches on leaf surfaces, leaf margins that roll inwards or
downwards, swollen and distorted leaves, galls, russetting and ‘witches’ brooms’ on
different ornamentals.
12.3.8 Aloe Gall Mite (Eriophyes aloinis Keifer)
Aloe gall mite is commonly known as ‘aloe wart mite’ or ‘aloe cancer mite’ and
found on aloes almost in all habitats. Mite infestation significantly reduces aesthetic
value of aloes (ER22). They begin feeding and initiating gall formation in spring as
the leaf or flower buds open. Burn, or put in plastic bag and dispose infested plant
parts in the trash. Heavy infestations of mite can be controlled by application of
pesticides or root drenching of plants with pesticides.
12.3.9 Southern Red Mite (Oligonychus ilicis (McGregor))
Southern red mite is a serious pest of ornamentals and causes a loss of plant vigour.
High infestation of this mite can lead to plant mortality. It is widely reported from
Brazil, Japan, South Korea, the Netherlands and the USA. Adult females are reddish
or pinkish looking. O. ilicis prefers azalea, camellia and holly. It feeds on the under-
surface of leaves causing bronzing and stipplings. Later, the infested leaves turn
grey or brown and fall prematurely.
Fig. 12.29 Broad
mite-damaged dahlia plant
12.3.10 Jasmine Mite (Aceria jasmini Channabasavanna)
Jasmine mite attacks tender leaves, stems and flower buds and produce white over-
growth, and affected plants remain stunted. The life cycle of mite takes 2–3 weeks,
incubation period lasts 3–5 days and nymphal development is completed in
8–11 days. High temperature and low relative humidity favour mite development.
12.3.11 Phyllocoptes fructiphilus Keifer
Phyllocoptes fructiphilus is a tiny eriophyid mite associated with most rose species
and cultivars. This mite is the vector of a devastating viral disease of roses called
‘rose rosette disease’ (RRD). The disease causes red discolouration of new growth,
elongated shoots, deformed blooms, witches’ broom of shoots and ultimately the
death of the plant (ER17). Pruning or removal of RRD-infected multiflora rose, and
the use of soft pesticides such as light horticultural oils and soaps, may preserve the
naturally occurring natural enemies of eriophyid mite.
Further Reading 271
12.4 Conclusions
Horticultural plants include fruit trees, vegetable plants and ornamental plants
grown both under outdoor and indoor conditions. Environmental horticulture also
includes nursery production. Economically important mite pests known from differ-
ent temperate and subtropical fruit trees are included. Salient information available
regarding description and identification, biology, ecology, damage caused in differ-
ent growth stages and periods of maximum activity are discussed on different fruit
trees for different spider mites, false spider mites, eriophyid mites and broad mites.
Most salient information on economic threshold values and measures for manage-
ment of different mite pests are mentioned under each fruit. Information on different
vegetable crops is arranged in different botanical families as many times mite pests
are found commonly as pests on plant species of the same family. So the vegetables
are arranged in families Alliaceae, Convolvulaceae, Cruciferaceae, Cucurbitaceae,
Leguminosae, Solanaceae, Malvaceae and Umbelliferae and other families which
include less commonly cultivated vegetable plants. Almost all plants grown as veg-
etables for consumption as fruits, corms, tubers or leaves are included here in this
portion. Recent information available regarding management of mites on different
vegetables is considered. In case of floricultural plants, information available on
mite pests of different floricultural plants is discussed under different mite pests
found causing similar type of damage simultaneously on different flowering plants
and involves almost similar management tactics.
Further Reading
I. Fruits
Allen JC (1979) The effect of citrus rust mite damage on citrus fruit drop. J Econ Entomol
71:746–750
Beettle GAC, Gellatley JG (1983) Mite pests of citrus. NSW Agfacts, Agdec 220–222
Beettle GAC, Robert EA, Vanhoff CL, Flack LK (1991) Effects of climate, natural enemies and
biocides on three citrus mites in Coastal New South Wales. Exp Appl Acarol 11:271–295
Bergh JC, French JV (2002) A method for harvesting and shipping live citrus mites (Acari:
Eriophyidae). Fla Entomol 83:370–374
Blimberg D (2008) Review: date palm arthropod pests and their management in Israel.
Phytoparasitica 36:441–448
Childers CC, French JV, Rodrigues JC (2003) Brevipalpus californicus, B. obovatus, B, phoenicis
and B. lewisi (Acari: Tenuipalpidae): a review of their biology, feeding injury and economic
importance. Exp Appl Acarol 30:5–28
Childers CC, French JV, Rodrigues JC (2011) An overview of Brevipalpus mites (Acari:
Tenuipalpidae) and plant-viruses they transmit. Zoosymposia 6:180–192
Cranham JE, Soloman MG (1981) Mite management in commercial apple orchards. East Malling
Res Stn Annu Rep 1980:171–172
Croft BA, Hoyt SC (1983) Integrated management of insect pests of pome and stone fruits. Wiley,
New York
Dhooria MS, Bhullar MB, Malik B (2005) Mite pests of citrus and their management in India.
AINP (Agricultural Acarology), Bangalore, p 28
Easterbrook MA (1996) Damage and control of eriophyoid mites in apple and pear. In: Sabelis
MW, Bruin J (eds) Eriophyoid mites – their biology, natural enemies and control. Elsevier
Science, Amsterdam, pp 527–541
Gerson U, Cohen E (1989) Resurgences of spider mites (Acari Tetranychidae) induced by syn-
thetic pyrethroids. Exp Appl Acarol 6:24–46
Gordon SC, Taylor CE (1976) Some aspects of the biology of raspberry leaf and bud mite
(Phyllocoptes gracilis). J Hort Sci 52:517–523
Gould WP, Rage A (2002) Pests of guava. In: Pena JE, Sharp JL, Wyoski M (eds) Tropical fruit
pests and pollinators. CABI Publishing, Wallingford, pp 295–312
Hoyt SC (1969) Population studies of five mite species on apple in Washington. In: Proc. 2nd Int
Cong Acarol, Budapest, pp 117–133
McCoy CS (1977) Horticultural practices affecting phytophagous mite populations on citrus. Proc
Int Soc Hort 2:459–462
Pena JE (1990) Relationships of broad mite (Acari: Tarsonemidae) density to lime damage. J Econ
Entomol 83:2008–2015
Pena JE, Duncan R, Vasquezi T, Hennesey M (1999) Guava arthropod seasonality and control of
fruit flies in South Florida. Proc Fla State Hort Soc 112:206–209
Readshaw JL (1975) The ecology of tetranychid mites in Australian orchards. J Appl Ecol
12:473–495
Walker GP, Vaulgaropoulos AL, Philips PA (1992) Distribution of citrus bud mite (Acari:
Eriophyidae) within lemon trees. J Econ Entomol 85:2389–2398
II. Vegetables
Capinara JL (2006) Integrated pest management for potatoes in the Western United States, vol
3316, Agriculture and natural resources. University of California, Berkeley
Capinara JL (2011) Hand book of vegetable pests. Academic, San Diego
Dhooria MS (1976) Unusual severe infestation of cauliflower by Tetranychus cinnabarinus at
Ludhiana. Indian J Pl Prot 3:211
Gerson U (1992) Biology and control of broad mite, Polyphagotarsonemus latus (Banks)
(Tarsonemidae). Exp Appl Acarol 13:163–178
Gill CK, Bhullar M, Dhooria MS, Brar PS (2006) Incidence of Thrips tabaci Lindeman and Aceria
tulipae (Keifer) on garlic in Punjab. J Insect Sci 19(1):88–90
Hofan H, Learmonth S, Wood P (2004) Common insect pests and diseases on vegetables in the
home garden, vol 4624, Bulletin, State of Western Australia. Dept. of Agriculture, South Perth,
p 42. ISBN 1448–0352
Howard RJ, Garland JA, Seaman WL (eds) (1994) Diseases and pests of vegetable crops in Canada:
an illustrated compendium. CAB Direct, Ottawa, p 554. ISBN 0-9691627-31
Karmaker K (1995) Comparative symptomatology of chilli leaf-curl disease and biology of tarso-
nemid mite, Polyphagotarsonemus latus (Tarsonemidae). Ann Ento 13:65–70
Ravenberg WJ, Van Lantern JC, Wolts J (1983) Developments in application of biological control
in greenhouses vegetables in Netherlands since 1979. Bull SROP, 1983/ VI /3: 36–48
Srinivasan P (2009) Insect and mite pests on egg plant: a field guide for identification and manage-
ment. AVRDC the World Vegetable Centre, Shanhua, Taiwan Publication No. 09-729-p 64
Further Reading 273
III. Floricultural Plants
Ahmed P, Rao NHP, Rao P (2000) Biology of yellow mites, Polyphagotarsonemus latus (Banks)
in chillies. Pestology 24:20–23
Childers CC, Rodrigues JCV (2005) Potential pest mite species collected on ornamental plants
from Central America at port of entry to the United States. Fla Entomol 93:100–106
DeMoraes GJ, Tamai MA (1999) Biological control of Tetranychus spp. on ornamental plants. In:
Lindquist EE, Sabelis MW, Angarita A (eds) Proceedings of International. Symposium. on cut
flowers in the tropics, vol 482, Acta horticulture. ISHS, Leuven, pp 247–252
Dhooria MS (1999a) Two-spotted spider mite (Tetranychus urticae)- a serious pest of roses in
polyhouses, and its control. J Acarol 14:84–87
Dhooria MS (1999b) Observations on the status of phytophagous and predaceous mites on differ-
ent ornamental plants in Punjab. J Acarol 14:88–89
Dhooria MS, Bindra OS (1977) Polyphagotarsonemus latus (Banks) – a mite pest of chilli and
potato in Punjab. Acarol Newslet 4:7–9
Dreistadt SH (2001) Integrated pest management for floriculture and nurseries, vol 3405, UC ANR
publication. Oakland University of California Statewide Integrated Pest Management
Programme, Oakland
Gerson U, Weintraub PG (2007) Mites for control of pests in protected cultivation. Pest Manag Sci
63:658–676
Hamlen RA (1978) Biological control of spider mites in greenhouse ornamentals using predaceous
mites. Proc Florida State Hort Soc 91:247–249
Kaur P, Dhooria MS, Bhullar MB (2006a) Development of two-spotted spider mite, Tetranychus
urticae Koch (Acari: Tetranychidae) on rose. J Res Punjab Agric Univ 43:117–120
Kaur P, Dhooria MS, Bhullar MB (2006b) Comparative efficacy of pesticides against two-spotted
spider mite, Tetranychus urticae Koch. J Insect Sci 19:190–194
Kaur P, Dhooria MS, Bhullar MB (2006c) Suitability of flowering annuals as hosts of two-spotted
spider mite, Tetranychus urticae Koch. J Appl Zool Res 17:24–28
Naegele JA, Jefferson RN (1964) Floricultural entomology. Annu Rev Entomol 9:319–340
1. https://www.youtube.com/watch?v=m8koYcupFzc Almond Pest Management:Web Spinning

Spider Mites
2. https://www.youtube.com/watch?v=6K3EdYveDBs Apple Mites
3. http://jenny.tfrec.wsu.edu/opm/displaySpecies.php?pn=330 Pear leaf Blister Mite
4. https://www.youtube.com/watch?v=Wg_IAqZiSAM How to monitor for web spinning spider
mites and their predators in Peach and Nectarine
5. http://www.growingproduce.com/video/pest-control/controlling-spider-mites-in-strawberries/
Controlling Spider Mites In Strawberries
6. https://www.youtube.com/watch?v=jSDPlqa-5pA Spider Mite infestation on Banana
7. https://www.youtube.com/watch?v=fAC_nAAwZeM Citrus leprosis
8. https://www.youtube.com/watch?v=GTKaHw9C-fw Leprosis outbreak in the citrus industry
9. https://www.youtube.com/watch?v=bGc_pvfMEJs Integrated management of Coconut
Eriophyid Mite
10. https://www.youtube.com/watch?v=es6qASCAYng Eriophyid mites on litchi leaf
11. https://www.youtube.com/watch?v=kVCcdLxF_fY Commercial Pest Control in Pepper
Greenhouse (Spider Mites), Part 1
12. https://www.youtube.com/watch?v=97P3Juu5IiQChilli Mite English Final_WMV V9.wmv
(male, female, eggs, & damaged fruit)
13. https://www.youtube.com/watch?v=U8Rstw7zZbo Cucumber plants attacked by spider mites
14. https://www.youtube.com/watch?v=J51GKLhDybQ Tomato Russet Mite.wmv

15. https://www.youtube.com/watch?v=31ktEhMkzKU Spider Mites damage on tomato
16. https://www.youtube.com/watch?v=GyUYqnt-OLc Mass rearing spider mites on bean plants
17. https://www.youtube.com/watch?v=H-lRxwLNbzM Rose Rosette Disease
18. https://www.youtube.com/watch?v=zSW130z_AZg (Rose mite control in Colombia)
19. https://www.youtube.com/watch?v=KUcylrYhyhs Tarsonemid Mite II
20. https://www.youtube.com/watch?v=jX7nrNTtxYI Recognizing Spider Mite Damage on
Orchids
21. https://www.youtube.com/watch?v=40Ul86mE4rk Life of a Cactus Mite
22. https://www.youtube.com/watch?v=BH2l3LnX550 Aloe Mite
Mite Pests of Field Crops
13
Abstract
A crop which is a volunteered or cultivated plant (any plant) whose product is
harvested by a human at some point of its growth stage is known as a field crop.
Field crops are grown on a large scale for food, clothing and other human uses.
Many species of spider mites and eriophyid mites are found in economic propor-
tions both where these crops are grown as annual or perennial crops. Some spe-
cies of spider mites appears as most important economic pests both in tropical
and temperate regions particularly under hot and dry conditions. Cereal crops
like barley, wheat, rice, oats, maize and sorghum, and pulse crops grown both in
summer and winter season, are all damaged by one or the other mite pest at dif-
ferent crop stages. Damage on summer cultivated field crops is particularly more
under hot and dry conditions. Besides causing damage to plants, eriophyid mites
are also vectors of wheat streak mosaic virus and pigeon pea sterility mosaic
virus disease. Barley yellow streak mosaic virus (BaYSMV) is also vectored by
brown wheat mite (BWM). Panicle rice mite (PRM) infestation is concentrated
on reproductive structures of plants during the milky stage of development result-
ing in deformation of the inflorescence and discolouration of rice hulls. Many
mite species are considered as invasive pests and may prove troublesome in ear-
lier uninfested areas.
Keywords
Cereals • Spider mites • Sterility mosaic disease
A crop which is a volunteered or cultivated plant (any plant) whose product is har-
vested by a human at some point of its growth stage is known as a field crop. Crops
refer to plants of the same kind that are grown on a large scale for food, clothing and
other human uses. They are nonanimal species or varieties grown to be for any other
economic purpose.

DOI 10.1007/978-981-10-1594-6_13
276 13 Mite Pests of Field Crops
13.1 Cereals
A cereal is a grass in the monocot family Poaceae (also known as Gramineae) culti-
vated for their edible grains. Cereal grains are grown in greater quantities and pro-
vide more food energy worldwide than any other type of crop. They are also known
as staple crops.
13.1.1 Barley (Hordeum vulgare; Poaceae)
Barley – a cereal grain – is the most important cereal crop in the world after rice,
wheat and maize. Barley is a very adaptable and widely grown as a summer crop in
temperate and as a winter crop in tropical areas. European Union, Russia, Ukraine,
Canada, Australia and the USA are the major producers of barley.
Brown Wheat Mite (Petrobia latens (Muller); BWM) Reported as a pest of both
barley and wheat during dry weather and drought cycles. Prolonged drought cycles
extending for multiple years or extended drought periods within a cropping season
favour the mite which also transmits ‘barley yellow streak mosaic virus’ (BaYSMV)
disease. BWM lays two types of eggs – red and white, which are attached to the soil
particles near the base of barley or wheat plants. Red eggs hatch after a brief incuba-
tion period, producing multiple overlapping generations. The white eggs are a rest-
ing stage and do not hatch during the summer. Cooler fall weather triggers the white
eggs to hatch after a 10-day incubation period. BWM moves from the soil to the
host plant to feed. Feeding produces a fine stipplings that cause leaves to take on a
water-stressed appearance turning first dusty grey and then yellow. Infested barley
plants appear scorched or bronzed and withered. Alfalfa, clover and some vegetable
crops like carrot and coriander are also reported as host plants of BWM.
Barley yellow streak mosaic virus (BaYSMV) is known to be vectored by

BWM. In general, infestation of BWM is not known to damage much barley plants,
but treatments against BWM seem justified when BaYSMV is found. Rainfall sig-
nificantly reduces BWM. Early planted barley is reported to be less damaged by
BWM than later emerging barley during drought years. Barley following barley has
been reported more favourable for disease development. In many areas, residue
destruction by burning has resulted in partial reduction of BaYSMV disease.
Blue Oat Mite (Penthaleus spp.; BOM) Blue oat mites are important pests of
seedlings of winter cereals (including oats and barley) but are restricted to the cooler
periods. BOM pierce and suck sap of leaves resulting in silvering of the leaf tips;
these mites take shelter on the soil surface when conditions are warm and sunny.
Many times BOM infestations result in green or greyish irregular patches in the
crop.
13.1 Cereals 277
13.1.2 Maize (Zea mays; Poaceae)
In the USA, maize is often referred to as corn, but the word corn is a misnomer.
Maize originated from Central America, but is distributed worldwide. Maize crop
can be grown in a wide range of climates in both temperate and tropical regions
throughout the world.
Spider Mites Oligonychus pratensis (Banks), Tetranychus urticae (Koch),

Tetranychus turkestani (Ugarov and Nikolskii) and Tetranychus pacificus
(McGregor)(ER16 and ER18 and ER19). Two types of spider mites, namely, Banks
grass mite (BGM) and two-spotted spider mite (TSSM), are commonly found on
corn including sweet corn. TSSM has vast host range and reported to feed on many
ornamental plants, vegetables, beans and many grasses. Both the species are similar
in appearance but differ in several biological characteristics. BGM usually appears
early in the season and feeds mostly on the lower leaves of the plants. But TSSM
appear in mid to late season, increase rapidly and feed over the entire plant (Sandhu
et al. 1975).
BGM primarily overwinters in the crowns of winter wheat and native grasses,
but TSSM primarily overwinters on some broad-leaved plants bordering the corn
field and crawl or is carried by wind to corn. The first evidence of mite feeding is
noticed on the top of the leaf where a yellow or whitish spotting of the leaf tissues
is found in areas where the mites are found feeding on the lower leaf surface. As
mite infestations develop, leaves may get severely damaged and the photosynthesis
is badly affected. Leaves may even be killed if mite infestation is very severe.
Effects on yield are most severe when mites start damaging leaves at or above the
ear level. Infestation may reduce corn grain yield due to poor seed fill. Both the mite
species produce webbings, but TSSM tends to produce more extensive webbings
than BGM. Banks grass mite appears often earlier in the season and remains longer
on the lower leaves before moving up the plants. TSSM usually appear later in the
season. Spider mite control decisions are based on many factors including the mite
species present, level of infestation, growth stage of the crop, cost of application and
market price of the crop.
Reduce moisture stress through timely irrigation; treat only heavily infested
areas of the field to allow recolonization by natural enemies. When the corn has
reached the full dent stage, it is unlikely to benefit from treatment against spider
mites. Commercial yield losses as high as 40 % for silage (dry matter) and 21 % of
grain losses from Colorado (USA) have been reported as a result of serious infesta-
tion of spider mites during hot and dry years. Late planting of corn or longer season
varieties suffer less mite damage than the ones which are planted earlier. The simple
guideline is to treat if damage is visible in the lower third of the plant, and mite colo-
nies are present in the middle third. Once the crop has reached the hard dough stage,
no economic benefit will result from the treatment. Sometimes feeding of eriophyid
mite, Eriophyes tulipae Keifer, on corn kernels causes a disease known as ‘kernel
red streak’, wherein discoloured corn ears show red striped pericarps.
13.1.3 Rice (Oryza sativa; Poaceae)
Rice plant originated from Asia and prefers a tropical or warm climate with a lot of
rainfall. It is mainly grown in China, India, Indonesia, Vietnam and Thailand.
Panicle Rice Mite (Steneotarsonemus spinki Smiley; PRM) Panicle rice mite
(PRM) is known in rice-producing areas of the world by a variety of common names
including the rice panicle mite, rice tarsonemid mite, rice mite, rice white mite and
spinki mite. PRM has been recognized as rice mite typically throughout the rice-
growing regions of the Asia. As the rice grain develops, more mites can be found
throughout the panicles. Peak activity of the mite feeding and multiplication are
found during the milky stage of development. The damage inflicted by rice mite is
typically concentrated on the reproductive structures of the plants and may include
deformation of the inflorescence and panicles as well as dehydration and necrosis of
panicle tissues, including discolouration of the rice hulls (ER15). Yield losses attrib-
uted to the rice mite range from 5 to 90 %. From China, 5–20 % yield reduction in
both early and late season rice crops has been attributed. From India, Japonica vari-
eties are reported more susceptible to PRM than Indica varieties suffering about
20 % yield loss. Some plant species belonging to families Cyperaceae and Poaceae
have also been reported as hosts of PRM, and some plant diseases have also been
associated with PRM infestations around the world. PRM has fast and efficient
reproduction. Females produce 50–70 eggs in their lifetime and have a short genera-
tion time ranging from 9 to 17 days at 25–30 °C. There can be 48–55 generations
per year under most ideal climatic conditions. PRM survive in environments with
high humidity. Higher densities of PRM are associated with high planting density.
Before planting a new crop, clean the new crop areas to remove any remaining mites
before planting a field; and, if possible, leave the fields fallow or at least 2 weeks
between crop cycles. Avoid staggered plantings in mite-infested areas.
Spider Mites (Oligonychus oryzae (Hirst)) O. oryzae is an economic pest infest-

ing rice in India and some other Asian countries. From India severe outbreaks of this
mite have been reported from Karnataka region resulting in considerable yield loss
on pre-kharif rice. Relatively low temperature (27 °C) and high humidity (85 % or
more) are conducive for the rapid multiplication of the mite, while relatively high
temperature (31 °C) had an adverse effect on mites. Some weeds like Panicum col
oratum, P. crusgalli and Echinochloa spp. have been reported as alternate hosts for
O. oryzae during September. Affected plants showed whitish patches on leaf sur-
faces, and in severe cases the leaves turned greyish white and dried.
13.1.4 Sorghum (Sorghum bicolor; Gramineae)
Sorghum is one of the top cereal crops in the world along with wheat, oats, corn,
rice and barley. It originated from Egypt but is now well spread to southern Asia and
13.1 Cereals 279
the Americas as well. It is an annual and is extremely tolerant to drought and is an

excellent choice for arid and dry areas.
Banks Grass Mite (Oligonychus pratensis; BGM) Sometimes large numbers of

BGM are found on sorghum especially in arid areas. Full-fed mites are deep green.
Mites construct webbings on undersurface of leaves; about 50 eggs are laid by a
female mite. Eggs are pearly white, spherical and hatch in 3–4 days. The life cycle
is completed in about 10–12 days under favourable conditions. Mite infestation
begins along the midrib region of the lower leaves; infested areas of leaves become
pale yellow initially and later become reddish on the top surface. Dense mite web-
bings are commonly found on undersurface of infested leaves. Increases in BGM
population generally occur after the emergence of sorghum grain heads. Large num-
bers of BGM occurring during early kernel development can reduce the ability of
sorghum plants’ make and fill grain, but not affected after kernels reach hard dough
grain. If BGM are very abundant, sorghum plants may lodge resulting in reduced
yield. Mite infestations commonly begin along field borders and may spread quickly
throughout the sorghum field. Hot and dry weather favours rapid build-up of mites
(ER17). BGM may also develop as induced (secondary) pests after insecticide
application for control of other key insect pests. Irrigations should be timed prop-
erly to avoid moisture stress. Insecticide applications should be considered when
30 % of leaves of most sorghum plants show mite damage symptoms. Destruction
of weeds which are favourite hosts of BGM around the sorghum field should be
done in fall or early spring. Another spider mite, Oligonychus indicus, also some-
times appears as pest of sorghum during drought conditions in India.
13.1.5 Wheat (Triticum aestivum; Gramineae)
Wheat is an important cereal crop, originated in the near East and Mesopotamia and
grows best in temperate climate. Canada, China, India, Russia and the USA are
leading producers of wheat.
Wheat-Curl Mite (Aceria tosichella Keifer; WCM) Wheat-curl mite is known as

a vector of important virus disease affecting wheat and corn in America. WCM is an
important pest of cereal crops in North America, Europe, Asia, the Middle East,
Africa and Oceania. Direct damage during high infestation includes discolouration,
curling or rolling of leaves, abnormal development of leaves and plant stunting.
Yield losses in wheat crops due to high populations/infestations of WCM can reach
around 30 %. However, the main damage caused by A. tosichella is due to the trans-
mission of wheat streak mosaic virus (WSMV) and high plain virus (HPV) diseases
(ER1 and ER2 and ER5). WCM occurs mainly on wheat, but its populations can
also develop on maize, sorghum, barley, oats, rye and on many grasses and weeds
of minor importance. At 25 °C mite population build up rapidly, but at 0 °C develop-
ment stops. Hot dry weather can reduce mite populations and virus problems sig-
nificantly, but survival of alternate hosts can prevent their complete elimination.
New infestations usually occur first along field margins and then spread. Mites on
corn plants feed and reproduce in the leaf sheaths. At ear formation mites can move
to feed on husks, silks and kernels. Volunteer wheat and grasses should be destroyed
to limit the vector survival; avoid late seeding. Cultural practices and resistant vari-
eties are the major tools in mite and disease management.
Banks Grass Mite (Oligonychus pratensis; BGM) Banks grass mite (BGM)
attacks a wide variety of grasses including turf, sorghum and wheat. At 24 °C, BGM
completes a generation in 10–12 days. In early stages mites are white salmon
coloured and become progressively darker green during their life cycle. Adults are
deep green, but overwintering forms are bright orange and may be found at the base
of host plants. These mites produce heavy webbings on undersurface of leaves for
their protection (ER2). Some plants are often killed, and kernel size is reduced as a
result of high mite infestations. Insecticide applications to field margins bordering
corn and alfalfa are often sufficient to prevent economic damage. BGM infestations
will cause the foliage to turn yellow initially; later as the infestation increases, the
leaves turn yellowish brown. Heavy webbing is typical of infestation by this mite.
Two-Spotted Spider Mite (T. urticae; TSSM) Two-spotted spider mite (TSSM)
incidence sometimes found on undersurface of leaves causes yellowing or silvering
of leaves. Patchy infestations (‘hot spots’) are characteristic of this pest in fields.
Sometimes mites are found suspended on silken threads and are thus dispersed by
air currents. Keeping dust to a minimum will also help prevent TSSM populations
from exploding. Number and speed of vehicles passing along roads adjacent to
wheat fields should be kept to a minimum. Heavily travelled roads can be mulched
with hay or other materials or sprayed with water to keep down dust. Proper irriga-
tion management of the wheat fields should be done to avoid mite build-up (ER2).
Brown Wheat Mite (Petrobia latens; BWM) It is mainly found on wheat, grass
hosts and other monocots such as onions. BWM is strictly a pest under dry weather
conditions, but may go to broad-leaved crops including alfalfa. BWM deposits two
types of eggs, diapausing (resting) and nondiapausing on soil particles near the
plant. Nondiapausing eggs hatch in about 10 days, but diapausing eggs require
moisture to initiate hatching; otherwise, they remain in dormant stage for an indefi-
nite period. BWM is mainly found on the upper surface of leaves. Plants heavily
infested with BWM have a stippled appearance giving yellowish look. Damage is
always associated with drought stress. Rainfall or irrigation will cause a dramatic
reduction in mite numbers. For proper management of BWM on wheat, weakly
monitoring should concentrate on field edges and corners where infestations are
likely to begin. Management of volunteer wheat is an important measure for pre-
venting BWM. Barley yellow streak mosaic virus (BaY SMV) is transmitted on
recrop of barley by BWM under dry conditions. Proper irrigation of wheat fields
will enable plants to tolerate mite damage. Sprinkler irrigation at the outset of mite
13.1 Cereals 281
population increases is many times recommended for control of both BWM and
BGM on wheat.
Winter Grain Mite (Penthaleus major (Duges); WGM) Winter grain mite has a
similar host range as of brown wheat mite (BWM) but may also infest legumes such
as alfalfa (ER2). WGM is larger in size and darker in colour, often having a dark
blue or green tinge as compared to the smaller BWM. Winter grain mite also lays
both diapausing and nondiapausing eggs. Mites lay eggs on soil particles near the
plants and on the lower leaves and stems of hosts like wheat, barley and grasses.
Injury by WGM can be prevented by rotating crops every 2 years. Mites are active
during cooler periods of the year (mid fall to late spring) with peak populations in
winter months. Infestations are comparatively more in fields treated earlier with
sludge or manure. Damage on young plants is more severe than on older established
plants. Damage may also be greater in plants stressed by nutrient deficiencies or
drought conditions. WGM is also found as a minor pest of turf grasses.
13.1.6 Oats (Avena sativa; Poaceae)
Oat is an annual grain crop grown for seed production used for livestock feed,
breakfast cereals and oatmeal. Oat is a spring-seeded crop and normally grown as
cover crop for alfalfa.
Banks Grass Mite (Oligonychus pratensis; BGM) BGM is an important pest of

oats. Mites look yellow to cream. Heavy mite population can kill small plants but
reduce kernel size in larger plants. Mites produce heavy webbings in which eggs,
larvae and adults are commonly found. Damaged leaves first become yellow, then
brown and finally necrotic. The overwintering form of mites is bright orange. With
the onset of winter, the mites move to wheat plant crowns where they feed until
spring when they disperse by ballooning on corn plants. Most banks grass mite
problems occur under hot and dry conditions, low humidity, absence of TSSM
(Tetranychus urticae) and lack of natural enemies.
Brown Wheat Mite (Petrobia latens; BWM) Sometimes, it appears as an impor-

tant pest of oats. They are oval shaped and dark red or brown. They spend the sum-
mer in the soil as white eggs resisting hot and dry conditions. In the fall as
temperature and moisture increases, eggs develop and hatch. Developing females
lay round red eggs which actively feed and reproduce. Feeding by BWM causes
mottled leaves owing to sap feeding and later yellow or bronze. BWM feeds during
the day, and peak mite population is also found during mid-afternoon on warm and
calm days. Mite population is suddenly reduced following driving rains (0.85 cm or
more).
Wheat-Curl Mite (Aceria tosichella; WCM) It is found mainly on wheat and has
also been reported on oats but seems to have minor importance.
13.2 Pulses
The term pulse is used by Food and Agriculture Organization (FAO) for crops har-
vested solely for the dry seed. Pulses belong to Fabaceae or Leguminosae family.
They exclude green peas. Groundnut and Soybean are grains but are grown mainly
for oil extraction. FAO recognizes dry beans like kidney bean (Phaseolus spp.), lima
bean (P. lunatus), mung bean (Vigna radiata), black gram (V. mungo), rice bean (V.
umbellata), dry broad beans, horse beans (Vicia faba), broad bean (V. faba) and
chick pea (Cicer arietinum); dry cowpea (V. unguiculata), pigeon pea (Cajanus
cajan), lentil (Lens culinaris) and lupins (Lupinus spp.) as pulses.
13.2.1 Pigeon pea (Cajanus cajan; Fabaceae)
Pigeon pea is a perennial legume, originated from India, and is mostly grown in
tropical regions. Highest production of pigeon pea is in Indian subcontinent, eastern
Africa and Central America.
Eriophyid Mite (Aceria cajani Channabasavanna) It is mainly reported as vec-

tor of pigeon pea sterility mosaic (PPSM) disease. However, direct mite feeding or
feeding through injecting toxins on the plants by mite feeding has not been estab-
lished. A single infected mite has been reported to effect the transmission of PPSM
disease in as short a period as 20 min. Healthy eriophyid mites, when exposed to
diseased plant leaves and then transferred to healthy plants, resulted in the develop-
ment of sterility mosaic symptoms. Greater mite intensity is reported on young
leaves compared to mature leaves. Top canopy of plants had more mite population
than other stratum of plant. Sterility mosaic disease is more prevalent in South
India. Common damage symptoms of PPSM disease are reduction of leaf size,
bushy growth of plants and yellowish development (mosaic type), and undersurface
of infested leaves becomes velvety to touch. Flower formation is inhibited, and
even, if formed, flowers wither and fall off resulting consequently in no seed forma-
tion. Incidence of mite and disease starts appearing 45 days after sowing. Uprooting
and destruction of mosaic-infected pigeon pea plants as well as the infected stubbles
of the previous crop will discourage the survival and carryover of the vector mite.
Weed (Atylosia spp.) many times harbours the vector, so discourage mite
development by uprooting this weed (Lateef and Reed 1990; Reed and Lateef 1990;
Shanover et al 1999).
Spider Mite (Schizotetranychus cajani Gupta) Sometimes S. cajani appears as a

minor pest of pigeon pea, causing yellowing of leaves (Figs. 13.1 and 13.2). Mainly
it feeds and breeds on lower surface of leaves. Females appear to be yellowish green
13.2 Pulses 283
Fig. 13.1 T. urticae

damaged pigeon pea
Fig. 13.2 S. cajani damaged pigeon pea plant

and mostly infest the crop from vegetative stage to pre-flowering stage. Mite feed-
ing causes reduction in photosynthetic activity. Severe mite infestations result in
speckling and drying of the leaves.
Spider Mite (Eotetranychus broodryki Meyer) Mostly it infests plants suffering

from mosaic at the late vegetative stage to the reproductive stage of the crop. Mite
looks greenish yellow and infests mostly midrib region and other main veins on
undersurface of leaves. Mite feeding results in characteristic yellow streaks all
along the midrib and outer veins on the upper surface of leaves. Sterility mosaic-
affected plants are more prone to mite attack. In some pockets of India (Gujarat,
Bihar, Uttar Pradesh and Haryana), sterility mosaic is a very important disease.
13.2.2 Cowpea (Vicia spp.)
Spider Mites (Tetranychus ludeni Zacher and T. urticae) They are commonly
found as pests on cowpea particularly during hot and dry periods (May–June), but
mite population declines suddenly in rainy period. Mites spin dense webbings on
the plants, and the infested plants attract a lot of dust particles adversely affecting
the photosynthesis (Fig. 13.3). Mite infestation is more in nitrogen-rich soils.
During severe infestations, mites congregate at the tip of the plant parts like leaves
and tender shoots and readily disperse through wind. As a result of mite infestation,
initially yellowish-white speckles appear which later turn yellow, curl, twist and fall
off. Heavy mite infestation leads to mortality of plants (ER8).
Fig. 13.3 Spider mite damaged bean plant

13.2 Pulses 285
Fig. 13.4 Broad mite damaged cowpea
Broad Mite (Polyphagotarsonemus latus (Banks)) Sometimes it infests and dam-

ages plant growth causing downward curling, more greening and development of
rusty-red colour on undersurface of leaves (Fig. 13.4). Mite infestation is sometimes
serious on nursery plants particularly in greenhouses, but damage under field condi-
tions is quite insignificant during warm and humid conditions.
13.2.3 Black gram (Vigna mungo)
Black gram is one of the important grain legumes in the rain-fed farming system and
can be grown under low moisture fertility conditions (Fig. 13.5).
Two-Spotted Spider Mite (Tetranychus urticae; TSSM) Sometimes it is found

seriously on black gram particularly during hot and dry months at maturity of crop.
In main season crop, mites do not generally develop into serious proportions.
Broad Mite (Polyphagotarsonemus latus) Sometimes it affected young new

growth and inflorescence producing malformed flowers resulting in reduced yield
(Gupta and Dhooria 1973).
13.2.4 Green gram (Vigna radiata)
It is native to Indian subcontinent and mainly cultivated in India, China, Thailand

and Bangladesh. Spider mites, T. urticae, sometimes cause minor damage. But
broad mite, P. latus, may cause significant damage during warm and humid period
(July–August)(Dhooria 2005b).
Fig. 13.5 Spider mite damaged mung bean field
13.2.5 Winter Pulses
Red-legged earth mite (Halotydeus destructor (Tucker), RLEM) and blue oat mite
(Penthaleus spp., BOM) are sometimes found in pest form on winter pulses, namely,
chick peas (Cicer arietinum), faba beans (Vicia faba), field peas (Pisum sativum),
lentil (Lens culinaris) and lupins (Lupinus alba and L. angustifolius), in Australia
particularly during seedling stages. BOM (Penthaleus falcatus Qin and Halliday,
Penthaleus major Duges and Penthaleus tectus Halliday) also attains pest status on
the above crops. RLEM feeding causes silvering on the upper surface of cotyledons
and leaves (Ridsdill-Smith 1997). Lupins (at epigeal emergence) are particularly
susceptible at emergence; and sometimes, if uncontrolled, mite damage may result
in mortality of seedlings. However, other pulses (hypogeal emergence) may recover
from the mite damage at emergence by reshooting from the base. Faba beans, field
peas and yellow lupins are particularly susceptible to foliar damage. On chick peas
and lentils, mite damage is also reported.
13.3 Fibre Crops
Fibre crops are field crops grown for their fibres and are traditionally used to make
paper, cloth or rope. Fibre crops are generally harvestable after a single growing
season, as distinct from trees, which are grown for many years before being har-
vested for wood pulp fibre, jute, flax and Indian hemp (stem-skinned fibres); and
cotton, coir, sisal and yucca (an agave) are other fibres.
13.3 Fibre Crops 287
13.3.1 Cotton (Gossypium spp.; Malvaceae)
Cotton plant is a shrub native to tropical and subtropical regions around the word.
Upland cotton (G. hirsutum) or Mexican cotton is the most important, and 90 % of
all cotton production is under upland cotton.
Two-Spotted Spider Mite (Tetranychus urticae; TSSM) TSSM is a complex and

includes T. cinnabarinus. As a result of mite feeding, light-coloured punctures or
spots are found, which cover large areas of the leaf. Feeding results in water loss.
When spider mite feeding is severe, defoliation and a total loss of squares and fruits
may occur. Other signs of the mite damage include russetting. Often mite-infested
areas can be spotted by having reddish or yellowish russetting or bronzing on under-
side of leaves (Gutierrez 1994). Resistance to TSSM feeding found in Gossypium
spp. (including G. barbadense, G. arboreus, G. herbaceum and G. austrae). Among
the cultivated cottons, G. hirsutum has several morphological and biochemical traits
that confer partial resistance, while G. barbadense varieties show high antibiosis. In
G. hirsutum, both constitutive and induced components of plant resistance have
been identified. Genotypes with lower sugar and starch content are generally poorer
hosts for spider mites. High populations of TSSM can affect the yield and quality of
cotton (Wilson 1993). The degree to which photosynthesis is reduced depends on
how many mites are on the leaf and for how long duration. In general mite popula-
tions which develop earlier in the cotton season and which develop to high densities
more quickly will cause the great losses (ER10 and ER12).
Spider Mite (Tetranychus turkestani) Females are straw coloured or greenish

with dark spots on each side of the body. Besides causing sucking damage to the
host, this mite also injects a toxin that causes severe damage to the plants and is
sometimes considered as a most destructive species. In the infested fields, leaves
may turn bright red and orange. Hot and dry weather is favoured for mite infesta-
tions. Very high relative humidity kills the spider mites during moulting, and wash-
ing them off leaves and favours the pathogenic fungus, Neozygites floridana, which
can cause rapid decline in mite populations.
Broad-spectrum pesticides reduce populations of mite predators which other-

wise help in keeping mites under check. Dust produced by vehicular movement on
unpaved roads adjacent to cotton fields and within fields during dry conditions
favours mite development. Plants suffering from nutritional stress particularly
potassium deficiency may be especially prone to spider mite infestations. Spider
mite outbreaks in cotton are related to mite population levels on corn and soybeans
fields where they overwinter and develop in May and June. Mites should be scouted
on a weekly basis starting in early June. Spider mite infestations can be determined
more rapidly by direct examination of suspected infested plants while walking in
the fields. One to ten mites per plant can be rated as light infestation, but 11–50
mites/leaf present on numerous plants when leaves are speckled, mottled yellow or
red are rated as medium infestation. But when more than 50 spider mites/leaf are
found on most of the leaves and on most of the plants, then the mite infestation is
rated as heavy. At this stage, many leaves exhibit reddish-brown colour.
Besides T. urticae and T. turkestani, other spider mites, namely, Oligonychus
gossypii Zacher (dark red in colour), T. desertorum (carmine in colour), T. gloveri
Banks, T. ludeni Zacher and T. pacificus, have been reported sometimes damaging
cotton plants. Petrobia latens has also been predominantly reported as a pest of
seedling cotton. Spider mites usually attack cotton during the latter part of the grow-
ing season. Hot and dry weather favours mite population increase. Since mites are
found in the same area year after year, monitoring for mites, fence rows and other
areas where mites were found last year needs critical examination. When chronic
problems occur, spot treatment may be required at 5–10 days interval for satisfac-
tory control. Treatment against mites should be given when 30–50 % plants show
infestation.
False Spider Mites Brevipalpus californicus (Banks) and B. phoenicis (Geijskes).

These are polyphagous mites, are very flat and have brick-red colour. These are
found on both surfaces of leaves along midrib portion and are less spread than spi-
der mites. All these mites are found in small populations and seem to have no eco-
nomic relevance on cotton.
Tarsonemid Mite (Polyphagotarsonemus latus (Banks)) It is distributed through-

out the tropics and is a greenhouse pest in temperate regions. A generation is com-
pleted in 4–5 days at 26–30 °C and at high relative humidity. A female lays about
3–8 eggs. It damages mainly young leaves, and lesions are caused by deformities on
leaf margins (ER11 and ER14). In cotton, mite incidence results in shortened inter-
nodes and results in reduced boll weight. P. latus is mainly a pest of cotton in humid
tropical regions such as Central Africa, Uganda and some areas of Brazil. In favour-
able climates, sometimes mite outbreaks cause about 11 % reduction in cotton yield.
Eriophyid Mite (Acalitus gossypii Banks) It is also known as cotton blister mite,
and its infestation produces hairy deformations on leaves and flowers. Leaves may
be crumpled and shoots distorted. Gossypium barbadense is very sensitive to this
mite, while G. hirsutum suffers least damage. Another eriophyid mite, Eriophyes
puttarudriahi (Channabasavanna), is reported feeding on cotton (G. herbaceum)
from India causing felt-like outgrowth on tender shoots. The infested plants may
remain stunted and boll formation is suppressed.
13.3.2 Jute (Corchorus capsularis; Malvaceae)
Jute is an important fibre crop. It originated from India, Burma and Africa and needs
a plain alluvial soil and standing water. Mainly it needs a monsoon climate (20–
40 °C and 70–80 % RH). India and Bangladesh are the major producers of jute.
13.4 Oilseed Crops 289
Spider Mites (Oligonychus coffeae Nietner) O. coffeae is a serious pest of coffee,

tea and Jute in India, Bangladesh, Sri Lanka, Australia and Indonesia. Females are
bright crimson dorsally and purplish or bronzy ventrally. Mite infestation is mainly
confined on upper surface of leaves; infested leaves show yellowish spots which
later turn brown. In case of severe infestation leaf fall may occur, heavily infested
plants become useless for fibre production.
Broad Mite or Yellow Jute Mite (Polyphagotarsonemus latus) Polyphago

tarsonemus latus is a major pest of capsicums, but is also found as pest on jute. Mite
activity is mainly restricted to top few leaves. In Bangladesh, P. latus is considered
as one of the major limiting factors of jute production. Both C. capsularis and C.
olitorius are susceptible to mite attack. The entire life cycle of mite is of very short
duration (3 days), which helps in its fast multiplication. The mites suck the sap of the
young leaves in the crown even before they are unfolded. Affected leaves present an
oily look but later turn deep green with coppery brown shades. The midrib curves
downward, and the leaf lamina rolls inwards from both sides. The secondary veins
wrinkle and give the leaf a rough and crumpled look; the affected leaves give a dis-
tinct sick look to the crop although normal leaves may grow below. The affected
leaves often drop off prematurely. The loss in yield is estimated at about 20 %. A
crop exposed to prolonged periods of high relative humidity with occasional drizzles
favours mite infestation, while heavy downpour of rain helps the crop regenerate
growth free from mite infestation (Rahman and Khan 2012).
13.3.3 Sunn Hemp (Crotalaria juncea; Fabaceae)
Eriophyid Mite (Aceria crotalaria Channabasavanna) It is reported infesting

sunn hemp crop during May–July in West Bengal (India). Mite infestation causes
twisting of apical leaves which become shortened, discoloured and ultimately yel-
lowish. Attacked part of the stem looks deep brown due to thick growth of brown
erinea. Internodes of mite-infested plants are shortened, and plants remain stunted.
13.4 Oilseed Crops
Rapeseed, canola, soybeans, sunflower, safflower, flax seed and mustard are impor-
tant oilseed crops.
13.4.1 Soybeans (Glycine max; Fabaceae)
Soybean is an annual plant and requires climate with hot summers (with mean tem-
peratures of 20–30 °C). It originated from East Asia; the USA, Argentina, Brazil,
Canada, China and India are important producers of soybean.
Two-Spotted Spider Mite (Tetranychus urticae) Tetranychus urticae has been

reported as a serious pest of soybeans. As a result of mite feeding, leaf spotting, leaf
droppage, accelerated senescence, pod shattering and yield reduction occur. Mite
damage first appears as small spots (‘stipples’); as mite feeding activity increases,
leaves become yellow, brown and eventually drop. From 40 to 60 %, reduction in
yield has been reported in case of high mite infestation during the late vegetative
and early reproductive growth stages. Mite infestation may cause yield reductions
as long as green pods are present. Dry and hot weather favours mite reproduction
and survival especially if accompanied by drought stress in the crop. When the
weather during June–August is especially hot and dry, mite infestations can reach in
damaging proportions. If hot and dry conditions continue during high mite infesta-
tion, even acaricide treatment may not produce adequate results; however, treatment
may be beneficial if significant pod or seed filling remains and leaves have yet not
turned yellow. In less rainfall regions and in sandy type soils, more spider mite out-
breaks are found, as in such soils chances of drought stress are more likely even
when irrigated. Disturbance of natural enemies through use of broad-spectrum pes-
ticides for insect pest control may also help in mite outbreaks. So mite infestations
occurring in the season should be scouted during the rest of the season.
13.4.2 Canola (Brassica napus; Brassicaceae)
Canola is a name applied to edible oilseed rape and belongs to the same family as
mustard, broccoli, Brussels sprouts and cauliflower. It is mainly grown in Australia,
Canada and the USA. Canola is widely adapted particularly to the cool extremes of
the temperate zones.
Earth Mites (Halotydeus destructor, Penthaleus spp.) The red-legged earth mites
(Halotydeus destructor; RLEM) and blue oat mite (Penthaleus spp.; BOM) are the
most regular and damaging pests of canola in southern Australia. The different spe-
cies often coexist. Adult mites rupture the surface of cotyledons and leaves, remov-
ing cell contents. Damaged plants may wilt and die in case of serious mite infestation.
More serious damage is observed particularly if growing conditions are poor.
Leguminous pastures serve mainly as major source of mite outbreaks. Sometimes
during spring, densities up to 30,000 mites/m2 are found (Ridsdill-Smith 1997).
Mites emerge from over-summering eggs (physiologically arrested or diapause

which survive through summer) in response to autumn rains and falling tempera-
tures. Juvenile mites feed on soil microflora (algal, mosses, bryophytes). RLEM may
have 3–4 generations during autumn to spring; BOM possibly have fewer. The use
of non-host crops like lentil, chickpea, wheat, barley and linseed prior to canola in
the rotation dramatically reduces the build-up and carryover of earth mites to densi-
ties below economically damaging levels in autumn. Intensive grazing of pastures in
the previous spring also reduces mite carryover. The cultivation of paddocks in early
autumn kills diapausing eggs and newly emerged nymphs. Spraying for RLEM at the
correct time before hatching of summer eggs proves useful (Anonymous 2005).
Clover Mite (Bryobia spp.) Feeding by clover mite is almost similar to that caused
by RLEM feeding. Whitish grey spots on the top of leaf surface are associated with
clover mite feeding; damage is quite high in years when there are autumn rains.
Early rains allow weed growth that lead to increased mite populations. All weed
hosts should be destroyed before seeding and/or applying on weed a herbicide with
the knockdown effect.
13.4.3 Groundnut (Arachis hypogaea; Leguminosae)
Groundnut also known as peanut is herbaceous in nature and seems to have origi-
nated from Paraguay in South America and is now grown throughout the tropical
and warm temperate regions of the world.
Peanut Mites (Paraplonobia spp.) Peanut mites are dark green to black oval with
light amber legs. They are relatively large mites (1.5 mm long) and fall off to the
ground when disturbed. Leaf feeding is characterized by fine silver stipplings. High
infestation of this mite causes shedding off lower leaves and subsequent death of the
infested plants. From Queensland (Australia), the mite is sometimes reported very
serious on peanuts; sprays can be initiated if more than 30 % of plants are showing
damage symptoms. Regular irrigations may reduce the risk of peanut mite
damage.
Two-Spotted Spider Mite (TSSM) It feeds on underside of leaves, causing a yel-

low speckling of leaves, and the foliage gradually turns from yellow to brown.
Large populations of mites sometimes are responsible for serious plant damage and
defoliation. Usually a fine webbing is associated with the feeding site of mites.
Initial infestations are small and spotty and are limited to areas 2–3 feet in diameter.
Major outbreaks of TSSM on groundnut have been reported during hot and dry
weather (on droughty/stressed plants). If weather conditions are favourable, certain
pesticides may also trigger mite outbreaks. Treatment decisions depend on judg-
ment and expectations of changing weather conditions and consider control mea-
sures under hot and dry climate conditions. Best results are obtained if treatments
are applied early in the infestation cycle before many eggs have been laid. Rainfall
and humid conditions are likely to lower mite numbers. Check regularly to monitor
changes. Irrigation is the best defence against TSSM injury/damage.
Spider mites, Tetranychus macfarlanei Baker and Pritchard and T. hypogaea

Gupta, have also been reported on groundnut. Infestation makes leaves to look yel-
lowish to brownish. Affected leaves often dry up and pod formation is adversely
affected. T. hypogaea infests groundnut leaves sometimes and adversely affect the
infested plants.
13.4.4 Castor Bean (Ricinus communis; Euphorbiaceae)
Castor bean is a fast-growing tender perennial with huge leaves that can be used as
a seasonal annual in cooler climates. Castor bean is native to tropical East Africa
around Ethiopia, but has naturalized in tropical and subtropical areas around the
world to become a weed in many places.
Spider Mites Tetranychus urticae, T. ludeni and Eutetranychus orientalis (Klein)

have been reported infesting castor plants in India. T. urticae colonizes on undersur-
face of leaves mainly during February–June causing yellowing and browning of
leaves (Ahuja 1994; Gupta et al. 1976). Ultimately severely mite-infested leaves dry
and fall off. E. orientalis mainly infests upper surface of leaves which attract a lot
of dust particles affecting photosynthesis (Fig. 13.6).
13.4.5 Sesame (Sesamum indicum; Pedaliaceae)
Sesame (commonly known as ‘Til’ or ‘Gingely’ or ‘SimSim’) probably originated

from Africa. India is the leading producer of the sesame.
Broad mite, Polyphagotarsonemus latus, infests mainly terminal one-third por-
tion of plants (Figs. 13.7 and 13.8). False spider mite, Brevipalpus phoenicis, is
sometimes found on undersurface of leaves. Light brownish spots are found on
infested leaves because of continuous feeding by mites. Sometimes spider mite,
Tetranychus urticae, appears in minor proportions during hot and dry weather
(Dhooria 2005a, b).
Fig. 13.6 Spider mite damage on castor leaf

Fig. 13.7 Broad mite

damaged sesamum plant
Fig. 13.8 Broad mite damaged sesamum field

13.4.6 Safflower (Carthamus tinctorius; Asteraceae)
Spider mite, Tetranychus macfarlanei, is reported in minor proportions infesting

safflower plants from India. Red-legged earth mites (H. destructor) and blue oat
mites (Penthaleus major) are also reported infesting safflower, and maximum mite
population is found during dry seasons. Purplish blue mites with bright-coloured
legs are found under clods of soils or on the underside of leaves during hot weather.
Acaricide sprays should be initiated before significant damage occurs during early
growth.
13.4.7 Sunflower (Helianthus annuus; Asteraceae)
Spider mites, Tetranychus urticae, and false spider mite, Brevipalpus phoenicis, are
reported in minor proportions particularly near harvest of the crop.
13.4.8 Jatropha (Jatropha curcas; Euphorbiaceae)
Tarsonemid mite, P latus, is sometimes reported in serious proportions during hot

and humid months from West Bengal (India). Mite attack is more serious on young
apical leaves. Infested plant parts curl and become twisted causing flowers and buds
to drop down. Crop growth is retarded.
13.5 Plantation Crops
13.5.1 Sugarcane (Saccharum officinarum; Poaceae)
Sugarcane is native to the warm temperate or tropical regions of South Asia mainly
cultivated in tropical or temperate climate in Australia, Brazil, Philippines, India
and Thailand.
Spider Mite (Oligonychus indicus (Hirst)) Sometimes O. Indicus is found as a

sporadic pest on sugarcane, maize and sorghum. Mites remain active on monocoty-
ledonous weed, Sorghum halepanse, throughout the year. Low humidity and high
temperature are favourable for build-up of this pest. These mites live and feed on
the underside of leaves; they form fine webs in which eggs are laid and young
nymphs develop. Leaves infested by these mites often develop a red russetting
colour (Fig. 13.11); mite infestation generally occurs during May–June. The lower
leaves of sugarcane are usually colonized first; damage may be of less significance.
However, prolonged heavy infestation accompanied by extensive damage to the
middle and upper leaves of young plants may reduce growth. Rainfall significantly
controls mites. Hot and dry climate and water-stressed plants are favourable for
mite build-up (Gupta et al 1975).
13.5 Plantation Crops 295
Fig. 13.9 S. andropogani

damaged sugarcane plant
Web Spider Mites (Schizotetranychus andropogani Hirst) S. andropogani has

been reported from India and Mexico on sugarcane infesting sometimes very seri-
ously. Weed plants like Dichanthium annulatum are also reported infested by web
spider mites. Mites live under thin webs in large colonies on either undersurface of
leaves on either side of the midrib. Feeding causes white blotches; mite infestation
is more on ratoon crops than on the main season crops (Figs. 13.9 and 13.10).
Population is more on sugarcane during post-monsoon period during August–
September; mite infestation is also reported on paddy leaves (Gupta et al 1972).
Spider Mite (Oligonychus stickneyi (McGregor)) Oligonychus stickneyi is

reported as occasional pest on sugarcane in Florida (USA), feeding and breeding on
undersurface of leaves. Mite-infested leaves develop a red russetting, and severe
damage may result in leaf death. Infestations generally occur during March–June;
lower leaves are colonized first. Damage may often be of little consequence; how-
ever, prolonged heavy infestation accompanied by extensive damage to the middle
and upper leaves of young plants may reduce growth. Mites prefer midrib regions
for feeding and development but spread to other areas afterwards. Along field bor-
ders, mite infestations are common which later spreads throughout the field.
Fig. 13.10 Close-up of S.

andropogani damage
Banks Grass Mite (Oligonychus pratensis) Banks grass mite also sometimes
infests sugarcane leaves and results in reduced production.
Eriophyid Mite (Aceria sacchari Wang) Aceria sacchari Wang is also sometimes
reported in serious infestations from India. A. sacchari feeds on the inner side of the
leaf sheaths forming a circular erineum. The blisters (about 9 mm diameter) are
initially light green but slowly turn rusty red and get hardened afterwards. In the
later stages of mite infestation, the outside of leaf sheath exhibits swollen scars.
Other eriophyid mites (Abacarus sacchari Channabasavanna and A. doctus Navia
and Flechtmann) are also reported from sugarcane in India and Brazil; A.
queenslandiensis is also reported from Australia as vagrant feeders.
Tarsonemid Mite (Steneotarsonemus bancrofti Michael) Steneotarsonemus

bancrofti Michael is also reported as a minor pest on sugarcane from India. Adult
mites are light greenish, and their feeding causes flat rusty clusters and scabby cor-
roded patches on underside of leaf sheath and on internodes. Mite feeding produces
transparent depression on young stalk, which later become rough and opaque, and
the whole stalk is covered with roughened surface.
Fig. 13.11 O. indicus

damaged leaf vs. healthy
sugarcane leaf
13.5.2 Cassava (Manihot esculenta; Euphorbiaceae)
It is also known as tapioca in India. Cassava typically grows as a shrub, is native of

South America and is extensively cultivated as an annual crop in tropical and sub-
tropical regions for its edible starchy roots.
Cassava Green Mite (Mononychellus tanajoa (Bonder); CGM) The cassava

green mite (CGM) is reported from the South America and is spreading quickly to
all cassava-growing areas through wind and the movement of infested planting
materials. Peak CGM densities occur during the first half of the dry season, with a
smaller peak occurring within about a month of the start of long rainy season. Active
stages of mite feed and reproduce on undersurfaces of the leaves by sucking sap
from cells causing yellow spotting of leaves which can increase from a few spots to
complete loss of green pigment in leaves. Severely damaged leaves dry out and fall
off (ER21). Because of reduced plant growth, accumulation of starch in the storage
roots is slowed, and sometimes even up to 50 % losses in root yields may occur.
Where leaves are eaten as vegetables by farmers, a corresponding leaf loss ensues.
Reduced growth and stunting of the tips is also responsible for contorted and thin
stems, thus affecting the planting material to be used for the next season. Young
plants are more infested than older plants; damage severity is greater during dry
season; and heavy rainfall can reduce CGM populations (Pillai and Palaniswami
1985; Yaninek and Hanna 2003).
Crops grown in poor soils are more susceptible to mite attacks. Always use clean
plant material for planting, and plant at the onset of the rains to encourage vigorous
growth and thereby increasing tolerance to mite attack. Cassava plants aged
2–9 months are most vulnerable to mite infestation. From Nigeria, practising inter-
cropping with pigeon pea has been reported to suffer less damage from CGM than
that grown on a pure stand. Predatory mites, namely, Typhlodromus aripo De Leon
and Amblydromalus manihoti (Moraes), introduced from South America – the home
of the CGM – have given effective control of CGM in Africa.
Red Spider Mites (Oligonychus gossypii and Tetranychus spp.) Several species
of red spider mites have been reported on cassava, mostly on the older leaves.
Infestation by these mites causes initially yellowish pin pricks along the main vein
of mature leaves. Spider mites produce protective webbings that are readily seen on
plant. Attacked leaves turn reddish, brown or rusty in colour. If mite infestation is
very high, leaves die and drop beginning with older leaves. Most damage occurs at
the beginning of the dry season. For managing these mites, conserve natural ene-
mies, and avoid planting next to infested fields. Avoid using broad-spectrum pesti-
cides particularly pyrethroids as these may lead to spider mite outbreaks.
13.5.3 Tea (Camellia sinensis; Theaceae)
Tea is an evergreen plant that grows mainly in tropical and subtropical climates.
Accurate identification of the mite pest is essential for making sound pest manage-
ment decisions. Early detection is often critical to eventual success in managing
mite pests and in reducing economic losses caused by them.
Red Spider Mites (Oligonychus coffeae (Nietner) The adults are about 500 μm
with a pink or red colour. The eggs are round, dark red and shiny. Mite population
grows fast under hot and dry conditions. A female lays 2–3 eggs per day on the
leaves, and fecundity ranged from 40 to 90 eggs. Egg stage lasts 3–8 days. The life
cycle is completed in 10–14 days. Nymphs and adults of spider mite suck the cell
contents producing tiny pale spots or scars. Mite-infested leaves have many tiny
white spots which are in fact the empty shed mites. Red spider mite is a polypha-
gous pest (Banerjee 1979; Cranham 1966; Das 1959; Hazarika et al. 2009).
Leaves of severely damaged plants turn reddish and drop prematurely. On upper
leaf surfaces as a result of extensive mite feeding, reddish-brown patches or bands
are visible starting along the midrib region and then spreading outward. In severe
cases of mite infestation, the entire leaf may become red. Crop losses from 17 to
46 % are reported. High temperatures, dry conditions and the absence of shade are
conductive to outbreak of this pest. Red spider mite is widely distributed on tea in
India, Bangladesh, Sri Lanka, Taiwan, Kenya and Uganda. When mite population is
very high, plants may be totally defoliated. Mite damage progresses from older
leaves upwards to the younger growth. Although mites and their eggs are present on
both leaf surfaces, the mites appear to prefer the upper leaf surface. Plants in the
establishment stages may be more damaged by the mite. Tea red spider mite also
infests cotton, castor, jute, mulberry, shade tree (Grevillea robusta) and some other
plants (Hazarika et al. 2009).
Pruning for red spider mites is less effective for management as compared to
scarlet mites. High-pressure water sprays and heavy rain dislodge mites. Generally
mite population explode after pesticide usage as they kill natural enemies of mites.
So application of broad-spectrum pesticides and pyrethroids should be used to the
minimum possible as they induce outbreaks of mites. Better field tending, balanced
nutrition and proper irrigations during dry season can manage red spider mites
properly.
Flat Mites (Brevipalpus phoenicis, B. australis, and B. obovatus;Tenuipalpidae) Flat

mites are also known as red and black flat mites or scarlet mite. Flat mites are impor-
tant pests of tea in Bangladesh, Sri Lanka, India and China. Adult mites are scarlet red
in colour, are obovate in shape and are about 300 μm size. Clusters of bright reddish-
orange eggs are found on leaves. In general, mites look light to dark green or reddish
orange; however, depending on temperature, females may have H-shaped black mark.
Flat mites are commonly found on banana, lemon, papaya, passion fruit and many
other plants. These mites are concentrated on the lower surface of leaves especially at
the leaf base and the petiole. Infested leaves have brown scurf followed by splitting of
petiole and defoliation. The scarlet mite, B. phoenicis, is reported in all tea-growing
areas causing 8–13 % crop losses. Pruning greatly reduces the population of scarlet
mites, and as the population growth of mites is quite slow, it rarely becomes abundant
until three or more years after pruning.
Broad Mite (Polyphagotarsonemus latus; Tarsonemidae) Broad mites are also

known as yellow mites and feed on plant sap causing scarring and distortion of the
leaves and stems. As a result of mite feeding, undersurface of leaves turns to a
brown, corky surface. Internodes get shortened and deformed. For mite feeding,
younger terminal growth (especially 2–3 leaves and the buds) is more preferred.
Plants in the greenhouse or nursery are highly susceptible. Twisting and stunting of
leaves (main terminal) are commonly observed. Females lay distinctly oval eggs
having 5–6 rows of whitish bumps. A good hand lens (10× or more) is needed to
observe these eggs. The life cycle is completed in about 4–6 days. The females are
about 150–200 μm long; males are slightly shorter and broader. A pale white stripe
runs longitudinally down the back of the female. Live mites are a light translucent
yellowish green.
For proper management of broad mites, sprays should particularly be directed at

the newest growth. Proper monitoring of mite during warm and humid period after
7–10 days particularly on terminal growth proves helpful in lowering the mite popu-
lation. Yellow mite is a polyphagous pest attacking tomato, cinchona, cotton, pulses,
potato, chillies, etc.
Pink Mite (Acaphylla theae; Eriophyidae) Pink mite, A. theae, is reported as an

important pest of tea causing considerable crop loss in India, Bangladesh, China,
Malaysia and Taiwan. The mite prefers young leaves and as a result of feeding
leaves turns pale and curls upwards. Severely infested leaves become leathery and
assume a ‘glossy buff’. Adult mites are orange coloured and are carrot shaped.
Mites are found mostly on the lower surface of tender leaves mainly on top 10–15 cm
of bushes. Life cycle of mite is completed in 7–10 days.
Purple Mite (Calacarus carinatus (Green); Eriophyidae) Purple mite is known

as an important pest of tea in South India. Mite-damaged leaves are characterized by
the ‘coppery brown’ discolouration. The infested leaves are covered with white cast-
off skins of mites along with the live mites. The entire leaf becomes dull (not
shining) and changes colour (remains green but tinted with bronze or purple colour).
Adult mites are microscopic, spindle shaped and purple in colour having five longi-
tudinal waxy ridges on the dorsal side. High mite infestation is found on the mother
leaves and maintenance leaves at the middle level of the tea bushes. Development
period lasts from 6 to 11 days. Assam tea is more susceptible to purple mite.
13.5.4 Cardamom (Elettaria sp.; Zingiberaceae)
Red spider mite is a minor pest and occurs on cardamom in small numbers during
summer months without causing serious damage and sometimes appears as pest in
nurseries. The mite becomes serious pest only in plantations where insecticide spray
is too frequent. Frequent spray of oxychloride on cardamom also induced red spider
mites. The mites spin webs on the undersurface of the leaves and live inside the
webs resulting in formation of characteristic white blotches on the leaves reducing
production of plants. The effective leaves look dirty white, dry up and lead to ulti-
mately mortality of infested plants (Anonymous 2009).
13.5.5 Areca Nut (Areca catechu; Arecaceae)
It is also known as areca palm and grows in much of the tropical Pacific Asia and
East Africa. False spider mite, Raoiella indica, is an important sucking pest on
young areca nut palms during dry weather. Eggs are reddish; mites feed on under-
surface of leaves. Spider mites (Tetranychus fijiensis and Oligonychus indicus) are
reported as a pest of areca nut from Kerala (India).
13.6 Fodder Crops 301
13.5.6 Rubber (Hevea brasiliensis; Euphorbiaceae)
The rubber tree requires a tropical or subtropical climate. Phytophagous mites,

Calacarus heveae (family Eriophyidae), Tenuipalpus heveae (family Tenuipalpidae)
and Eutetranychus banksi (McGregor) (family Tetranychidae), are reported infest-
ing rubber trees from some parts of Brazil.
13.5.7 Cashew (Anacardium occidentale; Anacardiaceae)
Cashew cultivation in the world is concentrated in the tropical portion of the globe.
India, Brazil and Vietnam are the major producers of cashew. Spider mites
(Oligonychus mangiferus) are reported mainly under prolonged drought conditions.
Mites feed on the undersurface of leaves, and affected leaves become whitish and
loose lustre. Normally mites appear as minor pests and many times no control mea-
sures are required.
13.6 Fodder Crops
Fodder crops are alternatively known as forage crops. Forage crops are grown as
annual or biennial and are grown to be utilized for cattle grazing or harvesting as a
whole crop. Herbaceous legumes (alfalfa or lucerne and clovers), maize, oats and
sorghum, are all used as fodder.
13.6.1 Alfalfa or Lucerne (Medicago sativa; Fabaceae)
Red-legged earth mite (Halotydeus destructor) and blue oat mite (Penthaleus major)
are among the main pests of lucerne. These mites feed on cotyledons and leaves of
seedlings and first-year plants and on leaves of older plants primarily in autumn,
winter and spring. Damaged seedlings and plants progressively show yellowing and
then whitening of cotyledons and/or leaves. Mite infestation can reduce vigour, pal-
atability and winter production of older plants. Heavily infested seedlings and first-
year plants may be killed or become stunted and weak. Both the mite species prefer
light, sandy loam and well-drained soils. They can be managed by timely irriga-
tions; and removal of other weeds growing along edges of fields during winter
reduces overwintering sites and early infestation of these mites.
Alfalfa spider mite (Tetranychus urticae) infests lucerne plants; and as a result of
mite feeding, initially stipplings on leaves are found; later infested leaves may des-
iccate and fall if too much mite infestation for more period is found (Fig. 13.12).
Timely irrigation helps in managing mites. Clover mite (Bryobia praetiosa) is found
feeding on alfalfa, clovers, peas and many weeds. These mites rarely cause much
damage, but their infestation caused a ‘silvered appearance’ due to loss of chloro-
phyll. Clover mites, however, become a problem when they invade houses/buildings
Fig. 13.12 Spider mite

damaged berseem twig
during March–April, fall and early winter months as these mites leave reddish stains
if they get crushed on curtains, bed spreads, walls, etc. Adult females lay small,
round, red eggs in bark crevices at the bases of tress or in cracks and crevices of
buildings in spring (before temperature reaches 32 °C). Eggs remain dormant dur-
ing the summer and begin to hatch as the temperature drops below 32 °C.
13.6.2 Rye Grass (Lolium sp.; Poaceae)
Winter grain mite (Penthaleus major) is an important mite pest of rye grass. Mite-
infested fields have greyish or silvery appearance caused by removal of plant sap and
chlorophyll from the plants. Maximum damage occurs from high mite infestation in
the fall and again in spring. Seedling grasses can be stunted or die if high mite popu-
lation appears on seedling stands in the fall and the winter is exceptionally cold.
13.6.3 Vicia (Vicia spp.; Fabaceae)
During cool and moist period (autumn to spring), red-legged earth mites (H. destruc
tor) are commonly found on Vicia species. Generally mite infestations are worse in
seasons with a cool showery spring and dry summers in Australia. Canola, peas and
lupins are also commonly infested by red-legged earth mites. Mite feeds by rasping
leaves (upper surface) with sharp chelicerae and sucks exuding plant sap. Infested
plants remain stunted producing little forage.
13.6.4 Timothy (Phleum pratense; Poaceae)
Timothy hay is a profitable cash crop and competes alfalfa in some cases as a fodder
for horses. It is a perennial grass native to most of the Europe.
Cereal Rust Mite (Abacarus hystrix Nalepa) Sometimes cereal rust mite infesta-
tion causes yield reduction from 30 to 70 %. These mites overwinter in the adult and
egg stages. Eggs from overwintering female mites are deposited in the leaf vein
groves and look like spherical yellow balls. Hatching of eggs begins in March, and
peak population of mite is found during April. Damage is most evident in April–
May when mites are abundantly found in the youngest tissues of plants. Mites prefer
cooler temperatures and tend to be less active during summer months. They are
found in the crowns of plant during winter. Damaged symptoms appear as retarded
growth, stunting and plant discolouration. Severe mite infestation causes substantial
yield losses and hay quality. A. hystrix is also known to vector ‘rye grass mosaic
virus’ (RMV) – a serious disease of temperate grasslands causing yield reductions
of pastures. Wheat-curl mites (Aceria tosichella) are also reported damaging
timothy.
Spider Mites (Tetranychus mcdaniel, T. urticae and Oligonychus pratensis) Spider

mite feeding can cause damage like leaf bronzing, stippling or scorching. In timothy
economic loss is caused by a drop in colour quality due to reduction in photosynthe-
sis. Water stress, wind and dust contribute to the potential for outbreak of mite popu-
lations. Winter grain mites (Penthaleus major) sometimes cause minor damage on
timothy.
13.7 Conclusions
Field crops refer to plants of the same kind that are grown on a large scale for food,
clothing and other uses by humans. Information available on various mite pests of
different crops grown as cereals, pulses, fibre crops, oilseed crops, plantation crops
and fodder crops is considered. Mite infestations wherever resulted in economic
losses particularly in case of most economic crops are discussed more elaborately.
Mite pests reported as major pests, most susceptible stage of the crop and the period
during which they are most damaging are given for each crop. Stress has been laid
on variable damage caused by different species of spider mites, false spider mites,
eriophyid mites and other mites because of their feeding damage; serving as vectors
of different diseases on many crops is mainly given, and information available on

their virus–vector relations is discussed. Biological information available regarding
different mite pests, type of damage caused by them, different stages of crop growth,
how they pass through unfavourable weather extremes and management options
under each situation is included particularly for most important mite species caus-
ing economic losses on main crops. Cultural practices playing useful role in man-
agement of mites on different crops are emphasized. Several mite species that may
cause potential damage under certain environmental conditions are also considered
under different crops.
Further Reading
Ahuja DB (1994) Seasonal incidence and chemical control of oriental mite, Eutetranychus orien
talis (Klein) on castor. Indian J Ent 56:1–5
Anonymous (2005) Identification and cultural control of insect and allied pests of canola. Bulletin
4650, Dept. Agri., 28 pages, State of Western Australia. (ISSN 1448–0352. year 2005). Dept.
of Agriculture, South Perth
Anonymous (2009) Cultivation Practices for Cardamom (Elettaria cardamomum). Spices Board,
Ministry of Commerce and Industry, Govt. of India, Cochin. 682 025, pp 40
Banerjee B (1979) Intra-tree variation in the distribution of the tea spider mite. Oligonychus cof
feae (Nietner). Acarologia 2:216–220
Cranham JE (1966) Tea pests and their control. Annu Rev Entomol 11:494–514
Das GM (1959) Bionomics of the tea red spider, Oligonychus coffeae (Nietner), on tea in north-
east India in relation to pruning and defoliation. Bull Entomol Res 57:433–436
Dhooria MS (2005a) Tarsonemid mite, Polyphagotarsonemus latus (Banks) (Acari:Tarsonemidae),
a serious pest of green gram (Vigna radiata) in Punjab. Insect Environ 9(3):103–105
Dhooria MS (2005b) Effect of different ecological parameters on infestation of tarsonemid mite
Polyphagotarsonemus latus on sesamum. J Appl Zool Res 16:137–138
Gupta SK, Dhooria MS (1973) Occurrence of Hemitarsonemus latus (Banks) on black gram and
its control in Punjab. Sci Cult 39:401–402
Gupta SK, Sidhu AS, Dhooria MS (1972) Occurrence of Schizotetranychus andropogani (Hirst) on
sugarcane in the Punjab and its control. Indian Sugar 22:335–337
Gupta SK, Dhooria MS, Sidhu AS (1975) Effect of food and temperature on the development,
longevity and fecundity of sugarcane red spider mite, Oligonychus indicus (Hirst). Acarologia
16:436–440
Gupta SK, Dhooria MS, Sidhu AS (1976) Seasonal abundance of Tetranychus telarius on castor in
the Punjab. Oilseeds J 6:16–18
Gutierrez J (1994) Acari: leaf -feeding mites. In: Mathews GA, Tunstall JP (eds) Insect pests of
cotton. CAB International, Wallingford, pp 407–424
Hazarika LK, Bhuvan M, Hazarika BN (2009) Insect pests of tea and their management. Annu Rev
Entomol 54:267–284
Lateef SS, Reed W (1990) Insect pests of pigeon pea. In: Singh SR (ed) Insect Pests of Tropical
Food Legumes. Valey, Chichester, pp 451–242
Pillai KS, Palaniswami MS (1985) Spider mites of cassava, vol 1, Technical bulletin series. Central
Tuber Crops Research Institute, Trivandrum, p 17
Rahman S, Khan MR (2012) Field reaction of some jute(Corchorus spp.) varieties to yellow mite,
Polyphagotarsonemus latus Banks. Asian J Biol Life Sci 1:101–103
Reed W, Lateef SS (1990) Pigeon pea: pest management. In: The pigeon pea Nene YL, Hall SD,
Sheila VK (eds). CAB International, Wallingford, pp 349–374
Further Reading 305
Ridsdill-Smith TJ (1997) Biology and control of Halotydeus destructor (Tucker)(Acarina:

Penthaleidae): a review. Exp Appl Acarol 21:195–224
Sandhu GS, Singh B, Dhooria MS (1975) Effect of rain on the population of Oligonychus indicus
(Hirst) (Acari: Tetranychidae) on different varieties of maize (Zea mays) in the Punjab, India.
Int J Acarol 1:10–13
Shanover TG, Romeia J, Minja EM (1999) Insect pests of pigeon pea (Cajanus cajan) and their
management. Annu Rev Entomol 44:77–96
Wilson LJ (1993) Spider mites (Acari:Tetranychidae)affect yield and fiber quality of cotton.
J Econ Entomol 86:567–582
Yaninek S, Hanna R (2003) Cassava green mite in Africa – a unique example of successful classical
biological control. In: Neuenschwander P, Borgemeister R, Langewald J (eds) Biological con-
trol in IPM systems in Africa. CABI Publishing, Wallingford, pp 61–75. ISBN 0-85199-639-6
1. https://www.youtube.com/watch?v=J3i5kzP1PkE Wheat Streak Mosaic Identification and

Management
2. https://www.youtube.com/watch?v=4aOXM88r_XM Mite identification and control in winter
wheat
3. https://www.youtube.com/watch?v=SeGSCZG_uAY Aceria tulipae on wheat
4. https://www.youtube.com/watch?v=KTKNOk65fXM Mite Movement
5. https://www.youtube.com/watch?v=uippXe-CCXA Management of wheat streak mosaic and
similar mite-transmitted diseases
6. https://www.youtube.com/watch?v=bxagRDR1Ft4 Spider Mites in Droughty Soybeans
7. https://www.youtube.com/watch?v=55qqfX_0c-U Spider Mites in Soybean
8. https://www.youtube.com/watch?v=GyUYqnt-OLc Spider Mite Farmer: Mass rearing on
beans
9. https://www.youtube.com/watch?v=lQFtezWyeAY Spider Mite Damage on Beans
10. https://www.youtube.com/watch?v=2hB84S7p6vY Sampling spider mites in cotton
11. https://www.youtube.com/watch?v=PexkYjLQnoU Broad mites in cotton
12. https://www.youtube.com/watch?v=oUcatfAOOQM Spider mites in cotton
13. https://www.youtube.com/watch?v=2hB84S7p6vY Sampling spider mites in cotton
14. https://www.youtube.com/watch?v=N0RxYTBZnOA Tarsonemid Mite I
15. https://www.youtube.com/watch?v=W0uIMvD_Sew Control of Panicle Mite in Rice fields
16. https://www.youtube.com/watch?v=swbn2w4GvVU Mites in corn
17. https://www.youtube.com/watch?v=eeIOtqOG6sg Spider Mites of Sweet Sorghum
18. https://www.youtube.com/watch?v=ThTI5IvFjYY Scouting for Spider Mites in Corn
19. https://www.youtube.com/watch?v=v1PgDdSzDiw Scouting for Two-spotted Spider Mites in
Field Corn
20. https://www.youtube.com/watch?v=O4PwbL_-N-k Spider Mites on Cannabis
21. http://www.plantwise.org/KnowledgeBank/Datasheet.aspx?dsid=34767 Cassava green mite
(Mononychellus tanajoa)
Mite Pests of Greenhouse Crops
14
Abstract
A greenhouse (also called as glasshouse) is a building in which plants are grown.
Size of greenhouse may range from small shed to industrial-sized buildings.
Commercial glass greenhouses are fitted with equipment such as screening
installations, heating, cooling and lighting which may be automatically con-
trolled by a computer. Greenhouse cultivation is common in Spain, the
Netherlands, the UK and many other European countries for growing costly
flowers and plants. In the USA, Korea and Israel, greenhouse cultivation is quite
common. Several species of spider mites damage a large number of greenhouse
vegetables and ornamentals worldwide. Cucurbits, French beans and a variety of
foliage and flowering ornamental plants are among the most affected crops in
greenhouses. Spider mites particularly damage begonias, cacti, chrysanthemums,
cyclamens, dahlia, petunias, roses, verbenas and zinnias. Hot and dry conditions
favour development and multiplication of spider mites. False spider mites often
damage orchids in greenhouses. Broad mites and cyclamen mites also sometimes
appear as important pests on different plants in greenhouses. Tomato russet mites
sometimes infest tomato and other solanaceous plants particularly in warm and
temperate areas and cause economic damage. Occasionally bulb mites belonging
to family Acaridae infest and damage flowers that are reproduced from bulbs,
corms and tubers. Some phytoseiid mites are available commercially through
insectaries and are being used successfully for biological control of spider mites
in greenhouses. Management of mite pests in greenhouses is discussed.
Keywords
Greenhouse • Management • Biological control • Phytoseiid mite
A greenhouse (also called a glasshouse) is a building in which plants are grown.

These structures range in size from small sheds to industrial-sized buildings. A
greenhouse is a structural building with different types of covering materials such as

DOI 10.1007/978-981-10-1594-6_14
308 14 Mite Pests of Greenhouse Crops
a glass or plastic roof and frequently with glass or plastic walls. Greenhouses can be
divided into glass greenhouses and plastic greenhouses. Plastics used are polyethyl-
ene film and multiwall sheets or polycarbonate. Commercial glass greenhouses are
fitted with equipment such as screening installations, heating, cooling and lighting
and may be automatically controlled by a computer (ER2).
Greenhouses allow for greater control over the growing environment of plants.
Depending upon the technical specification of a greenhouse temperature level, light
and shade, irrigation, fertilizer application and atmospheric humidity can be con-
trolled. Greenhouses may be used to overcome shortcomings in growing qualities of
a piece of land such as a short growing season or poor light levels, to enable certain
crops to be grown throughout the year that are important for the food supply of
high-latitude countries. In Spain largest complex of greenhouses cover almost
50,000 acres (200 km2). Greenhouses are often used for growing flowers, vegeta-
bles, fruits and transplants. Greenhouse cultivation is common in the Netherlands,
UK and many other European countries for growing costly flowers and plants. In
the USA, Korea and Israel, greenhouse cultivation is quite common. A number of
grape growers, rose cultivators and producers of coloured capsicum and many
growers of high-value ornamental flowers are switching to greenhouses for cultiva-
tion of their crops as they are more profitable (Albajes et al. 1999; Hussey et al.
1969).
The greenhouse environment is characterized by conditions that optimize plant
growth. However, it is not only plants that benefit from the stable greenhouse envi-
ronment but also herbivorous insects and mites. In addition, greenhouses often pro-
vide herbivores with unlimited amount of food plants in monoculture and lack of
natural regulating factors such as predators, parasitoids and diseases. On top of this,
crop cultivars have for generations been selected for quick growth and maximum
yields often resulting in reduction or even loss of resistance mechanisms against
herbivores. Hence greenhouse crops are quite vulnerable to herbivore attack.
Herbivores that are accidentally introduced into greenhouses or migrate into green-
houses through open vents will most likely find almost all biotic and abiotic condi-
tions in favour of their rapid population increase. Hence, herbivores from quite a
number of insect and mite pests have attained pest status on greenhouse crops.
14.1 Mite Pests in Greenhouses
14.1.1 Spider Mites
Spider mites (family Tetranychidae) feed mainly on the undersurface of leaves, by

inserting their stylets and sucking epidermal and mesophyll cell contents. Damaged
cells have a reduced number of chloroplasts, and this can be seen from the upper
surface of leaves as yellowing punctures that later coalesce and form bigger yellow-
ing areas; mite-damaged leaves fall off prematurely (Helle and Sabelis 1985).
14.1 Mite Pests in Greenhouses 309
Cytological damage is accompanied by chemical and morphological alterations in

the plants that leads to retardation of growth and yield loss (ER7 and ER8).
Several species of spider mites (mainly Tetranychus spp.) damage a large num-
ber of greenhouse vegetables and ornamentals worldwide. Cucurbits, French beans
and a variety of foliage and flowering ornamental plants are among the most affected
crops in greenhouses. Many other crops such as tomatoes may also be infested
severely by two-spotted spider mite (TSSM), Tetranychus urticae Koch; carmine
spider mite Tetranychus cinnabarinus (Boisd.) and the Lewis spider mite
Eotetranychus lewisi (McGregor) are commonly found on a wide variety of green-
house plants. In greenhouses mites are of great significance, they are relatively
small and spend most of their time on the undersides of leaves which makes them
difficult to detect and effectively control. Spider mites have a short life cycle (i.e.
populations build up rapidly), have wide host range and are usually resistant to most
of the pesticides used for their control. Because of above traits, spider mites have
been designated as most severe pests under greenhouse conditions. Understanding
the biology of individual species is crucial for attaining their effective control.
Spider mites particularly T. urticae feed on ornamental flowering plants like bego-
nias, cacti, chrysanthemums, cyclamens, dahlias, fuchsias, gerbera, geraniums,
Hibiscus spp., lantana, petunias, roses, verbenas and zinnias. Hot and dry conditions
favour development and multiplication of spider mites (ER5 and ER6).
Two-spotted spider mites (TSSM) overwinter as bright orange adult females on
old crop debris, greenhouse structures and other plants in spring and become active
when new plants are planted. Eggs hatch in 2–15 days depending on temperature,
but development at temperatures below 12 °C is reduced. A life cycle may be com-
pleted in 2–3 weeks depending on temperatures, and many generations per year are
possible in greenhouses. Females can lay 2–3 eggs/day or 50–60 eggs in its lifetime.
Under ideal conditions, huge populations of TSSM can build up rapidly if left
unchecked. TSSM may be a pest of quarantine importance on some export crops,
e.g., cut flowers (Van Lantern 1995).
14.1.2 False Spider Mites
False spider mites belong to family Tenuipalpidae and look flat orange to red with
black spots, and its adults are small (about 300 μm in length). Eggs are somewhat
flattened, red and sometimes ridged. False spider mites have sharp needle-like
mouthparts and, through them, pierce the epidermis of the plant and suck out the
juices. These mites have very slow rate of incubation and development particularly
during cold weather. Damage by false spider mites includes silvering of leaves and
gall formation on stems. These mites do not construct webs unlike spider mites.
Most important false spider mite species often found in greenhouses are Brevipalpus
phoenicis (Geijskes), Brevipalpus californicus Banks and Tenuipalpus pacificus
Baker. False spider mites often damage orchids in greenhouses.
14.1.3 Tarsonemid Mites
Tarsonemid mites belong to family Tarsonemidae and these mites feed preferably
on young or succulent plant tissue. Important mite species causing a nuisance in
greenhouse crops are broad mite, Polyphagotarsonemus latus (Banks), and the
cyclamen mite, Phytonemus pallidus (Banks). Broad mite is largely polyphagous,
but severe damage has been reported under different conditions on sweet peppers,
tomatoes and cucumbers particularly during warm conditions in the greenhouses.
Terminal growth of mite-infested leaves turns bronzy and shiny, and later this
growth dries and appears burnt. Females lay 1–5 eggs per day during their 1–2 weeks
of life. One female may lay 10–20 eggs in its lifetime. Female longevity is consider-
ably longer than that of males (Weintraub et al. 2003). Cyclamen mite has a com-
paratively narrow host range and affects strawberries and some ornamental plants in
many parts of the world. Persistent mite feeding causes stunting and distortion of
newly emerged strawberry leaves; flowers wither and dry. During heavy infestations
strawberry plants become stunted having a compact mass of crinkled leaves in the
centre, and the fruit is dwarfed and appears seedy. The action of wind, flying insects
and machines help in dispersal of mites within and between field movements of
strawberry (ER9).
14.1.4 Rust Mites
Wormlike mites belong to family Eriophyidae; they are also commonly known as
rust or gall mites. These mites are quite host specific and feed on epidermal cells
causing morphological and physiological alterations including gall formation and
other tissue distortions, toxaemias and severe non-distortive effects such as russet-
ting, browning or silvering of leaves and other green plant tissues. Such physiologi-
cal alterations may interfere with photosynthetic activity and result in alteration of
supply of nutrients and adverse hormones activity. In addition rust mites may also
serve as vectors of plant pathogens. Different morphological and physiological
effects as a result of rust mite infestation may also lead to reduction in crop yields
or to reduced aesthetic value.
Tomato russet mite, Aceria lycopersici (Wolffenstein), is probably the most
widespread and harmful species on greenhouse-grown vegetables. In the green-
houses, rust mite infests tomato and other solanaceous plants particularly in warm
and temperate areas. The same mite is also widespread and harmful in outdoor
tomatoes. Mites feed on epidermal cells mainly on the upper surface of the leaf,
which turns bronze after a few days of mite infestation. Severe infestation may lead
to fruit russetting and plant desiccation and eventually death of the plant. In most
susceptible cultivars of tomato, mites also attack flowers resulting in flower bud
death. Initially mite damage is noticed near the ground but later spreads upwards. In
greenhouses, rust mite is first located in some plants and later spreads to other
nearby plants, ultimately spreading throughout the greenhouse-grown tomatoes.
14.2 Biological Control of Mite Pests 311
14.1.5 Bulb Mites
Bulb mites belong to family Acaridae. Two most common bulb mite species reported
worldwide are Rhizoglyphus echinopus (Fumouze and Robin) and Rhizoglyphus
robini Claparede. These mites inhabit the soil or potting material and are occasion-
ally pests in the greenhouse where they infest and destroy flowers that are repro-
duced from bulbs, corms and tubers. Bulb mites usually occur in groups or colonies.
All developmental stages are present throughout the year. Adult mites are
500 μm–900 μm long and shiny white. However, legs are reddish brown. Bulb mites
have been recorded feeding on lily, narcissus, orchids, tulip, dahlia bulbs and on
gladiolus corms. They also infest onion bulbs, garlic and potato tubers. Hypopi
(plural of hypopus) also called heteromorphic deutonymphs form in response to
overcrowding or the area in which they are feeding deteriorates. Female can lay
400–700 eggs at favourable temperature (24°–27 °C). There are some reports that
these mites are not primarily destroyers of these plants, but they can take over a bulb
or corm that has been physically damaged (bruised) or has been infected with fungal
or bacterial pathogen. These mites can survive on numerous food sources and are
often shipped long distances on bulbs, corms and tubers wherever bulb mites are
discovered (Lesua et al. 2000). Discarding of infested stock is probably the best
solution. Before planting, bulbs must be examined thoroughly to detect bulb mites,
and only healthy bulbs should be planted. Preventive control measures, if available,
should also be tried for the treatment of susceptible plants like lily. Bulb mites can-
not withstand drought, and dry bulbs in storage are usually not attacked. Steam
sterilization and methyl bromide at low concentrations eliminated the mites from
soil.
14.2 Biological Control of Mite Pests
14.2.1 Through Predatory Phytoseiid Mites
For long time predatory phytoseiid mite Phytoseiulus persimilis Athias-Henriot has
been utilized for biocontrol of TSSM (Cakmak et al. 2009; Copping 2009). In the
recent past, predatory mite, Neoseiulus (=Amblyseius) californicus (McGregor) is
increasingly being used providing efficient results under greenhouse cultivation.
Phytoseiid mites, P. persimilis and N. californicus, are available commercially
through commercial insectaries. Galendromus occidentalis (Nesbitt), Neoseiulus
(Amblyseius) fallacis (Garman), Amblyseius andersoni (Womersley) and Amblyseius
swirskii Athias-Henriot are also being made available through the commercial
insectaries for biocontrol of spider mites in different parts of the world (Gerson and
Weintraub 2007, 2012; Weintraub and Palevsky 2008). The use of different preda-
tory mites in greenhouses against spider mites depends on food requirements,
developmental rates and reproduction (Zhang 2003). Based on these criteria, preda-
tory phytoseiid mites are classified into the following three categories (ER1)
(McMurtry et al. 2013):
Type I category. These mites are specialized predators of spider mites (Tetranychus
spp.). P. persimilis have been found as specialized predators of TSSM, they lay
their eggs on the webs produced by TSSM, and all mobile stages freely move
among the mite webs. There is a strong tendency of P. persimilis not to move
away or disperse from infested areas as long as live prey or spider mites are pres-
ent (Weintraub et al. 2007; Zhang 1990). Type I species are more efficient preda-
tors of spider mites than Type II and Type III phytoseiids.
Type II category. They are also commonly associated with spider mites that produce
dense webbings, but these mites also feed on other types of mites such as russet
and cyclamen mites and are not as specialized as mites in Type I category. These
mites have lower food requirements for their development as compared to phy-
toseiid mite species in Type I and Type III categories. G. occidentalis, N. fallacis
and N. californicus are representatives of Type II category. These mites have high
powers of aerial dispersal when live spider mites become scarce in their immedi-
ate vicinity.
Type III category. Species in this category include A. andersoni, A. swirski,
Neoseiulus barkeri Hughes, N. cucumeris Oudemans and Iphiseius (Amblyseius)
degenerans Berlese. These predaceous mites are generalist predators as they
have a much broader food range from various mite species, pollen and insects
such as whiteflies, thrips and mealybugs. Unlike Type I and Type II, Type III
predatory mites are not associated strongly with spider mite colonies. Fewer
predatory mites in fact are found on leaves with high mite density than on leaves
with a low density, and members of this type are adversely affected by dense
webbings of spider mites. Depressions on leaf surface, leaf hairs or other shel-
tered areas seem more important for these mites than the available food.
Cannibalism and preying on other predatory mites may be more common than in
Type I and Type II specialists.
For the success of particular predatory mite in a biocontrol programme, besides

traits mentioned under three above categories, many more factors can contribute to
the success of a particular predatory mite. Efficiency of response to pest infesta-
tions, diet habits of larvae and nymphal stages, behaviour towards other predatory
mites and environmental conditions determine the contributions of a predator in a
biocontrol programme, but still much more remains to be done for optimum utiliza-
tion of different predatory mites (Driesche and Vittum 1987).
14.2.2 Through Predatory Insects
14.2.2.1 Gall Midge, Feltiella acarisuga Vallot (Insecta, Diptera,

Cecidomyiidae)
The predatory gall midge (Feltiella acarisuga) is one of the most effective and
widespread natural enemies of spider mites particularly TSSM in a number of crop-
ping systems and can be useful in integrated pest management of spider mites that
attack greenhouse crops. Females lay shiny, translucent oblong eggs (0.10–0.25 mm)
14.3 Management of Mite Pests in Greenhouses 313
near prey mites on leaves. Larvae (0.2–2 mm) have four developmental instars and
feed exclusively on all developmental stages of spider mites and sometimes occur in
populations as high as 160 per cm2 of eggplant leaf. Pupae are fluffy white (1.0–
1.5 mm long) and are mainly found on the underside of a leaf next to a vein. Adult
is a pink-brown fly (2 mm length) with long legs. Reproduction and development
occur at 15°–25 °C, and eggs and larvae do not survive above 30 °C and below 30 %
relative humidity. F. acarisuga larvae have been reported to consume an average of
15 adult mites, 30 developmental stages or 80 eggs per day. Commercially F. aca-
risuga pupae are available from suppliers of natural enemies (ER10).
(http://www.anbp.org/)
(http://www.cdpr.ca.ga//doc/ipmnov/bensuppl/contents.htm)
14.3 Management of Mite Pests in Greenhouses
The greenhouse environment is not only nearly ideal for growing plants; it can pro-
duce outstanding pest infestations. Mites can be one of the outstanding pest infesta-
tions in many greenhouse crops and production systems around the world. They are
most notorious in long-term, protected growing situations such as greenhouse toma-
toes, stock plants or cut flowers (Gerson 2014; Maonen et al. 2010; Pena and
Osborne 1996). For suitable management of spider mites and other mites, following
control measures should be followed.
Monitoring Early detection of spider mite activity is important to achieve good

control. Look for stipplings caused by mite feeding (ER4).
Cultural Control Start clean, and remove weeds surrounding greenhouse environ-
ment regularly. Inspect for mites at all stages of plant growth regularly. Remove old
debris at the time of new planting. Avoid planting outdoor crops near greenhouses
as they may serve as sources of infestation.
Biological Control Numerous predatory mites are available and may be usefully
tried for mite pest control.
Chemical Control Soaps and oils work well by contact; new miticides often have
ovicidal activity.
Integrated Programmes By combining cultural and biological controls with

effective acaricides (Wittmer and Castilla 1995).
Alternate Chemical Controls Spider mites commonly have developed resistance

to miticides, so do not use the same compound again and again if satisfactory con-
trol was not achieved. Always use alternate chemical groups on a regular basis if
spider mites need continual control. Without adopting different cultural and biologi-
cal controls, chemical controls will eventually fail.
14.4 Conclusions
Two-spotted spider mite (Tetranychus urticae; TSSM) is most destructive under

greenhouse conditions. Mite feeding on spongy parenchyma, palisade parenchyma
and chloroplasts results in reduced chlorophyll and adversely affects photosynthesis
of affected plants. The webbing of the infested plant parts is one of the most char-
acteristic signs of heavy mite infestation. High temperature (more than 28 °C) and
low relative humidity (20–40 %) are suitable for heavy mite infestation and destruc-
tive population explosions. For proper management of TSSM, monitoring plants at
3–4 days interval for observing signs of mite feeding injury is most important. But
look for mites on the undersides of mature leaves especially along mid-vein region,
or gently tap leaves over a white piece of paper to dislodge mites to monitor more
easily. The introduction of infested plant material into greenhouses or allowing to
grow mite-susceptible plants near the outer perimeter of the greenhouse may give
rise to mite population explosions, especially during the summer. It is much easier
for a grower to prevent a mite infestation than to eradicate it once it is established in
the greenhouse. Over fertilization promotes succulent new growth which is more
susceptible to TSSM. In greenhouses that use drip irrigation, foliage remains dry
and TSSM population may be higher. But the use of occasional overhead irrigations
helps in managing mite populations. Thorough coverage of both the lower and
upper leaf surfaces is critical when applying pesticides. Insecticidal soaps and hor-
ticultural oils are effective against TSSM; however, certain plants may be sensitive
to both these materials. Most miticides are not effective against egg stage, so their
repeated applications are needed after 5–7 days. Mite growth regulator such as etox-
azole (Tetrasan) can also be applied tank mixed with pesticide before extensive mite
populations are found.
For TSSM control, phytoseiid mites Phytoseiulus persimilis, Galendromus occi-
dentalis, Neoseiulus californicus and Neoseiulus fallacis are available commer-
cially for releases in greenhouses. Temperature and relative humidity requirements
vary for each species. G. occidentalis, N. fallacis and N. californicus tolerate warmer
conditions (>29 °C, 30–40 % R.H.). These predatory mites persist even when TSSM
populations are quite low.
For successful and early control of broad mites and cyclamen mites on green-
house ornamental plants (peppers, tomato, begonia, dahlia etc.), look for symptoms
of mite damage. Cyclamen mite feeding causes inward curling of leaves, crinkling
and puckering. Critically examine buds under the microscope for their eggs and live
mites. In case of broad mite infestation, leaf edges curl downwards. Infested leaves
have twisted and distorted growth.
Further Reading
Albajes R, Gullino ML, Lantern JC, Elad Y (eds) (1999) Integrated pest and disease management
in greenhouse crops. Kluwer Academic Publishers, Dordresch, p 545
Cakmak I, Janssen A, Sabelis MW, Baspinar H (2009) Biological control of an acarine pest by
single and multiple natural enemies. Biol Control 50:60–65
Further Reading 315
Copping LG (2009) The manual of biocontrol agents. BCPC, Alton, p 851

Driesche RL, Vittum P (1987) Potential for increased use of biological control agents against
greenhouse pests in Massachusetts. Res Bull Massachusetts Agril Expt Sta 718:788
Gerson U (2014) Pest control by mites(Acari): present and future. Acarologia 54:371–394
Gerson U, Weintraub PG (2007) Review: mites for the control of pests in protected cultivation.
Pest Manag Sci 63:658–676
Gerson U, Weintraub PG (2012) Mites (Acari) as a factor in greenhouse management. Annu Rev
Entomol 57:229–247
Helle W, Sabelis MW (1985) Spider mites: their biology, natural enemies and control. Elsevier,
Amsterdam
Hussey NW, Read WH, Hesling JJ (1969) The pests of protected cultivation. Edward Arnold,
London, p 404
Lesua J, Conijn CGM, Sabelis MW, Straalen NM (2000) Biological control of the bulb mite,
Rhizoglyphus robini, by the predatory mite, Hypoaspis aculeifer on lilies: predator–prey
dynamics in the soil under greenhouse and field conditions. Biocontrol Sci Technol
10:179–193
Maonen RF, Vila E, Sabelis EW, Jansen A (2010) Biological control of broad mites
(Polyphagotgarsonemus latus) with the generalist predator, Amblyseius swirskii. Exp Appl
Acarol 50:29–34
McMurtry JA, Moraes GJD, Sourassou NF (2013) Revision of lifestyles of phytoseid mites (Acari:
Phytoseidae) and implications for biological control strategies. Syst Appl Acarol 18:397–320
(http://dx.doi.org/10.11158/saa.18.4.1)
Pena JE, Osborne L (1996) Biological control of Polyphagotarsonemus latus (Acari: Tarsonemidae)
in greenhouses and in field trials using introductions of predaceous mites (Acarina:
Phytoseiidae). Entomophaga 41:27–36
Van Lantern JC (1995) Integrated pest management in protected crops. In: David D (ed) Integrated
pest management. Chapman & Hall, London, pp 311–345
Wittmer CH, Castilla N (1995) Protected cultivation of horticultural crops worldwide. Hort
Technol 5:6–23
Weintraub PG, Palevsky E (2008) Evaluation of the predatory mite, Neoseiulus californicus, for
spider mite control under hot arid field conditions. Exp Appl Acarol 45:29–37
Weintraub PG, Kleitman S, Mori R, Shapira N, Paleveky E (2003) Control of broad mites
(Polyphagotarsonemus latus) on organic greenhouse sweet peppers with the predatory mite
(Neoseiulus cucumeris). Biol Control 27:300–309
Weintraub PG, Kleitman S, Alchanatis V, Palevsky E (2007) Factors affecting the distribution of a
predatory mite on greenhouse sweet pepper. Exp Appl Acarol 42:23–35
Zhang ZQ (1990) Two-spotted spider mites (Acari: Tetranychidae) and Phytoseiulus persimilis
(Acari: Phytoseiidae) on greenhouse roses: special distribution and predator efficiency. J Econ
Entomol 88:352–357
Zhang ZQ (2003) Mites of greenhouses: identification, biology and control. CAB International,
Wallingford
1. https://www.youtube.com/watch?v=Vcagnbm8dY8 Using beneficial mites to manage pests in

greenhouses
2. https://www.youtube.com/watch?v=hZwyWfqSck4&list=PLDE1DD3408997871C
Greenhouse Tomato (Structure)
3. https://www.youtube.com/watch?v=Wzkj5PEtkt0&index=3&list=PLDE1DD34089978
71C Redpath greenhouse buildings range
4. https://www.youtube.com/watch?v=JzvJLW_5SsI Monitoring Greenhouse Pests
5. https://www.youtube.com/watch?v=kVCcdLxF_fY Commercial Pest Control in The Pepper

Greenhouse (Spider Mites), Part 1
6. https://www.youtube.com/watch?v=U8Rstw7zZbo Cucumber plants attacked by spider mites
7. https://www.youtube.com/watch?v=−yolQljY_6I&index=4&list=PLZ53P1UKadIlhOk0Lp
WMPxuOBxYjLQ-EQ Spider Mite Circus
8. https://www.youtube.com/watch?v=GyUYqnt-OLc Spider Mite Farmer
9. http://www.greenhousegrower.com/video/production/crop-inputs/insect-control/broad-
mites/ Broad Mite
10. http://entnemdept.ufl.edu/creatures/beneficial/f_acarisuga.htm Feltiella acarisuga
Management of Mites Under Organic
Farming 15
Abstract
Organic forming is a method of crop production that involves much more than
choosing not to use pesticides, fertilizers and genetically modified organisms.
Organic farming presents many challenges; some crops are more difficult to
grow organically. For suppression of spider mites in greenhouses, proper man-
agement of temperature and relative humidity can be useful. The knowledge
obtained from biological, ecological and behavioural information can be used to
plan the management of mite pests. An organic farmer with experience can know
how to manipulate the natural enemies of spider mites to protect their crops. For
managing mite pests, cultural practices that suppress mite pests but encourage
population of their natural enemies should be used. Elimination of crop residues
can destroy mite pests and prevent them from damaging subsequent crops. Poor
water management (especially too little water) can predispose plants to spider
mites and many other insect pests. Dry weather, low humidity and temperatures
exceeding 30 °C are associated with spider mite growth. Good farm sanitation
can help to prevent introductions of mite pests from outside sources and slow
their movement within the farm. Regular monitoring for mite pests and their
natural enemies is very important for their management.
Keywords
Organic farming • Cultural control • Irrigation management • Crop residues
Organic farming is a method of crop and livestock production that involves much
more than choosing not to use pesticides, fertilizers, genetically modified organ-
isms, antibiotics and growth hormones. Organic production is a holistic system
designed to optimize the productivity and fitness of diverse communities within the
agroecosystem, including soil organisms, plants, livestock and people. The princi-
pal goal of organic production is to develop enterprises that are sustainable and
harmonious with the environment (Oelaf 1978).

DOI 10.1007/978-981-10-1594-6_15
318 15 Management of Mites Under Organic Farming
Organic farming promotes the use of crop rotations and cover crops and encour-
ages balanced host/predator relationships. Organic residues and nutrients produced
on the farm are recycled back to the soil. Cover crops and composted manure are
used to maintain soil organic matter and fertility. Preventative insect and disease
control methods are practised including crop rotation, improved genetics and resis-
tant varieties. Integrated pest management (IPM), weed management and soil con-
servation systems are valuable tools on an organic farm. Protection of the
environment minimizing soil degradation and erosion, maintaining long-term soil
fertility by optimizing conditions for biological activity within the soil, maintaining
biological diversity within the system and recycling materials and resources to the
greatest extent possible are the general principles of organic production. Organically
approved pesticides include ‘natural’ or other pest management products included
in the Permitted Substances List (PSL) of the organic standards (Bengtron et al.
2005; Gurr et al. 2004; Willer and Yussefi 2006). Organic farming presents many
challenges. Some crops are more challenging than others to grow organically; how-
ever, nearly every commodity can be produced organically.
Some seed treatments are allowed in organic production as, for example, the
inoculation of legume seeds with beneficial bacteria. Transplants, starts or seedlings
used to produce an annual organic crop must have been organically grown particu-
larly in case of organic culture of tomatoes, sweet potatoes, peppers, flowers grown
from transplants (e.g. cut flowers of roses and snapdragons) and many other similar
crops. Whenever possible, organic planting stock should be used for organic pro-
duction. In fact planting stock for annual crops is subject to the same requirements
as seeds for annual crops like garlic, white potatoes and flowers grown from bulbs
(e.g. daffodils or tulips sold as cut flowers). However, planting stock for perennial
crops may be obtained from nonorganic sources but must be under organic manage-
ment for at least 12 months before the first harvest of an organic crop as in cases of
tree fruits (e.g. apples, peaches, pears), nuts (e.g. walnuts, pecans), berries (e.g.
blueberries, cranberries, strawberries), grapes, asparagus, lilies and others (Flaherty
and Wilson 1999; Hesler et al. 1993; Wyss et al. 2005; Weibal et al. 2004).
15.1 Management of Mites in Greenhouses
In greenhouse crop production, proper management of temperature and relative

humidity can be useful in suppressing mite pests. Soaking/showering plants is most
easy and effective method to eliminate small population of spider mites. But a
steady stream of water shower helps in dislodging mites from the leaves. Water
showering can be done 2–3 times daily if desired. Mites usually disappear after a
week of showering (ER1–ER4).
By creation of unfavourable environment for spider mites which normally thrive
in hot and dry weather, indoor gardens can suffer from such conditions if there is
inadequate ventilation and too much use of large glow lamps. But by increasing
ventilation and using an air conditioner, the temperatures can be brought below
16 °C. Humidity can be raised by misting the plants several times daily. But take
15.2 Management of Mites on Organic Crops 319
care to know that what temperatures the plants can accept, as 16 °C may be too cold
for some plants (Cook and Pickett 2007). Several fungal diseases of insects and
mites may develop rapidly if water spray is done on plants.
Conventional greenhouses production can rely on pesticides for controlling
insects and mite pests. In contrast many organic producers focus on a system
approach for insect, mite and disease management by planting a diversity of crops
using screens to exclude insects and mite pests, by introducing predators to reduce
pest mites and by breaking the pest cycle using heat or cold. In order to break the
pest cycle, choose a time when the greenhouse is not in use for crops and create
conditions unfavourable for survival of pest mites. A greenhouse used in the winter
can be closed in summer and allowed to heat to high temperatures to kill pest mites
and insects. Conversely a greenhouse used in the summer can be opened in the win-
ter so that pests are killed by cold temperatures (Brandenburg and Kennedy 1987).
15.2 Management of Mites on Organic Crops
Before planting a crop, an organic crop grower needs to understand the mite pests
and beneficial organisms that are usually occurring on the crop being grown and
anticipate the pest problems that are likely to occur. The experienced growers act
before problems reach devastating levels because they know that pests are more
effectively controlled when their numbers are low. The major emphasis of insect
pest management in organic crop production systems should be placed on the use of
cultural and biological control methods (Hokanen and Lynch 2003).
Among different mite pests that are found damaging plants under organic sys-
tems of cultivation, two-spotted spider mite Tetranychus urticae (family
Tetranychidae) is the most important pest. Management of mite pests presents a
challenge to organic farmers. On organic farms, the focus is on the management of
mite pests rather than eliminating them. So success depends on learning biological
information about the conditions in which they can survive, and such information is
used to determine if mite pests can be deprived of some vital resource. Ecological
and behavioural information about how the mites interact with the environment and
with other insect and mite species, if available, can be used to shape a pest-resistant
environment. An organic farmer with experience can know how to manipulate the
natural enemies of spider mites to protect their crops. The knowledge obtained from
biological, ecological and behavioural information can be used to plan the manage-
ment of mite pests. So mite pests under organic cultivation may be controlled
through cultural, mechanical or physical methods, augmentation or introduction of
predators of the pest species, development of habitat for natural enemies of mite
pests and utilization of non-synthetic control measures. When these practices are
insufficient to prevent or control mite pests, then biological or chemical control
measures with substances approved for use in organic crop production may be
applied to prevent, suppress or control these pests (Brandenburg and Kennedy
1987).
15.3 Cultural Control
For managing mite pests, cultural practices that suppress mite pests but encourage
population of their natural enemies should be used. By paying attention to crop rota-
tions and crop residue incorporation, the grower can help prevent population build-
up of many insect and mite pests. The practice of growing the same crop continuously,
season after season, on the same piece of land is an invitation to various mites and
insect pest species. In some locations, pest problems are so severe that it is not eco-
nomically possible to grow certain organic crops using available technology (Gurr
et al. 2004; Landis et al. 2000).
Organic crop should be grown by providing optimum growing conditions as
many plants are able to withstand insect and mite feeding if they are actively grow-
ing and are able to compensate for some loss of foliage. Soil type and nutrient
content can affect crop vigour. Proper rotation of pest-susceptible main crops with
non-susceptible varieties and cover crops can keep mite pests at low levels.
Breakdown of plant residues can be enhanced by soil tillage, thereby allowing com-
plete decomposition of organic matter in a field and enhancing the control of bulb
mites and other soil insects like cutworms and root maggots. Pest barriers can be
incorporated by floating row covers and plastic tunnels which reduce access by
many mites and insects. The incorporation of a grass species into a rotation is often
recommended as grasses tend to be resistant to most of the mite pests.
Field borders or strips within the field that are planted to plant species and variet-
ies different from the main crop can provide habitat for beneficial arthropods and
can slow spread of pest mites in the field. Flowering plants along borders and edges
of fields can provide habitat and food for beneficial insects and mites. Weed man-
agement in borders can be a challenge, and the grower will need to check the bor-
ders to make sure that pest mites are not building up there and migrating into the
cropping areas. Sometimes small changes where mite-susceptible plants are located
or how they are watered can greatly influence their susceptibility to spider mite
damage (Wyss et al. 2005; Zhender et al. 2007).
Proper managing of fertilizer applications is another important cultural practice.
Spider mite populations are stimulated generally if the plant is receiving a higher
dose of nitrogenous fertilizer than is needed. Succulent tender growth is favourable
for development of most spider mite species.
Elimination of crop residues can destroy mite pests and prevent them from dam-
aging subsequent crops. Polycropping (planting crop mixtures) reduces many pests
because the presence of the host plant is not as apparent if the crops that are planted
do not house the same plants. In many tropical and subtropical areas, polycropping
is traditionally found. Polycropping can be a useful proposition if natural enemies
are retained in the crop.
15.6 Other Cultural Practices 321
15.4 Water Management
Poor water management (especially too little water) can predispose plants to spider
mites and many other insect pests. Irrigations should be properly given so that hot
and dry conditions conducive for spider mite development can be prevented. There
should be proper management of dust along roads and paths, as dusts create condi-
tions for outbreaks of mites. Drought conditions are favourable for two-spotted spi-
der mite outbreaks. Dry weather, low humidity and temperatures exceeding 30 °C
are associated with spider mite population growth.
Dust can be a problem along edges of orchards, vineyards or fields or along road-
sides because of lack of irrigation along these areas. Effective irrigation is critical in
many climates and may affect spider mite populations. Water-stressed plants allow
rapid development of spider mites than the plants under well-watered conditions. A
lower relative humidity typically favours more spider mites, although some species
do well under humid conditions. A periodic hosing of plants with a forceful jet of
water can physically remove and kill many mites as well as remove the dust that
collects on foliage interfering with mite predators. Disruption of webbing may also
delay egg laying until new webbing is produced. Planting and harvesting dates of
organic crops should be coordinated to avoid certain mite pests that would other-
wise build up in some crops. Avoid staggered planting of the same crop with succes-
sive planting near earlier ones. In some areas, midsummer and late season plantings
must be located in isolated fields. Destroy old crop residues soon after final
harvest.
15.5 Good Farm Sanitation
Good farm sanitation can help to prevent introductions of mite pests from outside
sources, slow their movement within farm and eliminate them when they are dis-
carded with crop materials that may harbour them. If transplants are purchased off
farm, then buy only from a reputed dealer and check very carefully before bringing
transplants to the farm. Quarantine any purchased transplants for at least a week,
and examine them carefully for mite pests and other insect pests daily. Culled plants
and produce are often piled near the field or processing area for later disposal. This
parasite can provide a suitable feeding and breeding site for mites and other insect
pests. These piles should be composted, buried or otherwise destroyed as soon as
possible.
15.6 Other Cultural Practices
By providing alternate (non-host) prey, availability of nectar and pollen from flow-
ers; refugia (non-crop area) contributing for survival and persistence of predatory
mites and insects, besides providing them overwintering habitat; and keeping the
area more plant rich in general help in better action of natural control. However, it
is not possible to provide a blueprint for building a workable system of natural con-
trol on a specific farm. Various strategies needed for conservation and augmenta-
tion, or supplement the actions of naturally occurring predators, should be
considered. However, careful planning is critical to the success of a natural enemies
release programme. But the natural enemy selected should be correct for the spe-
cific situation and specific pest; and timing and rate of application should be
correct.
15.7 Monitoring for Mite Pests and Their Natural Enemies
The correct identification of the mite pests and their natural enemies present in a
crop is very important. Identification of immature life stages such as eggs, larvae
and nymphs will greatly help the growers to prevent economic damage to the crop.
However, an experienced pest control advisor can alert a grower to potential prob-
lems before the crop losses occur. Growers can sometimes avoid treating crops
against mite pests if natural populations of mite pests, their predators and microor-
ganisms are effectively monitored. So it is important that the growers make regular
weekly checks in the field with a hand lens, inspecting the underside of foliage for
the first signs of activity of spider mites. More frequent monitoring should be done
during hot and dry weather which is most suitable for spider mite development. The
grower must identify tolerance levels or action thresholds for the particular mite
pest species on a crop. If there is a pest mite that feeds on the fruit or on a part of the
vegetable that is sold, the tolerance for damage is generally lower. In pest feeding
on portions of the plant that will not be sold, such as leaves of tomato plant, the mite
population at more higher levels can be tolerated (Thomas and Waage 1996; Van
Lenteren 2003).
Early detection of spider mites before damage is noticed is important. Spider
mites can be detected by taking a piece of white paper or cardboard and striking
some foliage over it. The dislodged mites can be seen walking slowly on the paper.
If crushed, most plant-feeding mites will produce a green streak. But if the mites
streak yellow or orange, they are likely by predatory mites.
15.8 Biological Control
Numerous beetle and thrip species are reported as predators of different spider
mites. Predatory mites (Phytoseiulus spp., Amblyseius spp. or Metaseiulus spp.) are
found under natural conditions. But they can also be purchased from reliable suppli-
ers and suitably released onto mite-infested plants. If the predators are used, avoid
application of pesticides that will kill them. Amblyseius spp. are active from 8 °C to
35 °C and can also survive on pollen. But to be effective, Phytoseiulus spp. require
a relative humidity greater than 65 % and temperatures above 20 °C (Brandenburg
and Kennedy 1987; Hokanen and Lynch 2003).
15.10 Chemical Control 323
Feltiella acarisuga (gall midges) are active predators of mites and can safely be
combined with predatory mites. The organic farmer should strive to conserve the
many naturally occurring beneficial organisms that are present in the crop. If one
finds it necessary to treat for a particular pest in order to prevent economic losses,
choose a selective spray material and use only enough of it to keep damage levels
within economically acceptable limits. Short residual pesticides will allow natural
enemies of spider mites to return to the field and crop control remaining on newly
introduced pest species.
There are many opportunities for enhancing biological control factors in organic
farming systems. Since most organic farms apply little or no harsh pesticides, it is
possible to build up large numbers of spider mites, predaceous insects and mites that
help control spider mite pests in many crops. When a grower is making the transi-
tion from conventional to organic production systems, in this period some biocon-
trol organisms can be most cost effective. But it should be sure that the introduced
pest is well adapted to the particular climate or site and that it is the correct species
for the pest that you would wish to control. It is important to buy a predator only
from a reputed supplier. Multiple releases of the biological control agents may be
required in order to bring pest mite populations to very low levels. Viability of bio-
logical control agents is affected through heat, cold, time of releases and diseases.
So pest management plans are site specific. Organic growers should develop their
own strategies based on their knowledge, available time and capital. Farmer can
manage their fields to provide suitable habitats to favour predaceous mites and
insects. This can be accomplished through conserving and augmenting population
of natural enemies of the mite pests.
15.9 Microbiol Control
The most effective natural control of two-spotted spider mite in soybean, corn and
many other cropping systems is the fungal pathogen, Neozygites floridana. This
fungus species attacks all stages of mites and is specific to spider mites. Infected
mites have a waxy or cloudy appearance. This mite killing fungus requires tempera-
tures cooler than 28 °C with 90 % relative humidity to produce infective spores.
Periods of at least 12–24 h of relatively cool, moist and humid conditions are neces-
sary for the fungal pathogen to disperse and infect a spider mite population in a
field. Mites die often 1–3 days of infection, and mite populations can decline quickly
once the pathogen has spread (Oelaf 1978; Zhender et al. 2007).
15.10 Chemical Control
A number of organically acceptable insecticides are available and each may be use-
ful in specific circumstances. If under certain unavoidable circumstances, the use of
organically acceptable pesticides can be initiated after consideration of the follow-
ing factors. The pesticide selected should have low mammalian toxicity as these
may present a health risk to the persons who must work in the crop after application.
Materials such as sulphur control effectively russet mites but can cause skin and eye
irritation during their application and even for sometimes afterwards. Second impor-
tant criteria of selected pesticide should be their minimal effect on natural enemies
of mites. Selected pesticides must be applied in a way to achieve adequate coverage
of the plant with the pesticide. Since most of the damaging spider mites are found
on the underside of the leaf surface, it must have direct contact with the spray mate-
rial for their successful control. In many cases, high volumes of water are necessary
to get the pesticide to the target. Often the best sprayer nozzle configuration will
direct the spray to the underside and interior of the plant. Higher pressure at nozzle
tips will break the water into smaller droplets. Air blast and electrostatic sprayers
can also provide excellent coverage without requiring high volumes of water needed
by conventional spray equipment. Proper timing of the spray to intercept the pest in
a susceptible stage is of great importance. Since most of the organically approved
pesticides degrade rapidly to the environment, so control of the mite pests may
require repeated applications (Brandenburg and Kennedy 1987).
Pesticides like acephate (Orthene) and abamectin (Avid) and horticultural oils
(summer oils) are effectively used in some cases particularly on homegrown plants.
Horticultural oils can be used on perennial and woody ornamentals during summer,
but avoid spraying flowers which can be damaged. But dormant oils are used to kill
mite eggs and dormant adults in the fall and spring. Insecticidal soaps are also effec-
tive against spider mites where webbing prevents penetration problems. Remember
that mites are very tiny, and soaps and oils work by contact only, so thorough cover-
age of the plants’ upper and lower surface is essential for good control. Spiromesifen
(Forbid) is selective against spider mites but conserves their natural enemies and is
reported useful on ornamental plants. Sulphur (different brands) has been found
effective against fungal diseases and some mites on some ornamental plants and
vegetable crops (Oelaf 1978).
15.11 Important Websites
• IFOAM – Organics International and its action group

(http://www.ifoam.bio/http://www.ifoam.org/)
• National Organic Programme (USDA NOP)
http://www.ams.usda.gov/nop/index.II.htm
http://www.ams.usda.gov/about-ams/programs-offices/national-organic-program
• Organic Materials Research Institute
(www.omri.org)
• Sustainable Agriculture Research and Education
(www.sare.org)
• ATTRA NCAT Sustainable Agriculture
(www.attra.ncat.org)
• Resource Guide for Organic Insect and Disease Movement
(http://www.nysaes.cornell.edu/pp/resource.guide)
• Biointensive Integrated Pest Management (IPM)

(https://attra.ncat.org/attra-pub/summaries/summary.php?pub=146)
• Ontario Ministry of Agriculture and Rural Affairs
(http://www.omafra.gov.on.ca/english/crops/facts/09-077.htm)
15.12 Conclusions
The knowledge obtained from biological, ecological and behavioural information

on different mite species on organic crops can be used to plan their management.
Hence mite pests under organic cultivation of crops may be controlled through cul-
tural, mechanical or physical methods, augmentation or introduction of predaceous
insects and predatory mites for control of pest mites, development of habitat for
natural enemies of mite pests and utilization of some non-synthetic controls.
Whenever these practices are insufficient to prevent or control mite pests, then bio-
logical, botanical or chemical materials or substances approved for use in organic
crop production may be applied to prevent, suppress or control mite pests (Willer
and Yussefi 2006). Pest management plans are site specific, and organic farmers
should develop their own strategies based on their knowledge, available time and
capital resources they can devote to pest management.
Cultural practices or specific crop production practices are implemented in the
initial stages of a long-term organic farm plan to reduce the likelihood of pest infes-
tation and damage. Making the crop unavailable to pests in space and time through
knowledge of pest biology (through crop rotations, manipulating timing of planting
or harvest, etc.), making the crop unacceptable to mite pests (through intercrop-
ping), reducing pest survival on crop by enhancing natural enemies (by increasing
crop ecosystem diversity and through habitat manipulation) and altering crops’ sus-
ceptibility to mite pests (by breeding mite-resistant or mite-tolerant cultivars and by
enhancing soil quality and fertility) are first-phase strategies to control mites. But
second-phase strategies for management of mites include conservation, biological
control and intercropping can be implemented after the strategies under first phase
have been employed. The inundative and inoculative releases as the role of biocon-
trol agents are the third-phase strategies for mite management on organic crops. But
using of approved pesticides of biological and mineral origin is the fourth-phase
strategy which can only be considered if damage by mite pests is unbearable on
high-value crops like vegetables and fruits. If vegetation is not managed to enhance
natural enemy impact, mite pest management will depend on input substitution
using third- and fourth-phase strategies. For an organic farmer, however, being
within a landscape dominated by conventional farms with a high pesticide input
may be disadvantageous. At present the researches available on management of
mite pests on organic systems are a few, but useful findings reported in integrated
mite management in different ecosystems must be considered for developing strate-
gies on organic crops. There is a need to do more research under different ecosys-
tems particularly on high-value crops on certified organic land. Future studies of the
effects of organic agriculture on natural enemies and the pest mites should be
encouraged particularly those that cover several spatial scales, including that of the
landscape, rather than confining only to within-field or even paired-field studies.
Further Reading
Bengtron J, Ahnstrom J, Weibull AC (2005) The effects of organic agriculture on biodiversity and
abundance: a meta-analysis. J Appl Ecol 42:261–269
Brandenburg RL, Kennedy GB (1987) Ecological and agricultural considerations in the manage-
ment of two-spotted spider mite, Tetranychus urticae (Koch). Agric Zool Rev 2:185–236
Cook SM, Pickett JA (2007) Pushing insects around: the use of ‘push-pull’ strategies in integrated
pest management. Annu Rev Entomol 52:375–400
Flaherty DI, Wilson LT (1999) Biological control of insects and mites of grapes. In: Bellows TS,
Fisher TW (eds) Handbook of biological control. Academic, New York, pp 853–869
Gurr GM, Wratten SD, Altieri MA (eds) (2004) Ecological engineering for pest management:
advances in habitat manipulation for arthropods. CABI Publishing, Wallingford
Hesler LS, Grigerick MA, Orage MJ, Palrang AT (1993) Arthropod fauna of conventional and
organic rice fields in California. J Econ Entomol 86:149–158
Hokanen HMT, Lynch JML (eds) (2003) Biological control: risks and benefits. Cambridge
University Press, Cambridge, pp 236
Landis DA, Wratten SD, Gurr GM (2000) Habitat management to conserve natural enemies of
arthropod pests in agriculture. Annu Rev Entomol 45:175–201
Oelhaf RC (1978) Organic farming: economic and ecological comparisons with conventional
methods. Allenhold Osmun, Monteclairf
Smith SM (1989) Plant resistance to insects. Wiley, New York, pp 286
Thomas M, Waage J (eds) (1996) Integration of biological control and host-plant resistance breed-
ing. Technical Centre for Agricultural and Rural Cooperation, Wageningen, p 99
Van Lenteren JC (ed) (2003) Quality control and production of biological control agents: theory
and testing procedures. CABI Publishing, Wallingford, p 327
Weibal FP, Haselis A, Schmid O, Willer H (2004) Present status of organic fruit growing in Europe.
Acta Hortic 638:375–385
Willer H, Yussefi M (2006) The world of organic agriculture: statistics and emerging trends 2006.
IFOAM, Bonn, pp 211
Wyss E, Luka H, Pfiffner L,Schlatter C, Ucholinger G, Daniel C (2005) Approaches to pest man-
agement in organic agriculture: a case history in European apple orchards. Organic Research
(May), Brisbane, pp 33–36
Zhender G, Gurr GN, Kuhne S, Wade MR, Wratten SD, Wyss E (2007) Arthropod pest manage-
ment in organic crops. Annu Rev Entomol 52:57–80
1. https://www.youtube.com/watch?v=u_u63n1umZ0. Predatory Mites for Organic Pest Control:

High-Tunnel Cucumbers
2. https://www.youtube.com/watch?v=9gU19BgJMXM. Organic Vegetable Production – Controlling
Spider Mites on Organic Strawberries
3. https://www.youtube.com/watch?v=unkPfLqkxvU. Spider mite control in the garden
4. https://www.youtube.com/watch?v=31ktEhMkzKU. Spider Mites
Mite Transmission of Plant Diseases
16
Abstract
Most plant viruses depend on vectors for their survival and spread. Mite-borne
plant viruses may cause severe or even crippling losses to many annual and
perennial crops in the tropics and semi-tropics which provide ideal conditions for
the perpetuation of viruses and their vectors. Many worm mites (Eriophyidae),
false spider mites (Tenuipalpidae) and spider mites (Tetranychidae) are known as
vectors of virus diseases infecting cereals, fruit trees, pulse crops and coffee
plants. Wheat streak mosaic virus (WSMV), wheat spot mosaic virus (WSPMV),
ryegrass mosaic virus (RgMV), fig mosaic virus, cherry mottle leaf virus, current
reversion disease, pigeon pea sterility mosaic disease and rose rosette diseases
are all vectored by different species of eriophyid mites. Virus diseases like citrus
leprosis and passion fruit green spot virus are vectored by false spider mites.
Barley yellow streak mosaic virus (BaYSMV) is vectored by spider mite,
Petrobia latens. Mango malformation disease (MMD) which was earlier reported
associated with bud mite Aceria mangiferae is now found to be caused by fungal
pathogen, Fusarium moniliforme var. subglutinans. Management of these dis-
eases through genetic host resistance, adoption of cultural practices, chemical
applications and various regulatory measures is suggested.
Keywords
Transmission • Viral diseases • Mango malformation • Monitoring
Most plant viruses depend on vectors for their survival and spread. Most vectors
(insects and mites) have piercing mouthparts that transmit plant viruses in either the
circulative virus (CV) or non-circulative virus (NCV). Non-circulative viruses are
carried on the lining of the cuticle of vector stylets. CVs cross the vectors’ gut, move
internally in the salivary glands (SG) and cross the SG membranes to be ejected
upon feeding. Most plant viruses depend on vectors for their survival because of the
presence of an impermeable cuticle that coats the plant epidermis, preventing entry

DOI 10.1007/978-981-10-1594-6_16
328 16 Mite Transmission of Plant Diseases
of virus particles. Many virus diseases may spread by contact or by vegetative

reproduction. Hemipterous insects (including aphids and leafhoppers) are well
adapted to their role as vectors by their capacity to pierce the epidermis and deli-
cately deposit viruses in the cytoplasm without risking the integrity of the plant cell.
Moreover, plants are rooted and lack independent mobility; therefore, many viruses
depend on insects and mites for transport among hosts (Jeppson et al. 1975).
Progress in molecular biology of viruses and their vectors has assisted in identi-
fying motifs in the viral genome, and in viral and vector proteins, thus adding to the
understanding of the process of virus transmission of insects and mites. Depending
on their period of retention of virus in their vectors, viruses are known as persistent,
semi-persistent and non-persistent. Non-persistent viruses are generally known as
stylet-borne, while persistent viruses are referred to as circulative. Non-persistent
viruses are acquired and inoculated during brief probing times and do not require a
latent period in the vector (as in aphids). Semi-persistent viruses need longer peri-
ods (hours) for acquisition and transmission than do non-persistent viruses; semi-
persistent viruses have a narrow range of vector species. However, they too need no
latent period and are lost when the vector moults. In persistent viruses, the longer
the acquisition and inoculation times, the higher is the rate of transmission. They
mostly have a narrow range of vectors, pass through moult and need a latent period.
Mite-borne plant viruses may cause severe or even crippling losses to many
annual and perennial crops in the tropics and semi-tropics which provide ideal con-
ditions for the perpetuation for viruses and their vectors. On occasions, mites are
responsible for transition from a non-spreading form to the epidemic form of dis-
eases. Many worm mites (Eriophyidae), false spider mites (Tenuipalpidae) and spi-
der mites (Tetranychidae) – all possessing piercing and sucking mouthparts – are
known as vectors of virus diseases infecting cereals, fruit trees, pigeon pea and
coffee plants (Jeppson et al. 1975; Slykhuis 1969). The plant diseases both viral and
fungal for which mites act as vectors are discussed below briefly.
16.1 Plant Viral Diseases Vectored by Eriophyid Mites
Eriophyid mites are very small (0.2 mm or 200 μm), four legged and worm type
having piercing–sucking mouthparts. They usually feed on young tissues in buds or
on leaves of their specific host plants. Some such mites cause leaf discolouration,
malformation, galls, bud blasting, swelling and other varied symptoms because of
their feeding, and these symptoms are often confused with symptoms caused by
some viruses. Manual transmission, graft transmission, tests with non-viruliferous
mites and the continued development of symptoms after the elimination of mites are
commonly used as tests for establishing the role of mite vectors in transmission of
different virus diseases.
16.1 Plant Viral Diseases Vectored by Eriophyid Mites 329
16.1.1 Wheat Streak Mosaic Virus (WSMV)
Wheat streak mosaic virus (WSMV) is distributed widely in Australia, Canada,

Mexico, Russia, the USA and some European countries. WSMV is a seed and mite-
borne virus that infects wheat causing severe leaf symptoms and reduced yields
(ER1 and ER2). The wheat curl mite (WCM), Aceria tosichella, vectors this virus.
It completes its life cycle in 8–10 days and under ideal conditions a single female
lays 12–20 eggs. Peak mite population is found at approximately 25 °C but its
development is reduced at 30 °C and below 15 °C. WCM is capable of surviving for
a few months in cold conditions but reproduction is significantly lowered (Slykhuis
1967). WCM reproduction is greatest following periods of rain and warm tempera-
tures (25–28 °C).
WCM infestation generally causes little direct damage in the field other than the
characteristic leaf curling and the occasional trapping of the flag leaf. However,
both nymphs and adults of WCM transmit wheat streak mosaic virus (WSMV). For
a mite to become viruliferous, the virus must be acquired during the two nymphal
stages, typically after at least 15–30 min of feeding on infected plant material. Once
infected the mite has the potential to transmit the virus to noninfected plants for at
least 7 days.
Wheat is the primary host of WCM, but the mite has been recorded on more than
60 other plant species (including barley, oats, annual grass and many graminaceous
weed plants). Infected volunteer wheat and alternate host plants provide both an
effective ‘green bridge’ refuge for WCM outside the growing periods of wheat and
serve as a potential sources of reinfestation and spread of WSMV to the successive
wheat crops. WCM do not colonize on plants having broad leaves. WCM is mainly
dispersed through wind. So WCM survival and dispersal are key factors influencing
the spread of WSMV. In addition to WSMV, WCM has been implicated as the vec-
tor (carrier) of at least five viruses, namely, wheat spot mosaic virus, wheat spot
chlorosis pathogen, cereal spotting, triticum mosaic virus and high plain virus. The
same mite in earlier literature has been referred to as Aceria tulipae, which is actu-
ally more commonly found on liliaceous plants.
Early infection of wheat streak mosaic virus (WSMV) is responsible for the
greatest loss of wheat yield. Plants infected between tillering and first node often do
not set any seed, while those infected between first node and booting typically have
reduced seed size. Infected seedlings often die prematurely, while later infection
causes progressively less damage with only slight losses expected when infection
occurs in spring growing periods. Serious outbreaks of WSMV can only occur if the
virus is present and WCM is abundant.
Plants with low levels of infection may show minor or no visual symptoms. But
plants infected with WSMV are typically characterized by leaf mottling and streak-
ing. Light green streaks running parallel to the leaf veins are the first signs of WSMV
infection. As the disease becomes more established, the streaks turn yellow and
develop into blotches, giving the leaf a green and yellow mosaiclike pattern. Tillers
on affected plants tend to be less erect than those on uninfected plants. Affected
plants can die prematurely or fail to grow, becoming stunted relative to healthy
plants. Heads on infected plants can be sterile and contain no seed or can have small
or shrivelled grains. Affected plants often occur in patches or at crop boundaries clos-
est to grasses that were growing when the crop emerged. Stunting symptoms are
much less obvious with late infection. Symptoms of WSMV typically develop at
temperatures above 10 °C, so they are masked during winter. WSMV can cause crop
failure in wheat when widespread infection occurs at the seedling growth stage.
WSMV is seed-borne at low levels (less than 1 %) in wheat seed. Transmission
through the seed has the potential to cause significant yield losses because of early
appearance of WSMV in wheat crop. Early widespread infection of young wheat
plants (approaching 100 % infection) is generally associated with greatest yield
losses from WSMV and can cause complete crop failure, as such crops produce
only small amounts of shrivelled grains. However, minimal yield loss occurs when
wheat crop becomes infected post-tillering.
Management Serious outbreaks of WSMV can only occur if the mite vector
(WCM) is abundant and a source of WSMV is present. So the management of the
disease is highly dependent on controlling WCM populations and sources of
WSMV. Following control options against WSMV should be followed:
• Control the ‘green bridge’ (volunteer crop cereals, e.g. wheat, barley, cereal rye,
oats and grassy weeds) as these hosts harbour both WSMV and WCM. So their
control needs to be done throughout the paddock (including along the fence line)
at least one month before sowing wheat.
• Sow healthy seed stocks of wheat.
• Avoid early sowing in virus risk conditions.
• Break the disease cycle by controlling over-summering volunteer wheat/cereals
and grasses within the field, on field boundaries and road sides, reducing the mite
populations invading autumn sown crops.
• Whenever it is possible, delay sowing of wheat in autumn until temperatures are
too low for mite activity.
Acaricidal control of mites is believed to be largely ineffective as WCM pre-

dominantly lives protected within the leaf whorls and is difficult to target and does
not respond well to the pesticides. Moreover, WCM lacks a diapause, an over-
summering egg stage that many mite species have. Eggs must die off or hatch to
continue development on alternate host plants and volunteer wheat. Even the herbi-
cides like glyphosate require up to 4 weeks to achieve full plant mortality, and in the
meantime dispersal of WCM through wind may occur on a new host.
16.1.2 Wheat Spot Mosaic Virus (WSpMV)
Wheat spot mosaic virus (WSpMV) is vectored by Aceria tosichella – an eriophyid

mite species, also found vectoring WSMV. Wheat spot mosaic virus is acquired by
nymphs as well as adults. In some cases, wheat fields are found simultaneously
having both WSMV and WSpMV diseases. Wheat spot mosaic virus disease causes
a very distinct mottling and spotting on many corn varieties and on barley, oats, rye
and a number of annual grasses. This virus disease is also reportedly vectored by an
eriophyid mite, Aceria tulipae, which is mainly found on liliaceous plants. At early
stages of infection, toxic effects of mite feeding are also attributed as cause, but in
WSpMV, symptoms continued to develop even after the plants were freed of mites.
The disease is not spread transovarially (Jeppson et al. 1975).
Virus-caused symptoms varied greatly in severity. In some cases extreme chloro-
sis and death of the emerging plants are reported. Plants infected simultaneously
with both WSMV and WSpMV were more severely diseased than plants infected
with either virus alone. Some of the mites from plants infected with both WSMV
and WSpMV carried both the viruses, but a higher percentage of the mites transmit-
ted WSpMV than WSMV.
16.1.3 Ryegrass Mosaic Virus (RgMV)
Ryegrass mosaic virus (RgMV) is a member of the potyvirus of plant viruses.

Ryegrasses are the most important hosts but other grasses such as cocksfoot
(Dactylis glomerata), fescue and annual meadow grass (Poa annua) can be infected.
RgMV has a worldwide distribution and is considered to be the most serious and
widespread sap-transmissible virus infecting Gramineae in Britain, Australia and
New Zealand. RgMV causes light green to yellow flecking and streaking of ryegrass
leaves which is followed by necrosis and death of the tiller Italian ryegrass (Lolium
multiflorum), some perennial grasses, rice and oats, but not on wheat and other hosts
of WSMV. Eriophyid mite, Abacarus hystrix, transmits RgMV. When reared on
diseased ryegrass, all stages of the mite except the eggs carried the virus. Non-
infective mites acquired the virus within 2–12 h of feeding on a diseased plant. In
the absence of a virus source, viruliferous mites lost infectivity in less than 24 h of
leaving infected leaves. All instars of A. hystrix transmitted the virus (Malligan
1960).
16.1.4 Agropyron Mosaic Virus (AMV)
Agropyron mosaic virus (AMV) has been found on Agropyron repens in some parts
of the USA and Canada. Eriophyid mite, Abacarus hystrix, has been reported as
vector of AMV in some parts of Canada and causes mosaic symptoms similar to
wheat streak mosaic virus (WSMV) disease (Slykhuis 1969). The virus is readily
sap transmissible and has particles similar to WSMV but is only distantly related to
WSMV serologically. AMV spreads from A. repens to wheat during the warm sum-
mer weather in Ontario region of Canada. Virus transmission by eriophyid mites
Aceria tulipae, Abacarus hystrix and Vasates mckenzie on Agropyron repens has not
been demonstrated.
16.1.5 Fig Mosaic Virus (FMV)
Eriophyid mite, Aceria ficus, has been reported as a vector of fig mosaic. Disease
symptoms include mottling, spotting and distortion of youngest leaves and later
vein clearing and a systematic mosaic of varying intensities. Sometimes disease-
infected leaves and fruits may even drop (ER4). Ten or more days are required for
appearance of disease. The fig mosaic virus is graft transmissible but not sap trans-
missible. FMV disease causes a serious problem for the fig growers in Egypt and
many other places. Biological and molecular characterization of FMV has been
reported recently from Egypt (Haffez et al. 2011).
Eriophyid mite, Aceria ficus, lives in buds during the dormant season. In spring,
mites spread to developing leaves, laying eggs among the thick mat of trichomes on
the lower leaf surface. Mites infest fruits, leaves, buds and young green twigs. Fig
mosaic virus has been reported from India, Germany and the USA. Transmission
occurred after access to an infected plant for only 5 min. Both nymphal and adult
mites are able to acquire the pathogen and transmission can occur within a few
hours after acquisition. Mites retain the ability to transmit after moulting and adults
remain inoculative 6–10 days after removal from infected plants, but the pathogen
is not transmitted transovarially. For management of FMV disease, never plant fig
cultivars that are propagated from mosaic infected trees. By controlling fig mites,
disease incidence can also be reduced.
16.1.6 Peach Mosaic
Virus disease is transmitted by an eriophyid mite vector, Eriophyes insidiosus.

Peach mosaic-affected trees have leaves which are often small, narrow, crinkled and
irregular in shape; leaves of affected trees have mosaic pattern. Internodes of
affected trees become shortened and a profuse growth of leaf axil buds is found.
Colour breaking of blossoms and rosetting of branches of peach and certain other
Prunus (plums, apricots and almond) host plants. and peach mosaic virus disease
have been reported from south-western USA and Mexico. Mites are found more
commonly beneath bud scales. Regular nursery inspection, removal of infested trees
and adopting different quarantine measures help in limiting the spread of the mite
vector of disease.
16.1.7 Cherry Mottle Leaf Virus (CMLV)
Cherry mottle leaf virus (CMLV) disease is transmitted by an eriophyid mite vector,
Eriophyes inaequalis. Disease symptoms include irregular chlorotic mottling, edge
tattering and reduction in size of leaves. CMLV has been reported from cherry
growing areas of the USA, Canada, European countries (where cherry grows) and
South Africa. Fruits on infested trees appear normal but they lack flavour and often
ripen late (ER7). Prunus emarginata is the only known wild host. Cherries grown in
the foothill locations where bitter cherry occurs are at greater risk of infection. With
infected wood CMLV is spread by budding and grafting and by the vector mite, E.
inaequalis. Trees on the orchard’s edge usually are infected first. Removal of
infected trees prevents new infections and also destroys stands of wild cherry near
commercial orchards. Virus-free nursery stock and scion should only be used.
16.1.8 Currant Reversion Disease
Blackcurrant reversion is a viral disease commonly found on blackcurrants (Ribes

nigrum) worldwide; it also affects redcurrants (R. rubrum) but not gooseberry (R.
uva-crispa). The disease is particularly widespread in blackcurrants in Eastern and
Central Europe, Russia, in some parts of the UK and Scandinavia and New Zealand.
In nature, the causal agent of the disease is transmitted between plants by the black-
currant gall mite, Cecidophyopsis ribis, but not through the seed. However, the dis-
ease can be transmitted between infectible Ribes plants by grafting. Two forms
(strains) of reversion are distinguished: the common European form (E) and the
more severe R form found in Finland and some other countries of erstwhile Soviet
Union. The two forms differ in the severity of symptoms expressed in blackcurrant
plants but the progression of the disease both in plants and crops is similar with each
of this form. Compared to leaves of healthy plants, the leaves of diseased reverted
plants are narrower, show a decreased number of the main veins and have larger but
fewer marginal serrations. Flower buds of infected plants develop strong malforma-
tions including the absence of stamens, elongation of the style and increase in num-
ber of petals. Affected flower buds are usually sterile causing a severe loss in fruit
productivity. The leaf markings often disappear as the leaves age and are usually not
evident on growth made during the summer (Jeppson et al. 1975; Slykhuis 1969).
For prevention and control of blackcurrant reversion disease, immersion in hot
water (46 °C for 10–20 min) is reported to eradicate the vector (gall mites). Planting
of healthy (certified) plant material away from sources of infection also proves help-
ful in controlling reversion disease. Successful chemical control of eriophyid mites
requires the accurate timing of applications to coincide with mite dispersal. But
blackcurrant reversion disease is the most destructive disease of blackcurrants, and
once established in the crop with its vector, it is very difficult to control.
16.1.9 Pigeon Pea Sterility Mosaic Disease (PPSMV)
PPSMV is characterized by a bushy and pale green appearance of plants, excessive

vegetative growth, stunting, reduction in leaf size, leaf distortion and mosaic of
leaves. Complete or partial cessation of flowering (sterile) occurs. The disease has
been reported vectored by an eriophyid mite, Aceria cajani, from India, Bangladesh,
Sri Lanka, Nepal and Myanmar (ER3). The mite is highly host specific, confined
mainly to pigeon pea and its wild relatives. Mites have a life cycle of about 2 weeks,
are found on the lower surface of leaflets and are found predominantly on
symptomatic leaves of PPSMV-infected plants. Some studies suggested a semi-

persistent mode of transmission of PPSMV by A. cajani. PPSMV causing sterility
mosaic disease (SMD) is considered as ‘green plaque of pigeon pea’. Vector mite,
A. cajani, is one of the major biotic factors, leading to heavy yield losses, and hence
poses a big challenge for pigeon pea production in the Indian subcontinent. Wettable
sulphur is reported to be very effective in controlling vector mites (Kulkarni et al.
2002; Latha and Doraiswamy 2008; Manjunatha et al. 2012).
The disease may be controlled possibly by removing all old plants in the vicinity
in annual crops and by removing all old pigeon pea plants growing as perennials in
the vicinity well before sowing the new crops.
16.1.10 Rose Rosette Disease (RRD)
RRV is a devastating disease and is a threat to virtually all cultivated roses (Rosa
spp.). Losses can occur in home and commercial landscapes. The causal agent of
RRD disease remains unknown. At present, researchers refer to the causal agent as
a virus or virus-like agent. RRD is vectored or transmitted by an eriophyid mite,
Phyllocoptes fructiphilus. Mites feeding on diseased hosts are able to acquire the
disease agent and transfer it to the new plants; RRD can be transferred to healthy
plants by grafting. The symptoms of RRD may include red colourations or mosaic
on leaves and shoots; leaf size reduction, distortion and severe roughening; and even
increased thorniness and reduction in flowering. Eventually death of the entire plant
affected by RRD occurs (ER8).
Mite vector is host specific (like most other eriophyid mites) on members of the
genus Rosa. Commonly the disease symptoms are known as ‘rosette of rose’ in the
USA, and such symptoms are reported from many rose species, cultivars and variet-
ies of wild and cultivated roses. Multiflora roses (Rosa multiflora) originally used as
fence is highly susceptible to RRD. Cultivated roses growing near infected multi-
flora roses have a high risk of infection. There is no cure for RRD once bushes
become infected. So, early detection is essential to prevent the disease to nearby
roses. Always purchase new roses from a reliable nursery after inspecting for RRD;
remove multiflora roses from within 100 metres of cultivated roses whenever pos-
sible. Adopt proper spacing of cultivated roses so that plants do not touch each other
and dispersal of disease vector may occur. Remove infected plants including roots
completely; remove and destroy any regrowth that occurs.
16.2 Plant Viral Diseases Vectored by False Spider Mites
In the mite family Tenuipalpidae, one or more species within the genus Brevipalpus,
Cenopalpus, Dolichotetranychus, Raoiella and Tenuipalpus are sometimes recog-
nized as serious economic pests. However, only three species within genus
Brevipalpus are known to vector one or more cytoplasmic- or nuclear-type plant
viruses (Childers and Derrick 2003; Childers and Rodrigues 2011). A brief account
16.2 Plant Viral Diseases Vectored by False Spider Mites 335
of virus–vector relations, biology of mite, disease symptoms and their management

is discussed below.
16.2.1 Citrus Leprosis
‘Citrus leprosis’ is caused by two viruses, namely, cytoplasmic type (CiLV-C) and
nuclear type (CiLV-N). Tenuipalpid mite, Brevipalpus phoenicis, has been reported
as the vector of citrus leprosis virus disease. Brevipalpus californicus and B. obova-
tus have been reported in some literature associated with citrus leprosis symptoms,
but only B. phoenicis has been proven to be an effective predator. The virus in B.
phoenicis is transmitted transtadially but not transovarially. Citrus leprosis has been
mechanically transmitted from citrus to citrus (Knorr 1968). The disease produces
chlorotic lesions on the fruit and leaves that do not result in systemic infections
(Bastianel et al. 2010). Differences in chlorotic patterns occur in different varieties.
Premature fruit drop, defoliations and death of the twigs can occur with devastating
results. Death of a twig or branch results when they become girdled by individual
lesions (ER5). The disease only spreads when both infected trees and mite vector
are present. If the mite vector is not controlled, CiLV-C can kill a tree within 3 years.
Citrus species especially sweet oranges (Citrus sinensis) and some mandarin
(C. eticulata) varieties are comparatively more infected by the mites which seriously
threatens orange production in prone areas. Citrus leprosis has been reported as an
important disease in Brazil, Argentina, Paraguay, Bolivia and Colombia (in South
America); Panama, Guatemala, Costa Rica and some other parts of Central America;
and North America (Florida, USA) (Childers et al. 2003a, b, c; Jeppson et al. 1975;
Knorr et al 1968).
16.2.2 Passion Fruit Green Spot Virus (PGSV)
Passion fruit green spot virus (PGSV) is a cytoplasmic virus disease. Tenuipalpid
mite, Brevipalpus phoenicis, is the only known vector of PGSV. In case of severe
outbreaks of disease, considerable leaf and fruit drop occurred and high populations
of vector mite are reported on passion fruit (Passiflora edulis). This disease has been
reported from Brazil. PGSV disease-affected mature yellow fruits showed charac-
teristic green spotting; patches of green spotting on the leaves are also found. Most
serious damage resulted from necrotic lesion that girdles the stems and kills the
plants. These spots may be uniformly green with a central necrotic depression. The
virus does not move systemically in the plant which resulted in subsequent death of
the plant and eventually in the destruction of the entire orchard (Kitajima et al.
2003).
16.2.3 Coffee Ring Spot Virus (CoRSV)
From Brazil and Costa Rica, tenuipalpid mite B. phoenicis has been reported as a
vector of coffee ring spot virus. Conspicuous localized ring spot lesions on both
leaves and berries are found. A reduced number of coffee berries are found resulting
in lowered yields. The disease caused premature fruit and leaf drop in affected
plants. CoRSV remains restricted to lesion areas. The spread of CoRSV can affect
the coffee industry throughout Central Africa (Chagas et al. 2003).
16.2.4 Orchid Fleck Virus (OFV)
Tenuipalpid mite, Brevipalpus californicus, is reported as vector of orchid fleck

virus from Australia, Brazil, Germany, Japan, Korea and the USA. Orchid fleck
virus (OFV) is a nuclear type of virus and its actual distribution is likely to be much
higher. OFV produces chlorotic or necrotic spots and rings on the leaf fronds of
many genera of Orchidaceae (Kondo et al. 2003).
16.3 P
lant Viral Diseases Vectored by Spider Mites
(Tetranychidae)
Barley yellow streak mosaic virus (BaYSMV) from Alberta (Canada) and Alaska
(USA) has been reported from dry land barley (Hordeum vulgare). Based on virus
particle shape and size, symptomatology, mechanical transmission, host range and
serology, BaYSMV is associated with the barley disease. BaYSMV disease out-
breaks are associated with recurring drought and are accompanied by infestations of
the brown wheat mite vector, Petrobia latens, which is found under drought condi-
tions. Brown wheat mite has been observed as a vector of BaYSMV. The disease
also reported transferred to Nicotiana benthamiana and Chenopodium quinoa by
the mechanical transmission with sap from diseased barley leaves. Early symptoms
of streak mosaic virus disease resemble the feeding damage of non-viruliferous
mites. Diseased barley and wheat leaves appear light green to pale yellow. As the
disease progresses, the leaves acquire light green to yellow dashes and streaks. In
addition some leaves display a uniquely diagnostic symptom pattern of yellow
banding wherein a wide band parallel to the mid-vein on one half of the leaf is dis-
coloured, whereas the other half is a symptomatic dark green. Most infected plants
exhibit moderate to severe diversing. Brown streaks appear on the leaves; in some
cases the extremely diseased plants die (Robertson and Brumfield 2000; Smidansky
1996).
16.5 Management of Mite-Vectored Plant Viral Diseases 337
16.4 Plant Fungal Diseases Vectored by Eriophyid Mites
16.4.1 Mango Malformation Disease (MMD)
Since long bud mite, Aceria mangiferae, has been reported to be associated with the
mango malformation disease, there has been controversy regarding the role played
by this mite and the fungal pathogen, Fusarium moniliforme var. subglutinans. In
recent studies, F. subglutinans has been recognized as the causal agent of mango
malformation, associated with floral and vegetative malformation of mango, and A.
mangiferae may play a role as carrier of fungal pathogen.
MMD has been reported from Australia, Africa (Egypt, South Africa, Sudan and
Uganda), India, the USA and some other parts of the world where mango is grown.
MMD spreads on a tree very slowly but if left unchecked can reduce mango yields.
Mango bud mite, A. mangiferae, has been shown to spread the disease within a tree
and not between trees. A. mangiferae is found mostly on apical growing buds rather
than on the lateral dormant buds. Within the buds, A. mangiferae are found in
peripheral scales than in the meristematic dome within the apical bud. The upper
and middle canopy of the tree reported are more preferred than the lower canopy by
mites. The mite is reported commonly on buds and inflorescences of the mango.
The main spread of MMD to new areas is by infected pruning equipment or vegeta-
tive planting material (ER6).
16.5 Management of Mite-Vectored Plant Viral Diseases
Once plants are infected, little can be done to free them from the virus. Although the
use of resistant varieties has been found to be the best economical and practical
approach for effective management of viral diseases, an integrated approach is essen-
tial in sustainable agriculture. Development of integrated management requires cor-
rect identification of the causative viruses, because symptoms can be misleading, and
adequate understanding of the ecology of viruses and their vectors. The following
approaches need to be considered for developing such an integrated management:
• Genetic host resistance. Resistant type of plants should be planted whenever they
are available. Recent advances in plant cell molecular biology and virology have
lead to the development of genetically modified plants with superior resistance
to some viruses.
• Cultural practices. Scouting and removal of symptomatic plants or known alter-
native weed or volunteer plants that may serve as a reservoir for a given virus.
Encourage rotation to non-host crops; always use clean or sanitized tools and
equipment at the farms. Some viruses are permanently relatively inactivated by
prolonged exposure of the infected tissue to high temperatures (at 38 °C) for
20–30 days. Heat therapy frees individual plants or cuttings of the virus and thus
the clean tissue is used as a propagative source, allowing large-scale production
of virus-free plants.
• Chemical applications. As such there are no chemical sprays or biological con-

trol approaches to eradicate viruses, although such approaches can be used to
control mite vectors.
• Regulatory measures. Management of mite vector populations in the field can be
difficult or impossible unless coordinated on a regional basis, but may be highly
effective in closed production systems such as greenhouses.
16.6 Conclusions
Mite-borne plant viruses may cause severe losses to many annual and perennial
crops in the tropics and semi-tropics where ideal conditions for perpetuation of both
viruses and their vectors are available. Many worm mites (Eriophyidae), false spider
mites (Tenuipalpidae) and spider mites (Tetranychidae) which possess piercing and
sucking mouthparts are known as vectors of virus diseases of cereals, fruit trees,
pigeon pea and coffee plants. Viral diseases like ‘wheat streak mosaic virus’
(WSMV), ‘wheat spot mosaic virus’ (WSpMV), ‘ryegrass mosaic virus’ (RgMV),
‘fig mosaic virus’ (FMV), ‘currant reversion disease’ (CRV), ‘pigeon pea sterility
mosaic disease’ (PPSMV) and ‘rose rosette disease’(RRD) are all vectored by eri-
ophyid mites and cause significant losses under ideal conditions. But virus diseases
like ‘citrus leprosis’, ‘coffee ring spot virus’ and ‘passion fruit green spot virus’ are
vectored by false spider mite (Brevipalpus phoenicis). Another virus disease known
as ‘barley yellow streak mosaic virus’ (BaYSMV) is reported vectored by spider
mite (Petrobia latens). In addition to virus diseases, a fungal disease, namely,
‘mango malformation disease’ (MMD), is also associated with eriophyid mite
(Aceria mangiferae). Information on damage symptoms and measures to manage
these diseases under field conditions is given under each disease.
Further Reading
Bastianel M, Novelli VM, Kitajima EW, Kubo KS, Bassanezi RB, Machado MA, Freitas-Astua
J (2010) Citrus leprosis: centennial of an unusual mite – virus pathosystem. Plant Dis
94:284–291
Chagas CM, Kitajima EW, Rodrigues JCV (2003) Coffee ring spot virus vectored by Brevipalpus
phoenicis (Acari: Tenuipalpidae) in coffee. Exp Appl Acarol 30:203–213
Childers CC, Derrick KS (2003) Brevipalpus mites as vectors of unassigned rhabdoviruses in vari-
ous crops. Exp Appl Acarol 30:1–3
Childers CC, Rodrigues JCV (2011) An overview of Brevipalpus mites (Acari: Tenuipalpidae) and
the plant viruses they transmit. Zoosymposia 6:180–192
Childers CC, French JV, Rodrigues JCV (2003a) Brevipalpus californicus, B. obovatus, B. phoeni-
cis, and B. lewisi (Acari: Tenuipalpidae): a review of their biology, feeding injury and eco-
nomic importance. Exp Appl Acarol 30:5–28
Childers CC, Rodrigues JCV, Derrik KS, Anchor DS, French JV, Welbourn WC, Ochoa R, Kitajima
EW (2003b) Citrus leprosis and its status in Florida and Texas, past and present. Exp Appl
Acarol 30:181–202
Further Reading 339
Childers CC, Rodrigues JCV, Welbourn WC (2003c) Host plants of Brevipalpus californicus, B.
obovatus, and B. phoenicis (Acari: Tenuipalpidae) and their potential involvement in the spread
of viral diseases vectored by these mites. Exp Appl Acarol 30:29–105
Haffez E, El-Morsi AA, Abdelkhalek AA (2011) Biological and molecular characterization of the
Fig Mosaic Disease. Mol Pathog 2(2). doi: 10.5376/mp. 2011.02.0002
Jeppson LR, Keiffer HH, Baker EW (1975) Mites injurious to economic plants. University of
California Press, Berkeley 614 p
Kitajima EW, Rezende JAM, Rodrigues JCV (2003) Passion fruit green spot virus vectored by
Brevipalpus phoenicis (Acari: Tenuipalpidae) on passion fruit in Brazil. Exp Appl Acarol
30:225–231
Knorr LC (1968) Studies on the aetiology of leprosis in citrus. Proceedings of the Conference on
International Organization Citrus Virology University of Florida Press, Gainesville,
4:332–341
Knorr LC, Denmark HA, Burnett HC (1968) Occurrence of Brevipalpus mites, leprosis and false
leprosis in Florida. Fla Entomol 51:11–17
Kondo H, Maeda T, Tamada T (2003) Orchid fleck virus: Brevipalpus californicus mite transmis-
sion, biological properties and genome structure. Exp Appl Acarol 30:215–223
Kulkarni NK, Kumar PL, Moniyappa V, Teifion A, Reddy DVR (2002) Transmission of pigeon pea
sterility mosaic virus by the eriophyid mite, Aceria cajani (Acari: Arthropoda). Plant Dis
86:1297–1302
Latha TKS, Doraiswamy S (2008) Detection of pigeonpea sterility mosaic virus, the causal agent
of sterility mosaic disease of pigeonpea in viruliferous mite vector by DAS-ELISA and
DIBA. Arch Phytopathol Plant Protect 41:537–541
Malligan TE (1960) The transmission by mites, host range and properties of ryegrass mosaic virus.
Ann Appl Biol 48:575–579
Manjunatha L, Ramappa HK, Mahantesha SRV, Gowda MB, Rajappa PV, Kavitha TR (2012)
Management of sterility mosaic disease (SMD) of pigeon pea. Plant Arch 12:1007–1012 (ISSN
0972-5210)
Robertson NL, Brumfield SK (2000) First report of Barley yellow streak mosaic virus-infected
barley in Alaska. Plant Dis 84:595
Robertson NL, Carroll TW (1988) Virus-like particles and a spider mite intimately associated with
a new disease of barley. Science 240:1188–1190
Slykhuis JT (1967) Virus diseases of cereals. Rev Appl Mycol 46:401–429
Slykhuis JT (1969) Mites as vectors of plant viruses. In: Maramarosh K (ed) Viruses, vectors and
vegetation. Interscience Publishers, New York
Smidansky ED (1996) Factors influencing the outcome of barley yellow streak mosaic virus:
brown wheat mite – barley interactions. Plant Dis 80:186–193
1. https://www.youtube.com/watch?v=uippXe-CCXA. Management of wheat streak mosaic and

similar mite-transmitted diseases
2. https://www.youtube.com/watch?v=J3i5kzP1PkE. Wheat Streak Mosaic Identification and
Management
3. https://www.youtube.com/watch?v=−Zo_RslCW4I. Sterility Mosaic Disease of Pigeonpea
4. https://www.youtube.com/watch?v=0J2DXrzD9ws. Fig Mosaic Virus
5. https://www.youtube.com/watch?v=fAC_nAAwZeM. Citrus Leprosis
6. https://www.daf.qld.gov.au/plants/health-pests-diseases/a-z-significant/mango-malformation-
disease. Mango malformation disease
7. https://www.google.co.in/search?q=Cherry+mottle+leaf+virus+disease&espv=2&biw=1024&
bih=640&tbm=isch & Cherry mottle leaf virus disease
8. https://www.youtube.com/watch?v=H-lRxwLNbzM. Rose Rosette Disease
Mite Pests of Mushrooms
17
Abstract
Mushrooms are highly proteinaceous and are used as food. The white button
mushroom is sold as fresh mushroom or is canned and made into soups, sauces
and other food products. Sciarid flies, phorid flies, pyemotid mites, and nema-
todes are the main pests of mushrooms and do most of the damage. A range of
the mite species affect the mushroom crop; some directly damage the fruiting
bodies while others are of predaceous nature and feeds on other small mites,
some may attach mycelium, fly eggs, nematodes and bacteria. Mycelium eating
mites can cause high yield losses. Mites occur in very low proportions in mush-
room houses where good pasteurized compost is used, but mites are commonly
found in poor hygienic conditions. Tarsonemus myceliophagus, Tyrophagus spp.
and Pygmephorus spp. are most important and are commonly found in different
mushroom houses. Measures for the management of mushroom pests includes
strict hygiene, using pasteurized compost and steaming the rooms in the end of
the cultivation cycle. Restricted use of some recommended compounds at low
dosages can be done for controlling mites in mushroom houses.
Keywords
Mushroom • Compost • Pasteurization • Sanitation
Mushrooms are very nutritious products that can be generated from lignocellulosic
waste materials (wheat straw, rice straw etc.), that are rich in crude fibre and protein.
In fact mushrooms also contain low fat, low calories and good vitamins. In addition,
many mushrooms possess multi-functional medicinal properties. Sciarid flies, pho-
rid flies, pyemotid mites and nematodes are the main pests of the mushrooms and
do most of the damage. A range of the mite species affect the mushroom crop; some
directly damage the fruiting bodies, some may attach the mycelium, and some of the
mites are predaceous in nature and feeds on other small mites, fly eggs, nematodes
and bacteria. Mite damage on the fruiting bodies often shows up as small cavities in

DOI 10.1007/978-981-10-1594-6_17
342 17 Mite Pests of Mushrooms
the stem and cap similar in appearance to bacterial pit disease. Mycelium eating
mites can cause high yield losses; mites are very small and are easily transported on
clothings and tools. Of all the mites that are encountered during the cultivation pro-
cess only a few species are counted as mushroom pests. Some mites feed on fungi
and bacteria that exist in great amounts in decaying organic materials; the others are
predators (Anonymous 2002; Fletcher et al. 1986; Kawaaiyan and Ramaswamy
1980).
Mites occur in very low proportions in mushroom houses where good pasteur-
ized compost is used, improved hygienic conditions followed, and a shorter crop-
ping cycle is taken. But mites are commonly found in poor hygienic conditions, and
when too long cropping period is followed. Mushroom mites, because of their small
size are often noticed when they are present in large numbers. Of the earlier stages
of composting large populations of various mite species may be found in the sub-
strate, hence there is abundant nutrition for practically all species at this stage. But
during the phase I of compost preparation process and its pasteurization, all mites
die. Appearance of mites on the farm is considered as an indication of either poorly
prepared compost, or its inadequate pasteurization. Most mushroom inhabiting
mites are predaceous in nature, feeding mainly on other species of mites as well as
nematodes. Some feed slowly on weed moulds and others on bacteria. But they are
all heat sensitive and are unlikely to survive effective phase II composting. Because
of variations in composting temperatures some mites inevitably survive. Mite spe-
cies which may be found as pests, or may assume the status of a pest in cultivation
of mushrooms are discussed below briefly.
17.1 Pest Mites
17.1.1 Tyrophagus spp.
They are slow moving, translucent, with long hairs on their body. If these mites are
abundantly found in mushroom culture, then they may eat small pits in the caps and
stalks. These pits thereafter may suffer from bacterial decomposition which breaks-
down tissues just below the surface and result in the skin collapsing leaving an open
pit (ER 5). These mites may also feed on mushroom mycelium, and can cause crop
losses when they are found in large numbers. Mites usually gain entry into the com-
post by clinging onto sciarid flies when the mites are in the migratory stage.
Migratory stages are normally produced when mites become over-crowded. These
mites do not pose a problem where efficient composting and peak heating takes
place. Organic debris found accumulating around the farm provides a breeding
ground for these mites. Sometimes Tyroglyphus putrescentiae (Schrank), and
Caloglyphus mycophagus (Megnin) have been reported causing damage to mush-
rooms, Agaricus bisporus. These mites also have a hypopus stage, which is resistant
to adverse conditions (Clancy 1981; Kheradmand et al. 2007; Petrova and Salmane
2000; Thapa and Seth 1982).
17.1 Pest Mites 343
17.1.2 Tarsonemus myceliophagus Hussey
This mite is considered as an important pest of mushrooms. They are so small

(0.2 mm long) and can not be seen with the naked eye. Looks shining light- brown,
and feed on the hyphae of mushrooms and many other competing fungi (olive green
Trichoderma, and Penicillium). Sometimes found in large numbers in the substrate
gnawing on the bases of mushroom stipes. Mites on feeding become rounded and
obtain a reddish-brown shade. In case of severe mite infestations, the whole area of
the mushroom may be detached from the growing surface. The base of the stipe is
damaged and discoloured into red-brown or rust-brown. T. myceliophagus repro-
duces rather slowly. Life cycle is completed in 8 days at 24 °C, and in 12 days at
22 °C. Exposure to 39 °C and above are reported lethal to this mite. These mites are
also suspected to spread the viral diseases of mushrooms. A female usually lays
about 1 egg per day, and female longevity may be about three weeks. Amount of
damage to the growing crop depends on the time and degree of mite infestation. If
large number of mites enter the batch at spawning, the consequences of their activi-
ties can be rather significant (Davis 1938; Hussey and Gurney 1967; Osborned and
Hamilton 1966).
The long-legged mushroom mites (Linopodes spp.) and a closely related species
of Ereynetes are also found on mushrooms. They are small, whitish with extremely
long forelegs (3-times the length of the body) which are waved about like antennae.
These mites move rapidly and hide in compost when disturbed. They are reported to
sever the roots of affected mushrooms, which tend to topple over and cause a red-
dish brown colouration.
17.1.3 Red Pepper Mites (Pygmephorus spp.; Pyemotidae)
Pygmephorus spp. got their common name ‘red pepper mites’, because of their
reddish-brown accumulations moving on the mushroom crops or the casing surface
(ER 1 & ER 6). These mites are not regarded as primary pests, but their presence is
usually an indicator that weed mould Trichoderma (green mould) is present in the
compost. These mites feed on various moulds but not mushrooms, but their pres-
ence indicates that the compost used in mushroom production is unsatisfactory. Red
pepper mites promote the spreading of green moulds not only in the infected rooms,
but also on the whole farm. Their presence in large numbers worsen the mushrooms’
marketable appearance and bring unpleasant sensations. Sometimes mites even
cause allergy to workers handling the mushroom cases. These mites reproduce very
fast, one female can lay up to 160 eggs within 5-days. They are yellowish – brown
in colour and are 0.25 mm in length with a flattened appearance. They are consid-
ered secondary pests. They often swarm on the casing and mushroom surfaces; and
where such swarming occurs the mushrooms are rendered unfit for sale. Red pepper
mites can also spread spores of Trichoderma from bag to bag. Mites can be trans-
ported by flies, air currents, tools, and equipment from one mushroom house to
another mushroom house. Proper preparation and pasteurization of compost usually
prevent the development of large populations of mites as few weed moulds will
grow which eliminates food for the mites. Proper sanitation of tools and equipment
can help reduce their spread (Clift and Teffolon 1981).
Pygmephorid mites namely Brennandania lambi (Krckzal), Pygmephorus sell-
nicki Krczal and P. mesembrinae Canestrini are also sometimes found in mushroom
cultures. Presence of these mites in large numbers is regarded as having nuisance
value only, and are indicators of the presence of weed moulds.
17.1.4 Other Mite Pests of Mushrooms
Tyroglyphus longior Gervais; Rhizoglyphus spp.; T. putrescentiae, Histiogaster

spp., Schwiebea sp.; Luciaphorus auricularia Gao, Zou & Jiang, Brennandania
lambi (Krczal), Pygmephorus spp., and Tarsonemus spp. are other mite species
occasionally found in pest status on mushrooms. Luciaphorus sp. (Family
Pygmephoridae) is sometimes found in severe proportions in mushroom cultivation
and causes severe production losses. Some other uropodid, eupodid, and histiosto-
matid mites may also be found under different systems of cultivation of mushrooms
(Clift and Teffolon 1981) but their role as mushroom pests needs investigations (ER
3 & ER 4).
17.2 Predaceous Mites
Hypoaspis miles Berlese feed on sciarid flies; Stratiolaelaps bituberous feeding on

T. putrescentiae, P. consanguineus feeding on sciarids; Arctoseius cetratus (Selnick),
Gamasus spp., D. fallax feeding on sciarids and nematodes infesting mushrooms,
are commonly reported as predaceous mites in mushroom cultures (Ali et al. 1997;
Enkegaard et al. 1997; Sobanhoa et al. 2009). However, reports about their effec-
tiveness as biocontrol agents of major mite or other insect pests of mushrooms is
lacking. Despite the fact that the predatory mites only benefit the mushrooms, they
cause discomfort for the working personnels because they easily stick to clothes and
hands causing unpleasant sensations. When predatory mites appear, it is advised to
test the whole batch of compost to detect their sources of food and take measures
against the real insect and mite pests of mushrooms. Predatory mites are mainly
carried in the mushroom farms through sciarid flies. These mites do not feed on
healthy mushrooms, but some may feed on damaged mushrooms. Histiostoma sp.
of mites is basically a bacterial feeder and may spread the bacteria on to healthy
mushrooms (ER 2).
17.3 Management of Mushroom Mite Pests
Following general measures to prevent the appearance of all kinds of mite pests dur-
ing mushroom cultivation can be adopted.
• High quality composting and compost pasteurization for the purpose of obtain-
ing selective mediums for mushroom mycelium.
• Maintaining strict hygienic conditions on the farm.
• Thorough cleaning of all machinery, equipment and rooms that were used during
the spawning process and spawn run.
• Maintain an environment that favours mushroom growth over its competitors
(collembolans, sciarid and phorid flies) and mites with proper temperature and
relative humidity.
• Controlling flies and other mite dispersing agents.
• Do not stretch the growing cycle beyond acceptable crop capability. For example
going from 4 to 3 or even 2 flushes can help reduce the insect and mite pests by
adjusting the time they have to develop.
• Steaming the rooms in the end of the cultivation cycle thoroughly including con-
tainers and shelves to exclude any possibility for the mites to survive.
• Using disinfectants to clean all working surfaces and implements.
• Screening of windows, doors, and ventilators with 20 mesh screens will check
the entry of mites from the outside environment.
• Immediate removal of spent compost from the farm.
17.4 Chemical Control of Mites
Several miticides/acaricides have been reported effective against mushroom mites.

Diazinon, dichlorvos, ethion, various pyrethrins, methoprene have been effectively
used both under laboratory and mushroom farms. Even citronella oil reported quite
effective against mites. On edible mushrooms, whatever chemical is used against
mite control its safety for human consumption should be first known. Only the most
appropriate pesticide be evenly used at the correct rate, and at the right time under
the right conditions. As part of an IPM programme, pesticides are used with moni-
toring of pests and at established thresholds. Avoid routine use of pesticides during
times of the year when the pest pressure is low during winter and early spring. Even
consider the effects on beneficial organisms in the timing of pesticide applications.
Compare the economic benefit expected from the pesticide application with costs of
control. Make sure the crop and pest are listed on the pesticide label of each pesti-
cide being used. Always adopt alternate use of pesticides to help prevent pest
resistance.
17.5 Conclusions
Edible mushrooms are highly nutritious, rich in crude fibre and protein. A range of
mite species affect the mushroom crop. Some mite species feed on fruiting bodies,
others feed on mycelium. A few mite species found on mushroom are predaceous in
nature and feed on small mites, insects, and nematodes. Their infestation sometimes
result in great yield reduction. Sometimes severe infestation of Tyrophagous
putrescentiae and Tarsonemus myceliophagus, directly or indirectly result in drastic

decline in mushroom production. Red pepper mites (Pygmephorus spp.) which are
not primary pests of mushrooms but their presence is an indicator that weed mould,
Trichoderma (green mould) is present in the compost. Appearance of mites is con-
sidered that the compost used in mushroom production is unsatisfactory, and mites
promote the spreading of green moulds in the mushroom farm. For controlling
mites, several acaricides like diazinon, dichlorvos, ethion, pyrethrins, methoprene,
and citronella oil have been reported effective. However, their safety for human
consumption should be known, and only the correct dose at appropriate time should
be applied. Avoid routine application of pesticides as many times pest presence is
low. Always compare the economic benefits expected from the pesticide application
with the costs of control. Maintain strict hygiene at the mushroom farms.
Pasteurization of composting material used for mushroom production is a big must.
Further Reading
Ali O, Dum R, Brennan P (1997) Biological control of the sciarid fly, Lycoriella solani, by the
predatory mites (Acari: Lalaepidae) in mushroom crops. Syst Appl Acarol 2:71–80
Anonymous (2002) Pennsylvania mushroom integrated pest management. Pennsylvania State
University in collaboration with US Department of Agriculture, University Park
Clancy G (1981) Observations on mites associated with low yielding crops of cultivated Agaricus
bisporus in Australia. Mushroom Sci 11:233–244
Clift AD, Teffolon RS (1981) Biology, fungal host preferences and economic significance of two
pygmephorid mites (Acarina: Pygmephoridae) in cultivated mushrooms. Mushroom Sci
11:245–253
Davis AC (1938) Tarsonemus spp. attacking mushrooms. J Econ Entomol 31:51
Davis AC (1944) The mushroom mite [Tyrophagus lintneri (Osborn)] as a pest of cultivated mush-
rooms, Technical. bulletin no. 879. ARS, U.S. Department of Agriculture, Washington
Enkegaard A, Sardar MA, Rodgaard HF (1997) The predatory mite, Hypoaspis miles: biological
and demographic characteristics on two prey species, the mushroom sciarid fly (Lycoriella
solani), and the mould mite, Tyrophagus putrescentiae. Entomol Exp Appl 82:130–146
Fletcher JT, White PW, Gaze RH (1986) Mushrooms (pest and disease cotrol). Intercept Publishers
Ltd., Newcastle, 122p
Hussey NW, Gurney B (1967) Bionomics and control of Tarsonemus myceliophagus in mushroom
compost. Entomol Exp Appl 10:287–294
Kawaaiyan S, Ramaswamy K (1980) A handbook of edible mushroom. Today and Tomorrow
Printers and Publishers, New Delhi, 123 p
Kheradmand K, Kamali K, Fathipour Y, Ueckerman E, Goltapeh EM (2007) Mite fauna associated
with button mushroom Agaricus bisporus in Karaj Region of Iran. Acta Entomol Sin
50:416–422
Osborne P, Hamilton GA (1966) Control of tarsonemid mite in mushroom sheds. J Stored Prod Res
2:140–141
Petrova V, Salmane I (2000) Some mite (Acari) species from mass rearing laboratories of com-
mercial mushrooms and beneficial arthropods in Latvia. Ekologia (Bratislava) 19:211–212
Sobanhoa S, Bussaman P, Chandrapatya A (2009) Efficacy of Xenorhabdus sp. (X1) as biocontrol
for controlling mushroom mites (Luciaphorus sp.). As J Food Ag-Ind 2009(Special Issue):S
145–S 154
Thapa CD, Seth PK (1982) Mushroom mites and their control. Indian J Mushrooms 8:45–52
Further Reading 347
Electron References (ER)
1. https://www.youtube.com/watch?v=bavqEcofOK0. Cheese mites and maggots

2. https://www.youtube.com/watch?v=hyP6czmysXI. Cannibal mites (Cheese mites)
3. https://www.youtube.com/watch?v=134aMOQwyhY. Cheese mites (Acarus siro) forming the
rind of a mimolette cheese
4. http://www.lumerias.com/video/acarus-siro-5-28548593?return_url=%2Fsearch%2Fuser%2FD
UWaj1xtwl%235-2854. Acarus siro Origine: mimolette
5. http://www.dailymotion.com/video/x1054zm_acariens-1_animals?from_related=related.page.
ext.meta2-only.b508f3d. Acariens 1
6. https://www.google.co.in/search?q=mushroom+mites&safe=strict&espv=2&biw=1024&bih=6
73&source=lnms&tbm= http://www.microscopy-uk.org.uk/mag/indexmag.html?http://www.
microscopy-uk.org.uk/mag/artjul00/ofmites.html. Mushroom mites
Mite Problems of Stored Foods
18
Abstract
Mites are common inhabitants of stored products of all kinds including grain and
grain-based commodities at all stages of processing. Some mites cause direct
damage by attacking grain or indirect damage by contaminating the foodstuffs.
Food contamination can make commodities unpalatable to livestock and even
unacceptable as human food. Practically all dried food products commonly
found in the home are susceptible, including bird feed and dry pet foods. More
than 112 different species of mites have been reported in storages of grains, flour
and other products. Important mite species found on different stored foods men-
tioning their appearance, ecological conditions preferred, foods damaged and
their development are discussed with an aim to use this information for their
early detection and management. Grain mite, cheese mite, dried fruit mites and
bulb mites are commonly found in storages. Grain itch mites are a source of
nuisance to people handling stored products causing transitory dermatitis.
Measures to manage mite pests in storages are elaborately discussed.
Keywords
Mite pests • Grains • Stored products • Packaged foods • Pet foods
Mites are common inhabitants of stored products of all kinds including grain and
grain-based commodities at all stages of processing. Some mites cause direct dam-
age by attacking grain or indirect damage by contaminating the foodstuffs. Food
contamination can make commodities unpalatable to livestock and even unaccept-
able as human food. Stored product mites can also be a cause of occupational dis-
eases like dermatitis or allergenic symptoms in workers handling the grains or food
processing industries. Many kinds of cereal products and other foods stored in
kitchen cabinets or elsewhere in the home may become infested with mites (Wrigley
et al. 2004).

DOI 10.1007/978-981-10-1594-6_18
350 18 Mite Problems of Stored Foods
Practically all dried food products commonly found in the home are susceptible,
including bird feed and dry pet foods. They may also be annoying in that they often
leave the infested food and crawl about in the house. A tolerance for the mite pests
or their excrements in the food approaches zero in most cases. The ability to store
food cereals, grains, dried fruit or nuts, a variety of packaged products, etc., in many
forms is a convenience in homes, restaurants, hospitals, warehouses, food process-
ing plants and grocery stores. If these goods are not stored/kept properly taking all
precautions in view, these foods might become contaminated by mites causing seri-
ous economic losses by feeding, by dispersing disease-carrying organisms in food
and by introducing toxins. Feeding of mites also results in lowered nutritional value
or their physical properties. Such food lacks aesthetic appeal and many people will
not think of eating food that contains some foreign object. Even the pets exposed to
stored product mites are at risk of sensitization and allergic reactions. To eliminate
mite infestations, it is necessary to identify the mite pest, and then finding out which
materials are destroyed by the mite, development of mite on different foods and
ecological conditions in which they survive/thrive is a big must to manage them
properly under different situations.
18.1 Stored Product Mites
The presence of mites makes grains and their products unacceptable or unattractive
to humans and livestock. So in a broad sense, mites are polluters of human and ani-
mal foods. Their role in the stored grain ecosystem might be of energy transformers,
grainivores, herbivores, fungivores, predaceous or parasitic role or scavenger–sap-
robe nature. In case of mites infesting stored grains, mites Tyrophagus putrescentiae
and Acarus siro have lifetime energy consumption of 2.5 m and 2.8 J, respectively.
Mites in general are capable of explosive population growth in favourable condi-
tions with an increase of 2500 times on one month as reported in several publica-
tions. More than 112 different species of mites have been reported in storages of
grains, flour and other products. Important mite species found on different foods in
stores are discussed below briefly regarding their appearance, ecological conditions
preferred and foods damaged, and their biology is discussed below with an aim to
use this information for their early detection and management (Griffiths 1960;
Kucerova and Stejskal 2009; Smiley 1991).
18.1.1 Pest Mites
18.1.1.1 Grain Mite (Acarus siro Linnaeus, Acaridae)

Grain mite is also known as cereal mite, flour mite, forage mite and meal mite. It is
earlier known as Tyroglyphus farinae and is also commonly known as flour mite. It
is the most common of all mite species occurring in stored products and is a serious
international pest of stored grain products and other foodstuffs (ER1 and ER2).
Because of its minute size (0.40 mm), slight infestations are often completely
18.1 Stored Product Mites 351
ignored. Mite attacks all types of dry farinaceous produce, grains, hay, cheese, fish-
meal, linseed, etc. (Hughes 1976; Zdarkova 1967).
Damage Causes direct damage by eating the produce; they usually do not pene-
trate bulk flour to more than 5–10 cm. Only the damaged grain is attacked, but once
inside the seed coat, the embryo is first eaten so that germination is impaired and
nutritive value is reduced. Heavily infested produce is tainted by a misty odour and
contamination of food is often the main damage. Flour mites in the flour can cause
‘bakers itch’, a dermal irritation. They are responsible for the mite dust on the flour.
Mimolette cheese in France is particularly ripened through the introduction of

grain mites Acarus siro during the ripening process. The fans of mite cheese are
convinced of its health-promoting effect. The culinary specialty is said to be benefi-
cial for digestive problems and diarrhoea, and its regular consumption presumably
helps you to household dust mite allergies. However, to confirm above it is consid-
ered important to understand what chemical, physical and microbial changes occur
within the cheeses because of mites. It is important to identify the mite species
present on each cheese variety to improve our understanding of their role in creating
the distinctive characteristics that set these cheeses apart from others.
Life History A female mite lays 20–30 eggs in a day onto the stored produce.
Under favourable conditions, a female may lay up to 800 eggs in her lifetime of
30–40 days. The incubation period lasts 3–4 days. About 25 °C temperature and
90 % of relative humidity are quite optimum for the suitable development of these
mites; life cycle under these conditions is completed in 9–11 days. Sometimes the
deutonymph stage is occupied by a hypopus. There are two types of hypopus – the
inert type is adapted for surviving unfavourable environmental conditions (dryness,
poor food etc.). Hypopus has reduced appendages and is capable of only feeble
movement. The active hypopus has well-developed legs and is adapted for dispersal,
either by its own efforts by transportations on the fur of rodents or by attachment to
an insect or an adult mite. Grain mite, A. siro, can be easily cultured and maintained,
so this mite may serve as a valuable model system for different studies under labora-
tory conditions. Molecular techniques (PCR and ELISA) and various chemical and
physical methods are commonly used for their early detection/infestation.
Appearance The adult mites are pear shaped and white with pinkish legs and have
a distinctive engorged basal segment on the foreleg; the body length varies from 0.3
to 0.4 mm and adults look like dust specks to the unaided eye. Since Acarus siro
does not have trachea, so the cuticle must be permeable to gases and to water vapour.
Lethal rate of water loss occurs at relative humidity lower than 60 %. The distribu-
tion of the grain mite in stored grain is conditioned by the relative humidity below
75 %. Fungi not only attract these mites to a food but are themselves good food
sources for them. On unbroken grain, fungi do not serve as any attraction but
weaken the hull permitting access to the germ where the mites can feed and
reproduce.
18.1.1.2 Tyrophagus longior

They are commonly associated with grain, hay and straw, but are also found on
cheese. However, they are not as serious as Acarus siro. T. longior has been impli-
cated in human pulmonary, urinary and intestinal acariasis. These mites are translu-
cent but the legs and chelicerae are more deeply tanned. The males are 0.33–0.53 mm
long, and the females are 0.53–0.67 mm long. In general, the life cycle requires
14–21 days.
18.1.1.3 Mould Mite, Tyrophagus putrescentiae (Schrank)

Mould mite was earlier known as Tyrophagus castellani and is one of the most
destructive pests of stored products. This mite is found most frequently on products
having relatively high fat or protein content such as linseed, dried egg yolk, peanuts,
cheese, ham, banana, spilled wheat, oats, barley and even on flour and mushrooms
(ER3). Grocers’ itch is found commonly in personnels who handled mite-infested
materials. The persons who handled mite-infested copra developed symptoms com-
monly known as ‘copra itch’. This mite has also been reported from uprooted
banana suckers.
Biology Mites are comparatively small; males are 0.28–0.35 mm in length and
females are measured 0.32–0.41 mm in length. This is a more slender mite than
other acarid mites. The life cycle requires 2–3 weeks at 23 °C and 87 % relative
humidity. Temperature thresholds for egg, larva, protonymph and tritonymph were
7.9, 7.4 and 11.1 °C, respectively. Under moist conditions (12–18 %) and warm
summer temperatures, a generation can be completed in 8–21 days. The mould mite
will breed readily above 30 °C. The mould mite is less tolerant to low temperature
and cannot develop below 10 °C; however, in an inactive state, this mite can survive
at 0 °C. At favourable temperatures and 90–100 % relative humidity, a female can
lay an average of 437 eggs.
18.1.1.4 Cheese Mite, Tyrolichus casei (Oudemans)

Cheese mite is commonly found in stored foods, cheese, grains, dried eggs, damp
flour, old honeycombs and insect collections. The body length varies from 0.45 to
0.70 mm. Life cycle (egg to adult) takes 15–18 days at 23 °C and 87 % relative
humidity. Mites appeared creamy white (ER7). A female may lay 20–30 eggs per
day. The total fecundity may be up to 800 eggs. Warm moist conditions are favour-
able for mite development. Mites cannot live under refrigerated conditions. The
cheese mite is also known to cause dermatitis to the workers who handle mite-
infested cheese. They can eat out small holes in cheese and have been cultured for
addition to Altenburger cheese to import a characteristic ‘piquant’ taste. Heavily
infested cheese is found covered with a greyish powder consisting of large numbers
of living and dead mites, cast skins and faeces. Some people consider such type of
cheese as ripe and particularly delectable. Mites prefer old cheese than the fresh.
For many cheeses, the presence of mites is highly undesirable but there are some
cheeses in which culture of cheese mites is artificially introduced. For example,
‘Milbenkase’ cheese in Germany is traditionally ripened by mites. The tradition of
ripening cheese through introduction of mite culture in Germany is for more than
500 years old in the region of Wurchwitz – a village near Leipzig.
18.1.1.5 Dried Fruit Mite (Carpoglyphus lactis Linnaeus)

The mite body is oval, slightly flattened and translucent. The legs and mouthparts
are pinkish. Adults are 0.38–0.40 mm long, and both sides are similar in appear-
ance. A female lays 4–5 eggs per day; eggs are laid attached to the substrate by
means of a stalk. A female can lay 280 eggs; adults live for 20 days. This mite spe-
cies is found principally on the dried fruits, jams, grains, sugar stores and products
containing lactic acid and also reported in honeycombs, fruit drink residue, wine
clinging to pieces of cork or forming a floating scum on the surface, rotting pota-
toes, flour, dried milk powder and caramel used in manufacturing sweets. C. lactis
has been reported to cause ‘dried fruit mite dermatitis’.
18.1.1.6 Thyreophagus entomophagus (Labailbene)

These mites feed on dried fruits, old flour and jams. They have elongated to oval
body and have colourless shiny cuticle and pale brown stumpy legs. Males are 0.30–
0.45 mm in length, while females are 0.45–0.60 mm in length. These mites are hav-
ing fairly slow movement.
18.1.1.7 Glycyphagus destructor (Schrank)

Glycyphagus destructor is also known as house mite. It is a cosmopolitan species,
but mostly found in temperate areas. These mites have been reported feeding on
flour, linseed, wheat, tobacco, hay and straw, sugar and cheese and reported to cause
dermatitis and asthma in man (ER4). They possess many long stiff setae which are
erect. The mite body is pear shaped, being somewhat constricted behind the fourth
pair of legs. Males are 0.35–0.50 mm long, while females are 0.40–0.56 mm long,
legs with needle-like tarsi; these mites have rapid jerky gait. They are considered to
be strictly mycophagous and are often associated with Acarus siro and also attracted
to sugar.
18.1.1.8 Grocers’ itch mite (Glycyphagus domesticus (Schrank))

Grocers’ itch mite is also known as house or furniture mite and is worldwide in
distribution. These mites are commonly found on flour, wheat, hay, linseed, tobacco,
sugar and cheese (ER5 and ER8). Its population may occur in enormous proportions
on rough furniture and on upholstered chairs stuffed with green Algerian fibre. As
its common name suggests, it may often cause ‘grocers’ itch’ in common associa-
tion with other acarid mites. As compared to G. destructor, its body is more rounded
and has uniform unhurried gait as against jerking. This mite bears long legs, males
are 0.32–0.40 mm long and females measure 0.40–0.75 mm.
18.1.1.9 Lardoglyphus konoi (Sasa and Asanuma)

Lardoglyphus konoi feeds only on high-protein diet and is recorded on cured fish
(dried), prawns (dried), molluscs, bones and fish meal, hides and sheep skins.
Hypopus is found associated with dermestid beetle larvae. Mites have glistening
body with truncate posterior margin, are light brown and have fairly rapid and jerky
movements. They are commonly found on cured fish in tropics and subtropics.
Hypopus (deutonymph) is commonly found in these mites and uses insects for dis-
persal purposes. Once the infestation of mite becomes established, its population
tends to increase exponentially. So its early detection in the cured fish is very impor-
tant for its management. Moisture content in equilibrium with 70–80 % relative
humidity is favourable for their development, and if humidities are considerably
reduced and well-dried fish is stored, then the chances of infestation by this mite are
lowered. The presence of salt in salted cured fish reduces the chances of mite
increase.
18.1.1.10 Bulb Mites (Rhizoglyphus echinopus (Fumouse

and Robin))
R. echinopus is a major problem in the storage of flower bulbs of lilies, narcissus,
onions, Amaryllis spp., Canna spp., Dahlia spp. and tulips. They are large, bloated,
smooth mites, with glistening body having reddish-brown legs and having slow
lumbering gait. They are found on bulbs of different plants and on very wet grains,
rotting potatoes, etc., and also found on all types of vegetables, mushrooms, decay-
ing grains and roots of cereal plants (oats, barley, rye and wheat). Sun drying of
stored bulbs for 6–7 h in tropical conditions freed the bulbs of mites.
18.1.1.11 Suidasia medanensi (Oudemans)

Suidasia medanensi is reported to be associated with wheat bran, rice bean, wheat
germ, groundnut and bees’ nests mainly from the United Kingdom, Europe, North
Africa and Southeast Asia. Suidasia spp. are quite small in size having dull frosted
appearance, bearing long tactile setae. Suidasia nesbiti often occur in wheat and in
grain stocks. Suidasia pontifica are also reported on dried and salted fish.
18.1.1.12 Tarsonemus granarius Lindquist

Tiny oval mites, with clear orange or yellow colour, sometimes look transparent. T.
granarius has been found in association with ageing stored wheat, oats, rice and rice
straw and also as pest of bulbs and growing plants (ER8). In laboratory studies, this
mite has been found to play a fungivorous consumer role in stored grain ecosystem.
Both preharvest and postharvest fungi (Alternaria, Aspergillus and Penicillium)
commonly found in stored cereals support populations of T. granarius. Populations
of T. granarius increased most rapidly at 30 °C and 90 % relative humidity. The
fungus Alternaria alternata has been found to be a favourable diet; the rm value of
this mite is 0.162 which is relatively low as compared to A. siro. As the incidence of
mites and fungi is often correlated in deteriorating grain, T. granarius may be used
as one of the biological indicators of microfloral activity and hence of overall qual-
ity of stored grain. Sometimes the mites that occur in the field also move to the store
house along with the harvested material. Sometimes eriophyid mite, Aceria tulipae,
is also found to play a destructive role in garlic when stored. Rhizoglyphus robini
has many times reported to damage onion bulbs.
18.1.2 Predaceous Mites
In addition to the different grain-feeding or mycophagous mites found on different

stored products, some predaceous mites which feed on grain mites or on different
stages of other stored product insects are also found. Important predaceous mites
and their role are discussed below in brief.
18.1.2.1 Cheyletus eruditus (Schrank)

Cheyletus eruditus occurs quite commonly among grain and grain products where it
lives by feeding on other mites chiefly Acarus siro. C. eruditus is white to pale yel-
low with diamond-shaped body; chalky white line runs lengthwise dorsally. They
have very powerful pincer-like palp. Front legs are long and used as feelers. These
mites are commonly found in farm detritus and are mainly predators of acarid mites
in storages. The life history is normal except that reproduction is parthenogenetic.
The younger stages may feed on eggs. Adult mites seize the prey with their power-
ful palps and suck the contents of prey through beak-like mouthparts.
Ecological balance is found between C. eruditus and A. siro and the environ-
ment. A. siro is very dependent on the water content of the food, and its numbers rise
enormously as the water content increases. C. eruditus population increases too
with the rise in population of their prey. However, C. eruditus can thrive at lower
water contents, but A. siro under such conditions can completely suppress the grain
mite. Cheyletus mites tend to be dominant in the summer, but Acarus species are
more dominantly found in the moist, cooler conditions of winter. Under conditions
which are unfavourable to A. siro, C. eruditus may completely suppress grain mites.
Sometimes Cheyletus malaccensis and C. lepidopterorum are also found commonly
associated with tyroglyphid mites.
18.1.2.2 Blattisocius tarsalis (Berlese)

This mite is a common predator of storage product pests in Europe and Russia and
is commonly found in association with Angoumois grain moth, Sitotroga
cerealella.
18.1.2.3 Paragarmonia dentriticus (Berlese)

Paragarmonia dentriticus is found in laboratory cultures of Tyrophagus.
18.1.2.4 Cheletomorpha lepidopterorum (Shaw)

It is reported to be feeding on Tyrophagus putrescentiae on wheat bran and on the
flour.
18.1.2.5 Acaropsis docta Berlese

Another cheyletid mite, Acaropsis docta, which feeds on the eggs of Trogoderma
granarium and Rhyzopertha dominica (important pests of stored cereals), has been
reported commonly in stored grains on laboratory cultures. Eggs of Corcyra cepha-
lonica are also fed upon by these mites. Breeding experiments on laboratory rearing
of A. docta on eggs of T. granarium revealed that 2.4, 4.6, 3.7, 2.6 and 4.4 eggs were
consumed, respectively, by a predatory mite larva, protonymph, normal deutonymph,

heteromorphic deutonymph and an adult female.
18.1.3 Parasitic Mite
18.1.3.1 Parasitic Mite (Acarophenax tribolii Newstead and Duvall)

Females of this mite are round and arched shaped and yellow in colour. Mainly
parasitizes stored grain pest, Tribolium castaneum, particularly in laboratory cul-
tures. Mites are found attached in regions where the cuticle is soft and congregates
beneath the second pair of wings, pierce the cuticle and suck the body contents.
18.1.3.2 Pyemotes ventricosus (Newport)

Pyemotes ventricosus is commonly known as the grain itch mite. This mite parasit-
izes larvae of many pest species and in that sense can be playing a beneficial role
too. But in laboratory rearing of certain pest species like pink bollworm (Pectinophora
gossypiella), they can be a source of nuisance to people who handle infested cul-
tures or foodstuffs for longer periods. It causes an irritating though transitory der-
matitis. The females are physogastric, the opisthosoma becomes enormously
distended with developing eggs and one female might produce more than 200–300
adults. P. ventricosus is known commonly as a parasite on some of the larvae of
Hymenoptera and Coleoptera and on the larvae and pupae of Lepidoptera. This mite
has been reportedly associated with stored grain insect pests like Sitophilus and
Cadra.
18.1.4 Scavengers and Saprobes
A tortoise-like uropodid mite, Leiodynychus krameri (Berlese), is often found in

damp, mouldy sprouting barley, wheat and corn in ships and warehouses and in the
litter of broiler houses. Several species of tydeid and tarsonemid mites are found in
grain debris on granary floors.
18.2 Temperature/Humidity Relations
Several species of mites belonging to families Acaridae and Tyroglyphidae can

assume pest status in case of stored flour, stored grains, packaged bakery, dried
fruits and pet food, depending on the temperature and humidity conditions occur-
ring in different types of storages. For most stored product insects, 25–33 °C is
optimal for growth and reproduction; at 13–25 °C or at 33–35 °C, insects are able to
complete their development and produce offspring. Both at temperatures lower than
13 °C and higher than 35 °C, insects eventually die. Lethal temperatures vary con-
siderably and depend on species, stage of development, acclimations and relative
humidity (Asplay et al. 2007; Sinha 1964).
18.4 Allergy Due to Mites in Storages 357
Mites are the smallest among the different stored product pests and are com-
monly found in grains and other commodities stored at 14–17 % moisture content.
However, because of their microscopic size, mites often go unnoticed at early stages
of infestation. In general mites are cold hardy and feed on broken grains, dockage
and moulds; they are well adapted for infesting stored products. Mould mites only
develop where there is a moisture or high humidity. Moulds infest dry foods (having
around 10 % moisture content) and serve food for mites and readily provide opti-
mum breeding ground for the storage mites.
Acarid mites penetrate into the hard grain and cause internal changes; whole
fragments of food are bitten of by the mouthparts with the help of their chelicerae.
The digestive system is adapted to the assimilation of hard particles with the aid of
characteristic enzymes. Their presence indicates impending seed spoilage and the
need for immediate attention. In contrast glycyphagid mites cannot gnaw through
hard parts of grain and feed on grain dust; only small particles of dry matter can be
taken up by the mouthparts mainly with the help of chelicerae. The digestive system
and enzymes are adapted to assimilating small particles. Their presence indicates
inadequate storage facilities with the possibility of infestation by more injurious
species.
18.3 Detection of Mites
The early detection of mite pests, their monitoring and ultimately their management
are integral features of any successful quality control or hygiene programmes
employed in storage and production premises. A specially designed BT mite trap
has been successfully used in the cereal and allied industry for early detection of
mite infestations. The BT trap also was successful in detection of mites at low levels
of mite infestations of flower bulbs and improves the control efficiency particularly
of lilies, narcissus and tulip bulbs. Many of the species of mites that attack flower
bulbs are also pests of stored cereals. Mites belonging to genera Tyrophagus,
Rhizoglyphus, Blattiscocius and Cheyletus have been detected successfully at early
stages of infestation under different storage systems and stored products (Wakefield
and Dunn 2005). Molecular methods (PCR and ELISA) and chemical and physical
methods (extraction through Berlese–Tullgren funnels, sieving and filth tests) are
the other commonly used approaches for early detection of mite infestation. But BT
traps have been found very successful under different systems of storages (Kucerova
and Stejskal 2009; Thind 2005). However, under experimental conditions, alarm
pheromones, aggregation pheromones and sexual pheromones have been isolated
and used for detection of stored product mites.
18.4 Allergy Due to Mites in Storages
Hypersensitivity due to storage mites is a worldwide problem. In many studies there

have been reports of asthma due to sensitization to the constituents of grain dusts,
grain rust, smuts and arthropods (mainly insects and mites), and fungi are some of
the constituents identified through skin tests as allergens in grain dust. Occupational
allergy and asthma attributed to stored product mites have been reported commonly
in workers who handle grains in elevators, farm grain stores, cereal workers and
stored hay and grains. Lepidoglyphus destructor, Acarus siro, Glycyphagus domes-
ticus, Tyrophagus putrescentiae and Chortoglyphus arcuatus species of mites are
mainly responsible for occupational allergy in storages. Cough, wheeze, asthma and
other types of allergies in grain handlers have been found (Chambers et al. 1999;
Ford and Platts-Mills 1987).
Steps can be taken to inhibit multiplication of mites and reduce exposure of pets
to stored product mites by way of consumption of food containing mites, as pets are
at higher risks of sensitization and allergic reactions to these mites. Storage mite
bodies and mite faeces are a source of potent allergens, and allergic reactions are
associated with the inhalation of the mites in either household or occupational envi-
ronments. The handling of foods in occupational settings has also been associated
on occasion with contact dermatitis. Although it is virtually impossible to totally
eliminate storage mites from the environment, so minimize their exposure to pets
and human beings.
18.5 Management of Mites in Storages
Stored product insects and mites are tiny and difficult to detect in bulk or packaged
food products. Therefore, they can be freely transported from processing plants to
warehouses, grocery stores, restaurants and household and institutional kitchens.
Even under the most carefully controlled conditions, some of these pests probably
will pass from one level of the food handling system to another either in egg, larval
or adult forms. Eradication at any one level is virtually impossible because of the
size and complexity of food distribution industry. Once an infestation occurs in one
commodity, it can quickly spread to others unless appropriate and timely control
measures are taken. Each entity in the complex maze of food distribution, from the
producer onto the consumer, must assume a role in the management of stored prod-
uct insects and mites. The most difficult part of managing stored product mites is
detecting their infestation. Large populations can develop before they are discov-
ered and a considerable damage might have already been done. The stored food may
have an odour, which has been described as minty, sweet or musty when it is infested
with mites. This odour may be the first indication that mites are present. Use a
microscope or hand lens to inspect stored products for mites that may be colourless
or cream coloured (Armitage et al. 1994; Hubert and Pekar 2009; Sinha 1966a, b;
Wilkin and Hope 1973).
18.5.1 Stored Grains
In dry climate, insects are dominant; mites and fungi are relatively less important.
In a subtropical climate, all three types of organisms are equally abundant. So
18.5 Management of Mites in Storages 359
storage of grains depends on the type of grain, harvesting, drying and storing prac-
tices of a country within a climatic region. So store the grains with less than 12 %
moisture content to retard development of mites. Both the qualitative and quantita-
tive nature of microbial infection and insect and mite infestation of stored grains are
profoundly influenced by the temperature and moisture components of the climate
of a country. Bacteria could be the dominant organisms for causing damage in
stored grains in humid and subtropical climates. But farmers and other authorities
in most countries always recommended proper drying of harvested grains either
through sunlight or by artificial heat, so that a moisture content of grains is kept less
than 12 % because at this level least damage is done by insects and fungi (Pulpan
and Verner 1965; Palyvos and Emmanuel 2006; Sinha 1966a, b).
Generally, a hot damp climate in the tropics provides the most favourable condi-
tion for rapid multiplication of the more destructive pests of stored products,
whereas in the temperate regions, a cool dry climate favours slower multiplication
of destructive species. So the atmosphere in the storage conditions/structures is con-
trolled in such a way that it is not conducive for the growth and development of
mites. On cereals pesticides like dichlorvos, fenitrothion, lindane, malathion and
permethrin are reported effective for application in the storages. Phosphine and
methyl bromide are the only grain fumigants in widespread use for fumigation of
grains (ER10).
18.5.2 Bulb Mites
Bulb mites, Rhizoglyphus spp. and Tyrophagus spp., are shiny creamy-white,
bulbous-appearing mites and measure 0.50–1.00 mm. They have four pairs of short
brown legs and look like tiny pearls with legs. They generally occur in clusters
inhabiting damaged areas under the root plate of onion bulbs or garlic cloves (ER9).
They have a widespread host range and feed on many kinds of bulbs, roots and
tubers and can infest bulbs in the storage or in the food. Bulb mites can survive in
the decaying vegetation in the field until it is completely decomposed. Bulb mites
damage bulbs by penetrating the outer layers of the tissue and allowing rotting
organisms to gain entry. Decaying cole crops, especially cauliflower, may harbour
very high bulb mite populations. No specific monitoring methods are available. Use
a microscope to examine fragments of undecayed vegetation in the soil or volunteer
onion or garlic bulbs for the presence of mites. Treatments are generally preventive
and should be considered for fields that are high in vegetative matter or that have
had previous bulb mite problems. No treatment thresholds are available. The fol-
lowing practices may be followed:
• Rapid rotation from one crop to the next fosters survival of mites on the leftover
vegetation in the soil from the previous crop. Decaying cole crops, especially
cauliflower, may harbour very high bulb mite populations. So uproot and destroy
them. Winter may reduce mite levels in the soil.
• Flood irrigation or heavy rains during the winter may reduce mite levels in the
soil.
• Avoid planting successive onion or garlic crops; growers must insist on clean
seed cloves.
• Hot water treatment of seed garlic before planting may reduce mite infestation.
• In field conditions, fallow the fields to allow complete decomposition of organic
matter; this reduces field populations of mites.
• Never dump new replacement foods (foods, flours, grain, etc.) on older unused
food. Avoid prolonged storage.
• Wash food containers frequently in detergent and hot water (54 °C). Dry com-
pletely before reusing them for food storage.
• Mites can be detected by sticking a piece of scotch tape, sticky side up on pack-
ages or on the shelves of storage areas.
• Always shop for dry goods from the store so that there may be high shelf
turnover.
• Store the goods at household level in airtight glass or plastic containers. Glass
containers should have rubber seal.
• Dry the goods in small quantities only if there is doubt of mite infestation.
18.5.3 Stored Packages in Groceries (Thind and Clarke 2001;

Thind and Wallace 1984)
• Carefully inspect all high moisture food and grain products. High moisture foods
purchased in bulk and stored in bags for long periods of time should be checked
regularly. Rotate food materials to remove the older items first.
• Store pet food only in a clean dry area. Never store under damp, poorly ventilated
conditions. If necessary, increase air circulation to reduce relative humidity and
prevent moulds and mildews. Relatively humidity below 55 % and moisture con-
tent below 12 % are the proper storage conditions. Ventilate and dry areas with a
dehumidifier or fan or by simply opening doors of a damp room.
• Periodically clean the storage areas especially cracks, crevices, shelving, etc.
Vacuum and wipe up any spilled foods, eliminating the foci of infestations by
cleaning with attention to horizontal surfaces such as beams and window ledges.
• Empty all vacuumed contents in a plastic bag and make sure it is discarded
outdoors.
• Locate the source of infestation in the stored product and eliminate it.
• To eliminate source of mite infestation, homeowners should supercool the prod-
ucts at −18 °C for 7 days in a deep freezer or superheated at 54 °C for 30 min in
an oven in shallow pans or 5 min in a microwave or disposed off in several heav-
ily wrapped paper bags for garbage disposal.
• Homeowners should remove all remaining food from the storage area. Place
uninfested food in plastic bags and secure them. Clean the floor, shelves, walls,
floors, etc., thoroughly with hot water or strong detergent, and store again only
after complete drying of floors.
Further Reading 361
18.6 Biological Control
There are some reports from laboratory experiments that through introduction of
predatory mites, Cheyletus eruditus, C. malaccensis, Acaropsis docta, Androlaelaps
casalis, Blattiscocius sp. and Pyemotes tritici, stored grain mites can be success-
fully controlled. But no solid evidence is found on utilization of these predatory
mites in controlling grain mites under any system of storage. But predatory mites,
which feed and develop on all stages of R. echinopus, are reported to suppress bulb
mite populations on lilies in laboratory conditions.
18.7 Conclusions
Already a lot of work has been done on the taxonomy, life history, physical limits,
feeding patterns and the community ecology of the major acarine pests of stored
grains, but there is still a gap in our knowledge of the trophic dynamic ecology and
energy budgeting of mite-infested stored grain ecosystems. The role mites play in
such ecosystems can be properly understood when system analysis can be made
with reliable quantitative information relating to involved species, on intrinsic rate
of increase and physical limits, energy budgets, ecological and physiological sur-
vival mechanisms such as hypopus formation and critical equilibrium humidity.
Studies on adaptability to grain quality and mass size, granary structure and chang-
ing environment that include both nourishing and pathogenic species of microflora,
common in the movement of grain and grain products, are needed. More research is
needed in all the areas listed above for the so-called less important species and as
well as for the major pest species. Still more work is needed on economic losses
caused in different regions of the world in granaries, warehouses and flour mills.
Further Reading
Armitage DM, Cogan PM, Wilkin DR (1994) Integrated pest management in stored grain: combin-
ing surface insecticide treatment with aeration. J Stored Prod Res 30:303–319
Asplay G, Steijskal VJ, Perkar S, Hubert J (2007) Temperature dependent population growth of
three species of stored product mites (Acari: Acaridida). Exp Appl Acarol 42:37–46
Chambers J, Thind BB, Dunn JA, Pearson DJ (1999) The importance of storage mite allergens in
occupational and domestic environments. In: Robinson WH, Rettich P, Rambo GW (eds)
Proceedings 3rd international conference on Urban Pests. pp 559–569. www.icup.org.uk/
reports/%5CICUP472.pdf
Diaz A, Okabe K, Eckenrode CJ, Villani MG, Oconnor BM (2002) Biology, ecology and manage-
ment of the bulb mites of the genus Rhizoglyphus (Acari: Acaridae). Exp Appl Acarol
24:85–113
Ford AW, Platts-Mills TAE (1987) Standardized extracts, dust mites and other arthropods (inhal-
ents). Clin Rev Allergy 5:49–73
Griffiths DS (1960) Some field inhabitants of mites of stored food products. Ann Appl Biol
48:134–144
Hubert J, Pekar S (2009) Combination of the antifeedant bean flour and the predator, Cheyletus
malaccensis suppresses storage mites under laboratory conditions. Biocontrol 54:403–410
Hughes AM (1976) The mites of stored food and houses, Technical Bulletin 9. Ministry of
Agriculture, Fisheries and Food. Her Majesty’s Stationery Office, London, 400 pp
Kucerova Z, Stejskal V (2009) Morphological diagnosis of the eggs of stored product mites. Exp
Palyvos NE, Emmanuel NG (2006) Seasonal abundance and vertical distribution of mites in flat
storage containing wheat. Phytoparasitica 34:25–36
Pulpan J, Verner PH (1965) Control of Tyroglyphid mite in stored grain by the predatory mite,
Cheyletus eruditus. Can J Zool 43:417–432
Sinha RN (1964) The effect of low temperature on survival of some stored product mites.
Acarologia 6:336–341
Sinha RN (1966a) Feeding and reproduction of some stored product mites on seed-borne fungi.
Sinha RN (1966b) Association of granary mites and seed-borne fungi in stored grain and in out-
door and indoor habitats. Ann Entomol Soc Am 59:1170–1181
Sinha RN (1973) Ecology and storage. Ann Technol Agric 22:351–369
Smiley RL (1991) Mites (Acari). In: Gorham JR (ed) Insects and mite prests in food: an illustrated
key vol. I, Agriculture handbook no. 655. United States Department of Agriculture, Washington,
DC, pp 3–44
Thind BB (2005) A new versatile and robust mite trap for detection and monitoring of storage
mites in the cereal and allied industries. Exp Appl Acarol 35:1–15
Thind BB, Clarke PG (2001) The occurrence of mites in cereal-based foods destined for human
consumption and possible consequences of infestation. Exp Appl Acarol 25:203–215
Thind BB, Wallace DJ (1984) Modified floatation technique for quantitative determination of mite
populations in feedstuffs. J Assoc Off Anal Chem 67:86–88
Wakefield ME, Dunn JA (2005) Effectiveness of the BT mite trap for detecting the storage mite
pests, Acarus siro, Lepidoglyphus destructor and Tyrophagus longior. Exp Appl Acarol
35:17–28
Wilkin DR, Hope JA (1973) Evaluation of pesticides against stored product mites. J Stored Prod
Res 8:323–327
Wrigley CW, Clark H, Walker CE (2004) Encyclopedia of grain science, vol 1. Elsevier (Academic
Press), Amsterdam
Zdarkova E (1967) Stored food mites in Czechoslovakia. J Stored Prod Res 3:155–171
1. https://www.youtube.com/watch?v=d0CcERpbcqs Under Microscope: Story on Flour

Mite Acarus siro (Acarina, Acaridae)
2. https://www.youtube.com/watch?v=R7llz-oOUxc Terrible Pest Flour Mite Acarus siro
(Acarina, Acaridae)
3. https://www.youtube.com/watch?v=ol70OSCOmbI Mould mites (Tyrophagus putrescentiae)
4. https://www.youtube.com/watch?v=5miHbbzKMm0 Glycyphagus domesticus
5. https://www.youtube.com/watch?v=_h8cr6cgjAg Dust mite
6. https://www.youtube.com/watch?v=j2IKBXKoV6Q House dust mites
7. https://www.youtube.com/watch?v=NctH7ZnBQ40 Cheese mites 3
8. https://www.youtube.com/watch?v=RWalcXkP2QM Grain mite (Chelhydra serpentine)
9. https://extension.umd.edu/Bulb-Mite-in-Garlic-Fields Bulb Mite Found in Problem Garlic
Fields
10. https://www.daf.qld.gov.au/plants/field-crops-and-pastures/broadacre-field-crops/grain-storage/
mites Mites in Stored Grains
Mites Predaceous on Pests of Agriculture
19
Abstract
Because of their small size and cryptic appearance, mites are difficult to detect
and hence in most cases their role in nature is overlooked. The role of parasitic
mites in reducing the population of honeybees (producer of honey) is well
known. The role of predaceous mites particularly of family Phytoseiidae is also
of great economic importance as regards management of spider mites on field
crops and in fruit orchards. Associations of different mite species and their eco-
nomic relevance on houseflies, mosquitoes, grasshoppers/locusts, scale insects,
whiteflies, psylla, thrips, caddisflies, collembolans, cockroaches, bark beetles,
stored grain beetles, moths and butterflies are discussed. Mite associations on
snails and slugs, symphylans, nematodes and earthworms and their economic
impact are also discussed. Predaceous mites belonging to mite families Acaridae,
Acarophenacidae, Anystidae, Arrenuridae, Ascidae, Bdellidae, Cheyletidae,
Cunaxidae, Limnesidae, Parasitidae, Pygmephoridae, Pterygosomatidae,
Tarsonemidae and Uropodidae are reported associated with insect pests of agri-
culture in different habitats.
Keywords
Predaceous mites • Insect pests • Grasshoppers • Psyllids • Leafhoppers
Acari, because of their small size and cryptic appearance, are difficult to detect and
hence in most of the cases their role in nature is overlooked. Once established in a
new area, because of their certain biological characteristics like high fecundity,
short developmental period, adaptation to different habitats in a locality, a myriad of
dispersal techniques and their diversified feeding habits, the mites may develop as
pests of crops and animals (including human beings) or may play the role of natural
control of pests by feeding on them both in terrestrial and aquatic habitats (Huggans

DOI 10.1007/978-981-10-1594-6_19
364 19 Mites Predaceous on Pests of Agriculture
and Blickenstaff 1996). The role of parasitic/predaceous mites in reducing the pop-
ulation of honeybees (producer of honey) is well known because of their economic
importance. Similarly the role of predaceous mites particularly of Phytoseiidae
family is also of great economic consequence. The relevance of parasitic mites in
declining bee populations and the role of different predaceous mites and insects on
phytophagous mites have been discussed under separate chapter in this book. Here
in this chapter, the role of different important mites (belonging to different families)
in influencing the population of some insect pests, nematodes, molluscs and other
acarines is discussed. Salient information about different mites which may be of
economic relevance in reducing or management of different insects is discussed
below.
19.1 Insects
19.1.1 Order Diptera
These insects have just one pair of wings; the hindwings are reduced to small club-
shaped balancing organs called halters. Gall midges, biting midges, mosquitoes,
sandflies, gnats, black flies, houseflies and tsetse flies are all dipterous insects.
Representatives are pollinators, parasites, predators or carry diseases that have a
huge impact on animals and man. Important information available in literature
regarding parasitic/predaceous role of different mites with respect to these insects is
discussed below.
19.1.1.1 Houseflies
Several species of macrochelid mites are reported feeding on eggs and immatures of
housefly. These mites provided effective control of houseflies in cattle manure and
sometimes even caused around 90 % reduction in fly population. Macrocheles gla-
ber feeds on Australian bushfly (Musca vetustissima) and sometimes provided total
control of flies. Another parasitic mite, Poecilochirus monospinosus Wise,
Hennessey and Axtell, feeds on larvae of housefly in manure. Life cycle of this mite
is completed in 17 days, and a female lays about 90 eggs in her lifetime. These mites
reported to provide effective control of houseflies in poultry manure mainly in late
spring and early summer. Macrocheles muscaedomesticae (Scopoli) infestation on
housefly is also reported in many publications. Uropodid mites, Fuscuropoda veg-
etans, are reported successfully controlling immatures of houseflies. Deutonymphs,
males and females of this mite feed on first instar larvae of houseflies and are also
reported controlling successfully adult population of houseflies. Results of some
laboratory studies indicated utilization of these mites in housefly control. This mite
is phoretic on dung beetles which facilitate its access to other suitable hosts.
Trombidiid mites, Allothrombium dipterae Ramaraju and Mohansundaram, are
found feeding on larvae of houseflies; Allothrombium muscaedomesticae
Vishnupriya and Mohansundarum are also found as parasites on the abdomen of
adult houseflies. Nymphal and adult stages of M. muscaedomesticae are reported
19.1 Insects 365
feeding on eggs of houseflies. Adults of M. muscaedomesticae and M. glaber are

reported using coprophagous flies for transport.
19.1.1.2 Mosquitoes
Arrenuridae Larvae of Arrenurus spp. are the most common parasites of mosqui-
toes. The larvae locate the host pupae and parasitize the adult mosquitoes as they
emerge. When adult mosquitoes land on water to oviposit, the mites detach and
continue their development in water exclusively on nulliparous mosquitoes.
Parasitization by Arrenurus madaraszi and A. pseudotenuicollis Wilson resulted in
high mortality in ovipositing females of Aedes and Culex species of mosquitoes.
Feeding and egg laying of mite-parasitized mosquitoes are adversely affected.
Water mites invariably feed on the same host species (both in the aquatic and aerial
stage). In many reports members of Arrenuridae family have been found to play an
important role in managing mosquito populations. The larvae of several mite genera
parasitize mosquitoes, while their nymphs and adults are free-living predators feed-
ing on mosquito eggs (Smith and Oliver 1986).
Hydryphantidae Sometimes hydryphantid mites are found on mosquitoes.

Parasitism by Panisopsis sp. curtailed flight activity and host-seeking behaviour of
mosquitoes, Aedes ventrovittis. Adults and nymphs of Piona nodata Muller are
found voracious feeders of mosquito larvae. One female mite may consume 100–
120 eggs; nymphs may consume five first instar larvae per day. But female adult
mite may consume 20–25 fourth instar larvae per day.
Limnesidae Parasitic mite, Limnesia jamurensis (Oudemans), is a great predator

of mosquito larvae; each mite consumed 6–8 larvae per day. These mites are in
general not specific predators of mosquitoes only.
19.1.2 Order Orthoptera
These insects have chewing mouthparts, and their hind legs are adapted for jumping.
Most species have toughened forewings to protect the larger hindwings. Important
representatives are grasshoppers, locusts, crickets, mole crickets, katydids, etc.
19.1.2.1 Grasshoppers/Locusts
Podapolipidae Podapolipus grassi Berlese is a common, most widespread ecto-

parasite of locusts. The mites infest third instar nymphs and subsequent stages.
Mites are found on the thorax or anterior abdominal segment and at the base of the
wings. The parasitic mite spends its entire life cycle on the locust dispersing only
during host mating. Mite infestation results in decreased host vigour. Podapolipoides
peruvensis Feldman-Muhsam is also reported as ectoparasite of grasshoppers
(Orphulella intricata and Allotryxalis strigata). Mites, P. madagascariensis and
P. poteri, are also reported as ectoparasites of grasshoppers in different reports.

From South India, P. nitidulae Ramaraju and Mohansundaram and P. hopperae are
also reported on grasshoppers (Branson 2003; Ramaraju and Mohansundaram
1996; Mohansundaram and Parmeshwaran 1989).
Trombidiidae Eutrombidium locustarum Walsh is an ectoparasite of many acridid

and tettigonid grasshoppers. Larvae parasitized even first instar nymphs. Up to 35
mites/nymph have been reported; mite feeding interfered in the normal flight of
adult as the wings were even broken in such cases. Mite nymphs and adults seek out
grasshopper eggs in the soil and devour them. A female mite consumed an average
of five eggs/day. One female mite is reported to lay up to 4700 eggs. As nymphs and
adults, these mites also prey on grasshopper eggs. Adult male mite requires about
three eggs to be able to reproduce, while female mite normally deposits about 4000
eggs, providing mite populations the potential to increase rapidly and substantially
as grasshopper populations increase. Larval mite infestation reduced survival of
grasshopper nymphs and adults up to 29 % and female reproductive output by about
47 %. The impact of ectoparasitism by mites could potentially help control grass-
hopper numbers if percent infestation of grasshopper can be increased. Mite abun-
dance seems to be limited by soils’ drainage (the poorer the drainage, the fewer the
mites). Rearing of mites under laboratory conditions is difficult, so advanced bio-
logical studies need to be done before using these ectoparasites for control of grass-
hoppers under field conditions (Rees 1973).
Another trombidiid mite, Eutrombidium trigonum Hermann, has also been

reported in literature as important ectoparasite of grasshopper, Hieroglyphus nigro-
repletus, leading to death of its host if the mite population per grasshopper is 40 or
more. Trombidiid mite, Eutrombidium rostratus (Scopoli), is also reported parasit-
izing locusts (Locusta migratoria) in laboratory cultures. In addition to above, there
may be many more reports of ectoparasitism by mites on different orthopterous
insects, but only the salient information has been given above (Mohansundaram and
Permeshwaran 1989).
19.1.3 Order Hemiptera
Bugs range from minute wingless insects to giant water bugs. All bugs have piercing
and sucking mouthparts in the form of a long rostrum made up of slender stylets
sheathed by the labium. Bugs, leafhoppers, aphids, lantern bugs, cicadas, whiteflies
and scale insects are all members of order Hemiptera. Information important for
insects of agricultural importance where important parasitic mites play some role in
their management is given below.
19.1 Insects 367
19.1.3.1 Scale Insects (Superfamily Coccoidea)

They are sedentary, wingless females and are flat and oval or elongate. Their bodies
may be covered with waxy secretions that form a soft or scalelike covering. Many
species are significant pests of crops such as citrus trees and coffee plants.
Camerobiidae Neophyllobius sp. is reported feeding on first instar nymphs (crawl-

ers) of California red scale (Quadraspidiotus ostraeformis).
Cheyletidae Cheletogenes ornatus (Canestrini and Fanzago) is reported as a pred-

ator of first instar nymphs of armoured scale insects. An adult female consumes
about 90 crawlers in its lifetime.
Cunaxidae Cunaxoides sp. is reported as an important predator of oyster shell

scale (Lepidosaphes ulmi) on apples.
Erythraeidae Leptus sp. and Bochartia sp. are reported to feed on mango pest
(Drosicha mangiferae). Balaustium putmani Smiley is reported to feed on eggs of
scale insects.
Eupalopsellidae These mites are commonly associated with armoured scale

insects. Eupallopsellid mite, Saniosulus nudus Summers, is reported actively feed-
ing on Florida red scales (Chrysomphalus aonidum). From 40 to 50 eggs are pro-
duced by one female mite and each female consumed about 260 eggs. The
predaceous mite also actively predated upon sugarcane scale insect (Melanaspis
glomerata). Mites crawled under the mother scale insects and attacked the emerging
crawlers there. S. nudus is reported to have potential for biological control of scale
insects.
Hemisarcoptidae These mites are often associated with armoured scale insects.
Hemisarcoptes malus Shimer is reported feeding on eggs of a number of scale
insects. Hemisarcoptes coccophagus Meyer, H. cooremani Thomas and H. malus
are the major mortality factors of the oyster shell scale insect (Lepidosaphes ulmi)
infesting apple, date-palm scale (Parlatoria blanchardi) and citrus scales. Female
mites feed on adults and eggs of scale insects; its introduction in New Zealand in
1987–1988 has resulted in about 80 % reduction of population of scale insects. H.
coccophagus prefers ovipositing scale females. The decreasing order of preference
for the scale stages as reported was young females, immature males and second
instar nymphs.
Pyemotidae Pyemotes herfsi Oudemans is reported to parasitize soft scale

(Saissetia oleae) – a citrus pest in Argentina.
19.1.3.2 Whiteflies (Family Aleyrodidae)

The wings of these insects are either white or mottled, with a distinctive dusting of
white powdery wax over the surface. Many whitefly species are serious pests
particularly during warmer months. Well-known species include Trialeurodes vapo-

rariorum and Bemisia tabaci.
Phytoseiidae Phytoseiid mites, Amblyseius swirskii Athias-Henriot and Euseius

rubini Swirski and Amitai, are reported as effective predators of tobacco whitefly
(Bemisia tabaci) (Wyoski and Cohen 1983).
19.1.3.3 Psylla (Family Psyllidae)

Psyllas are also known as jumping plant lice variously coloured, look-like small
leafhoppers but with longer antennae. The two pairs of oval wings are held rooflike
over the body. The head has a short three-segmented beak. Some psyllids are signifi-
cant plant pests.
Erythraeidae Balaustium putmani Smiley has been reported as a predator of pear

psylla (Psylla pyricola).
19.1.3.4 Aphids (Family Aphididae)

The aphids are small, soft-bodied and mostly green, pink, black or brown. The
abdomen usually carries a pair of short tubes, called cornicles.
Acaridae Acarid mite, Tyrophagus putrescentiae (Schrank), is reported feeding on

grape phylloxera (Daktulosphaira vitifoliae) and preferred them over fungi and
dead arthropods.
Anystidae Anystid mite, Anystis agilis (Banks), has been reported in literature as
an important predator of woolly pine needle aphid.
Thrombidiidae Thrombidiid mite, Allothrombium pulvinum Ewing, in some parts

of China has been reported as important parasite of aphids. Adult female may con-
sume two to three aphids in an hour. Releases of mite at the rate of three or more
mites per aphid controlled successfully the aphids. This is a univoltine species and
at 20–30 °C has been reported to complete its development in about 75 days. In
literature, mass rearing of trombidiid mites has been recommended under labora-
tory conditions for releases and successful control of its prey. Allothrombidium
fuliginosum Hermann is the most widely distributed species. Adult mites feed on all
instars of aphids but certainly prefer immobile stages.
19.1.3.5 Leafhoppers (Family Cicadellidae)

Leafhoppers are generally slender, with broad or triangular heads. Many species are
brown or green, have distinctive hind tibiae and jump very well. Leafhoppers are
pests of vital crops such as rice and corn.
Anystidae Anystid mite, Anystis agilis, has been reported as an important predator
of grape leafhoppers.
19.1 Insects 369
19.1.3.6 Bugs
True bugs belong to suborder Heteroptera. All bugs have piercing–sucking mouth-
parts in the form of a long rostrum made up of slender stylets sheathed by the
labium. Many bugs that suck sap are serious crop pests. Mites reported predaceous/
parasitic on these bugs are discussed hereunder.
Pterygosomatidae Some species of pterygosomatid mites have been reported as

parasites on blood-sucking pests (Triatominae). Pimeliaphilus plumifer Newell and
Rickman, has been found parasitizing blood-sucking bugs. Mites completed their
life cycle in 60 days at 30 °C and 15–20 % relative humidity. On an average 60 eggs/
female are laid. As reported in some studies, when more than 25 mites/adult bug are
found, then they kill their host within one hour.
Otopheidomenidae Hemipteroseius adleri Costa is reported parasitizing red fire-

bugs (Pyrrhocoris apterus). All the stages of mites are found on the infested bugs.
Mites are found mainly in the anterior part of the bugs’ abdomen on the surface of
the tergites under the insects’ wings. The mean number of mites per host is reported
as 4.8 (range 1–56). Sometimes moulted skins of the mites, the excrement of mites
and even dead mites are found on the infested bugs. Juvenile stages of host insect
were free of mites. Besides H. adleri, other otopheidomenids found parasitic on
Pyrrhocoridae and Lygaeidae bugs are H. womerslei Ewans, H. indicus {Krantz
and Khot}, H. parvulus Treat and H. sabbaticus Treat. Ectoparasitic mite,
Hemipteroseius vikrami Menon, has been reported on bugs, Dysdercus koenigii and
D. cingulatus (Menon et al. 2011).
19.1.4 Order Thysanoptera
Thysanoptera are commonly known as thrips. They are small, slender insects, usu-
ally with two pairs of narrow, hair-fringed wings and distinctive mouthparts which
include a pair of mandibles in which one is small and the other is needle-like. Many
thrips are plant pests especially of cereal crops. Associations of various mites with
thrips are discussed below.
Phytoseiidae In addition to being spider mite predators, some phytoseiid mites are
found predaceous primarily on thrips. Phytoseiid species, Amblyseius cucumeris
(Oudemans) and A. mckenziei Hughes (also known as A. barkeri), are reported feed-
ing on western flower thrips and the onion thrips (Thrips tabaci) which are serious
pests in greenhouses. If the phytoseiid mites are released in earlier stages of thrip
infestation, these mites can eliminate thrip population in greenhouses. Releases of
10–25 predatory mites per leaf of sweet peppers infested by T. tabaci are recom-
mended in greenhouse cultivation. But for cucumber leaves, suppliers recommended
50 predatory mites per plant. But more detailed studies are required considering
cost–benefit ratios of releasing predatory mites for thrip control. Phytoseiid mites,
Euseius hibisci and Galendromus (=Typhlodromus) occidentalis (Nesbit), are also

reported predaceous on different thrips.
Anystidae Anystis agilis are reported in many research publications to control cit-
rus thrips.
Pyemotidae Adactylidium sp. has also been reported parasitizing the eggs of
thrips, Gynaikothrips ficorum – a pest of ornamentals.
19.1.5 Order Trichoptera
The members are commonly known as ‘caddisflies’. They are slender, dull adults
and look mothlike, but unlike moths their bodies and wings are covered with hairs
but not scales. Antennae are long, thin, multisegmented; mouthparts are weakly
developed. The aquatic larvae usually pupate inside cases made with sand grains
and twigs. Water mites have been reported as parasites of caddisfly larvae.
Hydrachnidia reported as parasites of different trichopteran insects are
Philopotamidae, Psychomyiidae, Lepidostomatidae, Hydryphantidae (Protzia spp.),
Momoniidae (Momonia sp.), Sperchontidae (Sperchon sp.), Unioncolidae
(Unioncola spp. and Koenikea spp.) and Aturidae (Albia spp.). In most cases, mite
larvae are found attached to adults, but in family Limnephilidae and Leptoceridae,
representatives are also found on larvae.
19.1.6 Order Thysanura
They are primitive, wingless insects with elongate, flat bodies that may have scales
on their surface. They have simple mouthparts and may have small widely separated
compound eyes or no eyes at all. The abdominal segments have projections called
styles. They have representatives commonly known as silverfish or booklice and
occupy a wide range of habitats.
Cunaxidae Cunaxid mite, Cunaxa capreolus Berlese, is an important predator of

silverfish. At 30 °C, one generation of mite is completed in about 4 weeks. One
adult mite is reported to consume about 230 booklice during its development, each
female laid on an average of 45 eggs.
19.1.7 Order Collembola
Collembola are commonly known as springtails. They have a ventral tube on the
undersurface of the abdomen and in some species used for gripping smooth sur-
faces. Predaceous mites reported feeding on collembolans are discussed below.
19.1 Insects 371
Anystidae Anystis salicinus is reported as an effective predator of lucerne flea

(Sminthurus viridis).
Bdellidae Bdellid mite, Bdellodes lapidaria Kramer, is reported many times

actively feeding on lucerne flea. Another bdellid mite, Bdella depressa Ewing, has
been reported feeding on springtails.
Parasitidae Members of this family are mostly predators of different soil pests.
Parasitic mite, Pergamasus longicornis (Berlese), is reported feeding on soil
collembolans.
Laelapidae Hypoaspis sp. has been found in different research publications to

prey upon collembolans.
19.1.8 Order Blattoidea
Blattoidea are commonly known as cockroaches. They have an oval, flat shape; the
head is often covered by shield like pronotum. They possess two pairs of wings;
forewings are usually tough and cover large membranous hindwings. They are
adapted to human habitats and thriving in warm, unsanitary conditions. The follow-
ing mite associations have been reported commonly.
Pterygosomatidae Pimeliaphilus plumifer Newell and Ryckman and some other

Pimeliaphilus spp. are found infesting cockroaches under laboratory culture condi-
tions. This species also infests blood-sucking bugs. Egg capsules produced by heav-
ily parasitized cockroaches even fail to hatch and hence host population is reduced
considerably. P. podopolipophagous has frequently been reported infesting cock-
roach breeding cases. The eggs are laid by the mite either singly or in batches on the
soft wood or other objects in the breeding cases; occasionally they were also laid
glued on the wings of cockroaches. If more than 20 mites per individual were found,
then there are chances of mortality of mite-infested cockroaches.
19.1.9 Order Coleoptera
Coleoptera are commonly known as beetles. They have toughened forewings, also
known as elytra. The elytra may be short, but in all species they meet down the
middle of the body. Most beetle species are herbivorous, but there are many scaven-
gers and predators. The role of mite associations with different beetles as found in
literature is given below.
19.1.9.1 Bark Beetles

They are fairly small beetles (0.25–0.70 cms) that unusually develop on the cam-
bium under the bark of trees. Adult beetles first cut distinctive galleries in which
they mate and lay eggs. More than 100 species of mites, representing 80 genera and
42 families, are reported on bark beetles (Dendroctonus frontalis) alone. Mites asso-
ciated with bark beetles (Lindquist 1967; Lindquist 1969) are discussed below.
Laelapidae Laelapid mites, Dendrolaelaps quadrisetus Berlese, Histiogater

arborsignis Woodring, Proctolaelaps fiseri Samsinak and Proctolaelaps hystricoi-
des Lindquist, are found in a large number of habitats. These mites also tend to ride
on a large number of insects. D. neodiseius associated with bark beetles may also be
having mutualistic association with beetles Sometimes Proctolaelaps xyloteri
Samsinak from 22 to 33 % larvae per gallery found. P. eccoptogasteris Vitzthum
preyed on eggs and larvae of beetle (Phloeotribus scarabaeoides). At times 10–15
ovigerous females emerged on one prey.
Pygmephoridae Pygmephorus spp. have been reported feeding on bark beetles.
Pyemotidae Pyemotes scolyti (Oudemans) and P. dryas Vitzthum are known as

parasitic on bark beetles.
Tarsonemidae Iponemus spp. are egg parasites of beetles (Ips and Orthotomicus).
Tarsonemoides spp. are specialized as egg parasites of bark beetles (Ipps spp.).
Many other members of families Digmasellidae, Ascidae and Uropodidae;

Tarsonemoidea and Acaroidea superfamilies are known associated with bark beetles
in different parts of the world (Moser 1975).
Acarophenacidae Paracarophenax spp. have been reported as egg parasites of

bark beetles.
19.1.9.2 Stored Grain Beetles

Acarophenacidae Acarophenax lacunatus (Cross and Krantz) is reported as para-
sites of Rhyzopertha dominica and Cryptolestes ferrugineus which are pests of
stored grains. A. tribolii parasitizes the grain beetle, Tribolium spp. Mites are found
on the cuticular region of the host (intersegmental region) beneath the wings. The
highest parasitism leads to a significant decline in populations of beetle species.
Ascidae Blattisocius tarsalis (Berlese) mites preferred eggs of beetles Tribolium

spp. and Rhyzopertha dominica.
Cheyletidae Acaropsis docta (Berlese) feeds on stored product insects like

Tribolium granarium, T. castaneum, R. dominica and Lasioderma serricorne.
Female mites prepare a net of web around them; the development is completed in
14–15 days. The male and female consumed an average of 10 and 20 eggs of
Trogoderma spp. Acaropsis sollers have also been reported to feed on Trogoderma
eggs; younger or fresh eggs were more preferred for predation than older eggs.
19.1 Insects 373
Pyemotidae Straw itch mite, Pyemotes tritici Oudemans, has been reported as an
effective biological control agent for Oryzaephilus mercator and Lasioderma ser-
ricorne (both Coleoptera) and some lepidopterous larvae. P. herfsi Oudemans para-
sitizes Tribolium castaneum and is known to locate the beetle pest up to 7 cm depth
of bran.
19.1.9.3 Mites Predaceous on Other Coleoptera

Larvae of water mite, Eylais extendens Muller, are found as voracious predators of
cockchafer beetles, Haliplus spp. Rhynchopolipus (=Tetrapolipus) rhynchophori
(Ewing) are reported as parasite of palm weevil (Rhynchophorus palmarum).
Urodinychus sp. and Fuscuropoda sp. have been found under elytra of red palm
weevil. Predatory acarid mite, Sancassania sp., is reported to feed on eggs of sugar-
cane white grub (Holotrichia serrata) (Moser 1975). One generation of mite was
completed in 4 days. Adult mites moved deep into the soil up to a depth of 18 cm
when the soil was dry, but failed to move down when the soil was wet. Macrocheles
spp. have been found attached singly to the hind leg of the beetles, Corpus spp.
Allothrombium cicindelae Ramaraju and Mohansundaram are found parasitic on
cicindellid beetles. On dung beetles, Tarsopolipus ramakrishnai Ramaraju and
Mohansundaram is reported feeding on the beetle. Acarophenacid mite, Acarophenax
manhunki Steinkraus and Cross, is reported as a parasite of lesser meal worm
(Alphitobius diaperinus). On corn rootworm, Diabrotica undecimpunctata an acarid
mite, Tyrophagus putrescentiae was reported feeding.
Sometimes uropodine mites are found associated with doryline and ecitonine
army ants in the neotropics. Some mites are known to be kleptoparasites, e.g.
Antennophorus grandis on the ant, Lasius flavus. True ectoparasitism has been dem-
onstrated for uropodine mite, Macrodinychus sellnicki Hirchman and Zirngiebl-
Nicol (Dinychidae), which slowly sucks the pupae of the Neotropical ant,
Paratrechina fulva into shrivelled little houses.
19.1.10 Order Lepidoptera
Lepidoptera are commonly known as moths and butterflies. They possess tiny over-
lapping scales on the body and wings, with multisegmented antennae. Butterflies
have club-ended antennae, which moths usually lack. The mouthparts usually form
a proboscis for taking nectar. In larval stage, the majority have chewing mouthparts
and are herbivorous. Mites associated with moths and butterflies (Treat 1975;
Housewart et al. 1980) are discussed hereunder.
Ascidae Ascid mite, Blattisocius tarsalis, prefers eggs and first instar larvae of
Indian meal moth (Ephestia sp.). B. tarsalis is reported effective against Ephestia
cautella, in bagged maize in Kenya.
Pyemotidae Straw itch mite, Pyemotes tritici, has been reported as an effective
biocontrol agent of Indian meal moth and fig moth (Cadra cautella). P. herfsi is
known to parasitize the pink bollworm, Pectinophora gossypiella – an important
pest of cotton crop – and caused up to 85 % mortality of their host. Pyemotes ventri-
cosus (Newport), commonly known as ‘hay itch’ mites, feeds on lepidopterous lar-
vae. The life cycle takes from 5 to 30 days, depending on temperature and relative
humidity.
Erythraeidae Two erythraeid mites, Balaustium dowelli Smiley and Erythraeus

whitcombi Smiley, are reported feeding on eggs of boll worms (Noctuidae). The
mite destroyed 15 eggs in about 6 h and are reported to be most injurious under hot
and dry conditions (Smiley 1966).
Cheyletidae Cheletomorpha lepidopterorum Shaw are sometimes found on moths

and prey on some other parasitic mites carried by the insect.
Laelapididae Dicrocheles hippioides Treat (=Myrmonyssus phalaenodectes) is

the most interesting parasite of the thoracic tympanic organs of certain noctuids
chiefly those of the genus Leucania. Its colonies, whether founded by a single
female or by several, occupy and destroy the function of one but not of both the
hosts’ ears. Each female can produce 80–100 eggs. The young develop to maturity
on the original host and the fertile females then seek another moth on which to
repeat the cycle.
19.1.11 Order Hymenoptera
Sawflies, ants, bees and wasps are common representatives. Most members have
two pairs of membranous wings, joined in flight by tiny hooks. The ovipositor of
female bees, ants and social wasps has evolved into a sting. Metamorphosis is com-
plete. Too many associations of mites with bees are reported and will be discussed
in a separate chapter as the bees have great economic relevance. Mites are also
found associated especially with hymenopterous families Formicidae, Apidae and
Bombidae and acarid and europodid mites.
19.2 On Snails and Slugs
Snails and slugs are known as gastropods; they are close relatives of clams, mussels
and other molluscs. They differ from arthropods (insects, mites, spiders, etc.) in
lacking distinct segmentation or an external skeleton. Their body is soft and moves
by means of a broad muscular ‘foot’ that covers the underside. On slugs, a large lobe
called the mantle is present on the front half of the back; this is covered by a hard
shell in snails. Slugs and snails typically feed at night, avoiding sunny, drying con-
ditions. During the day most slugs migrate to sheltered areas under debris and in soil
19.3 On Symphylans 375
cracks. They have an extraordinary diversification of habitats. Representatives live

in gardens, in woodland, in deserts and in small ditches, rivers and lakes (Thrk and
Phillips 2009).
Ereynetidae Riccardoella limacum (Schrank), commonly known as slug mite, are

commonly found on wild slugs and snails. They live on the body surface within the
mantle cavity of a variety of terrestrial molluscs. In case of infestation, small whit-
ish mites can be seen moving rapidly over the slugs’ body, often present in consider-
able numbers without any apparent injury to the host mollusc. Mites hide under the
shell rim and congregate in the pneumostome. This mite has been reported as a true
parasite feeding on the blood of its host. Slugs, Arion ater, Milax gagates and
Agriolimax agrestis, are the preferred hosts of slug mite. Snails are also reported as
the preferred hosts of slug mite. A snail is reported bearing up to 60 mites but with-
out any observable damage.
Uropodidae Fuscuropoda marginata (Koch) is reported influencing the popula-

tion of slugs Laevicaulis alte, which are serious pests of agriculture in some circum-
stances. The mites fed on mucus from the body surface of the slug. Mites feed by
making holes (0.3–1.0 mm in depth and 0.2–0.4 mm in diameter) on the dorsal
surface of the slug body; these holes in the course of time turned to cancerous spots
and the infested slug becomes inactive in 40–44 h and died on the third day. At
times as many as 18 mites are observed feeding on a slug. Mites preferred adult
slugs, but juveniles were not spared. F. marginata may help in controlling slug, L.
alte.
Unioncolidae Unioncola diversities and U. scutigera are sometimes found para-

sitic on freshwater edible bivalve molluscs (Lamellidens marginalis). These mol-
luscs are a source of food and are a biological indicator species of freshwater. Mites
are found as parasites on both external and internal environment of the host.
Unioncola intermedia on Anodonta anatina and U. ypsilophora on A. cygnea are
found as parasites. Unioncolid larvae hatching from eggs kept in dark at 12–20 °C
live for at least one week. From one to seven females are found per host, but never
more than one male. The largest number of nymphs and females is reported during
winter months.
19.3 On Symphylans
They are small, soft-bodied myriapods, belonging to order Symphyla and are usu-
ally white or pale brown or grey looking creatures. The head bears a pair of long and
thread-like antennae and three pairs of mouthparts. Behind the head is a trunk, made
up of 14 segments. Symphylans live in soil or leaf litter and feed on the plant
material.
Parasitidae Pergamasus quisquiliarum mite is closely related to plant root system

and effectively regulates the garden symphylans, Scutigerella immaculata – a soil-
borne pest of many crops. The mite completed one generation in 17 days, while the
symphylan required about 87 days to complete its life cycle. Female mite deposited
33 eggs near the plant roots. About 15 symphylans have been reported consumed by
one mite in its lifetime.
19.4 On Nematodes
The nematodes or roundworms (phylum Nematoda) are the most numerous multi-
cellular organism on earth. A handful of soil will contain thousands of the micro-
scopic worms – many of them are parasites of insects, plants or animals. Nematodes
have been characterized as a tube within a tube. In size they range from 0.3 mm to
over 8 m.
Galumnidae Pergalumna spp. are reported to thrive on nematodes and have poten-
tial as biocontrol agent.
Laelapidae Hypoaspis spp. are reported actively feeding on plant parasitic nema-
todes and are currently being exploited. These mites lay their eggs in soil which
hatch in 1–2 days. Nymphs and adults feed on soil-dwelling insects, other mites and
nematodes. Currently commercially available cultures are shipped as adults. Release
rates of 10,000–25,000 mites/acre are recommended by commercial sources for the
effective control of soil organisms including nematodes.
Uropodidae Fuscuropoda vegetans (DeGeer) have been reported successfully

feeding and developing on nematodes. Mite population was found during autumn
and spring; however, their population was very low during winter. The mite is
reported to be phoretic on dung beetles which facilitate their access to other suitable
hosts.
Nematophagous mites have been used as potential biocontrol agents of root-knot

and fungus-feeding nematodes. Oribatid mites Fuscozetes sp., Galumna spp. and
Nothrius spp. have been reported feeding on free-living nematodes. Some of the
astigmatid mites – Tyrophagus putrescentiae and Histiostoma bakeri Hughes and
Jackson – are also reported feeding on nematodes. Nematophagous mites seem to
be abundantly available in the soil ecosystem and are potential biocontrol agents of
plant parasitic nematodes.
19.5 Mite Associations on Earthworms
Earthworms belong to phylum Annelida and class Oligochaeta. They are tube-
shaped, segmented creatures commonly living in soil. Their activity in the soil aer-
ates and mixes the soil and is constructive to mineralization and nutrient uptake by
vegetation. Some natural enemies including some parasitic mite associations inter-
rupts their proper development. Mite associations found in literature are reported
hereunder.
Anoetidae Histiostoma murchiei Hughes and Jackson and H. berghi Jenson are
reported parasitizing cocoons of earthworms. All stages of mites except the hypopus
occur inside the cocoon. Males develop earlier than females. The incubation period
is about 3.8 days; the larval stage lasts 1 day. The first nymphal stage lasts from 3 to
12 days. Once the hypopus is formed, it remains inside the protonymphal skin or
emerges and moves about the cocoon. Once the hypopus gains entrance into the new
cocoon, it develops into the tritonymphal stage which lasts for only one day. H.
murchiei is reported to parasitize about 20 % of the cocoons of the earthworms,
Apprectodea caliginosa, and about 7 % of cocoons of the earthworm, Lumbricus
terrestris, are also found infested by the above mite.
Uropodidae Sometimes reports of mite problems in earthworm beds are reported.

Uropodid mite, Uropoda (=Fuscuropoda) agitans Newell, is reported parasitizing
cocoons of earthworms. Sometimes earthworm beds can become so badly infested
with mites that the worms’ food supply is endangered and the worm population
declines.
19.6 Conclusions
In nature many associations of different mite species belonging to different mite

families with the insect pests of different crops, pests of veterinary importance and
in soil are reported with different insects, nematodes and earthworms. Many preda-
ceous mites belonging to mite families Acaridae, Acarophenacidae, Anystidae,
Arrenuridae, Ascidae, Bdellidae, Cheyletidae, Cunaxidae, Erythraeidae,
Eupalopsellidae, Galumnidae, Laelapidae, Limnesidae, Parasitidae, Pygmephoridae,
Pterygosomatidae, Tarsonemidae, Trombidiidae and Uropodidae are reported as
predator/parasites on insect pests of agriculture in different habitats; however,
detailed experiments are lacking and observations reported are on the basis of con-
ditions of a particular place. Detailed studies on different prey–predator interac-
tions, their mass multiplication keeping in view the cost of production and their
release rates for field application are urgently needed. Earlier, most of the observa-
tions are confined to small sample size and that too under laboratory conditions.
Further Reading
Branson DH (2003) Effects of a parasite mite on life-history variation in two grasshopper species.
Evol Ecol Res 5:397–404
Housewart MW, Jennings DT, Brekett LP, Brann TB (1980) Parasitic mites (Acari Erythraeidae)
on spruce budworm moths (Lepidoptera: Tortricidae). Can Entomol 112:193–197
Huggans JL, Blickenstaff CC (1996) Parasites and predators of predators in Missouri, Reserch
Bulletin 903, University of Missouri Agricultural Experiment Station. University of Mossouri,
Columbia, 40 pp
Hunter PE, Rosario RMT (1988) Associations of Mesostigmata with other arthropods. Annu Rev
Entomol 33:393–417
Lindquist EE (1967) Mites parasitizing eggs of bark beetles of the genus Ips. Can Entomol
96:125–126
Lindquist EE (1969) Mites and the regulations of bark beetles. In: Proceeding 2nd International
Congress Acarology, 1967, pp 389–399
Menon EP, Joshi S, Hussain M, Ramamurthy VV (2011) A new species of Hemipteroseius (Acari:
Otopheidomenidae) parasitic on Dysdercus (Hemiptera: Pyrrhocoridae) in India. Zootaxa
2800:53–636
Mohansundaram M, Parmeshwaran S (1989) Mites infesting the grasshoppers occurring in differ-
ent ecosystems in Coimbatore. J Biol Control 9:28–30
Moser JC (1975) Mite predators of the southern pine beetle. Ann Entomol Soc Am 68:113–116
Ramaraju K, Mohansundaram M (1996) New species of Podapolipus, Podapolpides and
Tarsopolipus (Acari: Podapoliidae) from South India. Int J Acarol 22:33–41
Rees NE (1973) Arthropod and nematode parasites, parasitoids, and predators of Acarididae in
America North of Mexico, Technical Bulletin No. 1460. U.S. Department of Agriculture,
Agricultural Research service, Washington, DC, p 228
Smiley RL (1966) Further descriptions of two erythraeids predaceous upon cotton bollworm eggs.
Proc Environ Soc Wash 68:25–28
Smith BP (1988) Host-parasite interaction and impact of larval water mites on insects. Annu Rev
Entomol 33:393–417
Smith IM, Oliver DR (1986) Review of parasitic associations of larval water mites (Acari:
Parasitengona: Hydrachinidia) with insect hosts. Can Entomol 118:407–472
Thrk FA, Phillips SM (2009) A monograph of the slug mite – Riccardoella limacum (Schrank).
Proc Zool Soc London 115:448–472
Treat AE (1975) Mites of moths and butterflies. Cornell University Press, Ithaca, 362 pp
Wirth WW (1997) A review of the pathogens and parasites of the biting midges (Diptera:
Ceratopogonidae). J Wash Acad Sci 67:60–75
Wyoski M, Cohen M (1983) Mites of the family Phytoseiidae (Acarina: Mesostigmata) as preda-
tors of the Japanese bayberry whitefly, Parabemisia myricae Kuwana (Homoptera:
Aleyrodidae). Agronomie 3:823–825
1. http://www.redwormcomposting.com/worm-bin-creatures/parasitic-worm-bin-mites/. Parasitic
Worm Bin Mites
2. https://www.youtube.com/watch?v=9bwFcRiT9so. Balaustium (Erythraeidae) on a Dead
Mosquito
Further Reading 379
3. https://www.youtube.com/watch?v=n9SI2sE4dSk. Limonica, Kopperts latest predatory mite

against thrips and whitefly
4. https://www.youtube.com/watch?v=FqV7Cj_BtW8. Mites living in Fruit Fly culture medium
5. https://www.youtube.com/watch?v=JOqXAPxx50k. Predatory mites
Biological Control of Phytophagous
Mites 20
Abstract
Biological control is the beneficial action of predators, parasites, pathogens and
competitors in controlling pests (including insects, mites, weeds and plant dis-
eases) and their damage. Different species of phytophagous mites including spi-
der mites, false spider mites, eriophyid or worm mites and bulb mites are
naturally controlled by different predaceous mites, mite predaceous insects and
acaropathogens (including viruses, fungi, bacteria, etc.). Predaceous mites are
also commonly known as micropredators, and they belong to families
Phytoseiidae, Stigmaeidae, Bdellidae, Cunaxidae, Tydeidae, Anystidae,
Cheyletidae and Erythraeidae. Some representatives of Phytoseiidae and
Stigmaeidae have been reported very useful in managing pest mites on different
fruit crops, field crops and ornamental plants both under field conditions and
under protected cultivation. Phytoseiid mites Neoseiulus californicus, Amblyseius
fallacis, Phytoseiulus persimilis, Metaseiulus occidentalis and Amblyseius swir-
skii are reported to control pest mites, white flies, scale insects and thrips in some
conditions. Insect predators particularly beetles, Stethorus spp. and Oligota spp.,
and thrips have also been reported managing successfully pest mites. Microbial
diseases caused by viruses and fungi (order Entomophthorales) are also reported
controlling mite pests in some circumstances. Some spider mites, galumnid
mites and eriophyid mites are also reported to reduce damage by weeds.
Keywords
Predaceous mites • Microbes • Weeds
Biological control is the beneficial action of predators, parasites, pathogens and

competitors in controlling pests (including insects, mites, weeds and plant disease)
and their damage. Biocontrol provided by these living organisms (collectively
known as ‘natural enemies’) is especially important for reducing the numbers of
pest mites. It can be an important component of integrated pest management (IPM).

DOI 10.1007/978-981-10-1594-6_20
382 20 Biological Control of Phytophagous Mites
The intent of biological control is not to eradicate pests but to keep them at tolerable
levels at which they cause no appreciable harm. Conservation, augmentation and
classical biological control (also called importation) are tactics for harnessing the
effects of natural enemies. Biological control may occur naturally in the field or
result from manipulation or introduction of biological control agents by people.
Weeds are the unwanted plants and compete with main crops for nutrients, moisture
and sunlight resulting in decreased crop quality, serving as alternate hosts of mite
pests, higher production costs and reduced crop yields. These unwanted plants
(weeds) are sometimes highly damaged by some host-specific mites resulting in
their destruction. So weed control is also considered from natural control point of
view in this chapter. To get the greatest benefit from both natural control and bio-
logical control, it is necessary to recognize natural enemies and understand their
characteristics such as life cycle, preferred host mites and efficiency at controlling
pests. The management of mite pests under different ecosystems (including both
field and greenhouses) is considered separately below (Albajes et al. 1999;
Dalrymple 1993).
20.1 Biological Control of Phytophagous Mites
Different phytophagous mites including spider mites (family Tetranychidae), false

spider mites (family Tenuipalpidae), eriophyid or worm mites (family Eriophyidae)
and bulb mites (Acaridae) are naturally controlled by different predaceous mites,
mite predaceous insects and acaropathogens (including viruses, fungi, bacteria,
etc.) (Bjornson 2008; Castillo et al. 2015). Different natural enemies and acaro-
pathogens are discussed below briefly.
20.1.1 Predaceous Mites
Predaceous mites are also commonly known as micropredators. They belong to

families Phytoseiidae, Stigmaeidae, Bdellidae, Cunaxidae, Tydeidae, Anystidae,
Cheyletidae and Erythraeidae (Walter and Proctor 2013). A brief account of differ-
ent mite families and their important representatives found as important predators of
different phytophagous mites is given below (ER12).
20.1.1.1 Phytoseiidae
Phytoseiid mites are free-living terrestrial mites commonly found on foliage, bark
and humus in all parts of the world including from alpine and arctic tundra to tropi-
cal jungles. Most species move much more rapidly than their prey and appear shin-
ing when alive. They may be having white to brown look; the colour of the material
fed upon makes them appear grey or black. Phytoseiidae are approximately of the
same size as spider mites.
Phytoseiid mites are most effective and widespread predators of plant-feeding
mites; their larvae are less mobile than the protonymphs and other advanced stages.
20.1 Biological Control of Phytophagous Mites 383
The most effective species are capable of producing 30–60 eggs during their life,
which is generally equivalent to the egg laying capacity of their prey. Phytoseiid
mites are also active throughout the year on evergreen trees and shrubs in warmer
climates; however, under such conditions the non-availability of food supply is the
limiting factor. Phytoseiid mites show a great diversity in feeding habits; some are
preferably carnivorous and other species prefer pollen, nectar and crawlers of scale
insects, whiteflies and thrips. Some species also feed on fungi and even honey dew
from aphids, scales, whiteflies and mealybugs. Phytoseiids that can survive on food
species other than prey mites have a better chance of surviving adverse conditions.
A brief information about most important phytoseiid species found useful in con-
trolling pest mites is given below (Del Fosse et al. 1978; Hoy and Jeyaprakash 2008;
Stenseth 1979; Gerson 2014).
Neoseiulus californicus Neoseiulus californicus is reported to control two-spotted

spider mite, Tetranychus urticae, on various crops in temperate and subtropical
regions around the world (ER7 and ER11) and are found on many plants including
avocado, citrus, cassava, corn, grapes, strawberries, several vegetable crops and
many ornamental plants. They prefer warm (10–33 °C) temperatures but can toler-
ate much colder temperatures for short periods and can tolerate a wide range of rela-
tive humidity (40–80 %), but prefer high humidity. Eggs are football shaped about
0.04 mm in length and are pale whitish. Larvae are translucent; adults are about
0.1 mm in length and are oval in shape. Both the nymphal and adult stages feed
actively on their prey; the total developmental time may range from 4 to 12 days
depending on the temperature. Adult female mites live for about 20 days.
N. californicus have been reported in different studies feeding on prey mites T.

urticae, Oligonychus pratensis, O. persea, Panonychus ulmi, Phytonemus pallidus
and Polyphagotarsonemus latus; many of them are serious crop pests (ER3 and
ER6). Predatory mites are also known to feed on thrips infesting many crops. N.
californicus is commercially used against two-spotted spider mites in different parts
of the world on avocado, citrus, dwarf hops, grapes, raspberries, roses, many orna-
mental plants, strawberries and several vegetable crops in greenhouses as well as
under field conditions. In case of strawberries, releases at the rate of one female per
strawberry plant is reported to maintain T. urticae below threshold level (5% of
leaves infested).
Amblyseius fallacis Releases during June and July at the rate of 1000 and 2000
mites per peach tree infested with spider mites, Panonychus ulmi and T. urticae in
Ontario (Canada), revealed that significant control of prey mites is not received in
the year of release. Even in the following year, significant increase in phytoseiid
mites not recovered. But trends revealed that A. fallacis can be a useful predator in
some fruit orchards. However, the role of phytoseiid mites in controlling both the
prey mites needs more investigation.
Phytoseiulus persimilis Phytoseiid mite, P. persimilis, is known as a fast action

predator and has been used for the control and management of various spider mite
pests under both greenhouses cultivation and under open field conditions. They are
tiny (0.5 mm long), hunter-orange-coloured mites and lay eggs among webs of spi-
der mites. Immature development takes about 8 days, and adults may have longevity
of 36 days at optimum temperatures (between 25 and 30 °C) with a relative humid-
ity of 60–90 %. Cooler temperatures hamper the development and reproduction of
mite. P. persimilis is mass reared in insectaries and is shipped as adults and imma-
tures in vermiculite or as adults and immatures (including eggs) on bean leaves
(ER2 and ER9). The vermiculite product is flowable and easy to distribute. This
predaceous mite has been successfully used for controlling common red mite,
Tetranychus cinnabarinus and T. urticae, and the Pacific mite, Tetranychus pacifi-
cus. But there is one drawback in utilization of P. persimilis in cases when no spider
mites on their hosts are available for their consumption; in such cases these mites
being specialist predators of spider mites may quickly starve to death. Unlike N.
fallacis which can support its needs with pollen, or N. californicus which can sur-
vive longer without eating anything for their utilization as biological control, one
may have to provide mite food all the times (ER8).
Successful control of spider mite, T. urticae, has been reported on strawberry

when P. persimilis were released at the rate of 3 mites/m2 (ER5). At least two
monthly releases of this mite on strawberry crops maturing at normal time (from
February to early March) and early crop (maturing in December–January) are
needed in Korean climates. For more effective control of spider mites on strawber-
ries, predatory mites should be released early in the cropping season when spider
mite population is fairly small. However, if predatory mite is released at the time
dense population of T. urticae is found, the control effect is limited.
Metaseiulus (=Typhlodromus or Galendromus) occidentalis This predaceous mite

has been reported as an obligate predator and one of the successful biological con-
trol agent of spider mites in vineyards, almond and apple orchards in the USA. M.
occidentalis is marketed commercially as a biological control agent and is recog-
nized having a potentially worldwide role in integrated spider mite control pro-
grammes. Well-fed mites do not show aerial dispersal behaviour indicating that
food availability may be a component in stimulating aerial dispersal. M. occidenta-
lis have been genetically manipulated through artificial selection to produce pesti-
cide resistant or nondiapausing strains. Like many arthropods, this predatory mite
contains multiple genomes including genomes of several microbial symbionts as
well as its own mitochondrial and nuclear genomes. M. occidentalis can reduce pest
mite populations at low densities over long periods of time because it has a rapid
development time (about one week at 25 °C) and a high reproductive rate (20–30
mites/female). M. occidentalis is commercially mass reared and released to control
spider mites on strawberries in the USA. The mite has also been introduced into
Australia and New Zealand to control pest mites on apple and peach orchards in a
classical biological control programmes.
Amblyseius swirskii This mite has been widely used to control thrips and whiteflies
(which serve as carriers of broad mites on their bodies) in sweet pepper crop in
Israel and Spain.
Amblyseius tetranychivorus Amblyseius tetranychivorus is also reported to keep

the spider mites at low populations on several crops in India.
20.1.1.2 Stigmaeidae
Stigmaeid mites are red to yellow, ovoid or elongate mites whose dorsum is covered
with several plates. These mites live in the soil and on plants and are usually preda-
tors of different species of mites; some stigmaeid mites have commonly been
reported feeding on eriophyid, tetranychid and tenuipalpid mites. Two genera,
namely, Agistemus and Zetzellia, are commonly found on many plant species.
Zetzellia mali is reported feeding on European red mite (ERM) and apple rust mites
in many orchards and contributes towards biological control of mites. But it has a
drawback of feeding on the eggs of the phytoseiid mites, M. occidentalis and
Typhlodromus pyri. Z. mali feeds mainly on eggs and immature stages of ERM and
on active stages of apple rust mite. Eggs of Z. mali are round, lemon yellow but
turns reddish yellow after feeding on ERM. These mites are almost oval in outline
but are more pointed at the rear end and are slightly smaller than the spider mites.
Females overwinter in cracks and crevices on twigs or bark or under empty scales
of San Jose scales. Z. mali can be easily seen and monitored by sampling leaves.
However, no detailed studies on their predator prey ratio have been reported. The
mite is mainly found in large numbers in unsprayed orchards and is also reported
resistant to some organophosphate compounds but, unlike M. occidentalis and T.
pyri, is susceptible to endosulfan. Agistemus longisetus on apple, A. exsertus on
tomato russet mites and Aculus schlechtendali have also been commonly reported
feeding on their preys.
20.1.1.3 Anystidae
Members of this family are fast runners. They are large, reddish soft-bodied mites
and prey on some pest mites and small insects. Anystis baccarum has been reported
feeding on a wide variety of small insects and mites (ER1) and is commonly found
on blackcurrants, cereal crops, apple and other fruit orchards associated with erio-
phyid and spider mite pests. Their peak population is found during spring after
coming out of their overwintering sites. Females lay eggs in batches of 20–24 under
the bark on the trunks of the trees or in the soil surrounding the base under field
conditions and are reported to play an important role in controlling apple rust mites.
Anystis agilis have been reported feeding on crawlers of purple scale, Lepidosaphes
beckii, in Florida. A. agilis consumed an average of 870 females of pest mite
throughout its life. A. salicinus has been introduced from southern France into
Australia to control red-legged earth mite, Halotydeus destructor.
20.1.1.4 Cunaxidae
Members of these mites are quite fast runners and may be red, yellow or brown in
appearance. They are indiscriminate feeders on small insects and mites on diverse
crops and in many other habitats. Cunaxa capreolus is commonly reported as preda-
tor of oriental spider mite, Eutetranychus orientalis. At 30 °C one generation is
completed in about a month. A female mite is reported to consume about 470 spider
mites in its life. Cunaxoides olivieri is observed in some studies to feed on gall mite,
Caleptrimerus vitis, on grapevines.
20.1.1.5 Tydeidae
They are small, soft-bodied fast-moving mites found commonly on plants and in the
soil. Homeopronematus anconai is reported as a good predator of tomato russet
mite, Aculus lycopersici. Lorryia formosa is also reported to feed on the soft scales,
Saissetia oleae, and on honeydew produced by scales in citrus groves.
20.1.1.6 Cheyletidae
Cheyletidae are slow-moving white-, yellow- or orange-coloured mites having a
prominent claw complex with palp tarsus carrying sickle and or comb-like setae, as
in Cheyletus eruditus and Cheletogenes ornatus. Hemicheyletia bakeri and
Cheletogenes ornatus have also been reported feeding on spider mites and other
pests.
20.1.1.7 Erythraeidae
They are large reddish mites having two pairs of prodorsal sensilla, many dorsal
setae and thumb–claw process with long straight chelae. The larvae are parasites of
some arthropods, and the nymphs and adults are predaceous on small arthropods
and on eggs of European red mite, Panonychus ulmi. Many Balaustium spp. have
been reported feeding on scale insects and immature stages of mites.
20.1.1.8 Bdellidae
Mouthparts of these mites are snout-like carrying long palpi which terminate with
strong setae but lack the palpal thumb–claw complex. The prodorsum has two
prominent sensilla. They are medium to large mites (0.5–3.5 mm) with finely stri-
ated integument. Bdella depressa is reported as a voracious feeder of some spider
mites and springtails. B. longicornis is reported feeding on spider mites infesting
grapevines. Bdellodes lapidaria is found as an effective predator of lucerne flea,
Sminthurus viridis. Besides some other species have been reported feeding on scale
insects.
20.1.2 Insects Predaceous on Mites
Insects found predaceous on mite pests of plants have often been reported as macro-
predators (ER12).
20.1.2.1 Stethorus spp. (Coleoptera, Coccinellidae)

They are small beetles (1.0–1.5 mm long) with black, brown or yellow appendices
and are present in many ecosystems including rainforests, dry savannas, orchards
and various crops (ER10). They feed both as larvae and adults and are specialized
predators of spider mites and false spider mites. Stethorus indicus, S. bifidus, S.
japonicus, S. gilvifrons, S. pauperculus and S. tridens have been reported commonly
feeding on different spider mites. Several species have been reported as effective
biological control agents in IPM systems to control spider mites on peppers and
cucumbers in greenhouses where temperature ranges from 20 to 30 °C (Biddinger
et al. 2009).
20.1.2.2 Oligota spp. (Coleoptera, Staphylinidae)

Most of these beetles live in decaying plants or in fungi under the bark of trees
where they prey upon mites and small insects. Both larvae and adults prey upon
spider mites. The adults are small (1–2 mm long) elongated insects generally black
or brown with the abdomen projecting from the short elytra. The development time
varies from 21 to 30 days and the longevity is 30–35 days in different species. A
female beetle in its lifetime lays from 40 to 300 eggs. During development a larva
feeds up to 20 mites/day, while an adult beetle may feed up to 10 mites/day. Oligota
oviformis has been found to play quite beneficial role in controlling spider mites in
orchards. O. flavicornis is also reported preying on spider mite, Panonychus ulmi.
O. pygmaea is considered as an important predator of tea mite, Oligonychus cof-
feae, in India.
20.1.2.3 Thrips (Thysanoptera, Thripidae)

Thrips (0.2–5 mm long) are elongated and have two pairs of wings that are held over
the back at rest, and many species are plant feeders or feed on pollen or fungi. Some
species of thrips prey on spider mites or other small arthropods such as thrips, coc-
cids and whiteflies. Representatives of the families Aeolothripidae, Phlaeothripidae
and Thripidae have been reported feeding on spider mites in different agroecosys-
tems. Scolothrips indicus and S. sexmaculaltus (both Thripidae) and Aeolothrips
fauri and Leptothrips mali (both Phlaeothripidae) have been reported as acaropha-
gous. They consume eggs and other stages of mites. The developmental period var-
ied from 8 to 26 days, fecundity varied from 12 to 220 eggs/female and longevity of
female varied from 21 to 49 days (Gilstrap and Oatman 1976).
20.1.2.4 Other Mite Predaceous Insects

Some species of heteropterous bugs belonging to families Anthocoridae, Miridae
and Lygaeidae are also reported as facultative predators of spider mites. Anthocoris
nemorum, Orius longisetosus and Blepharidopterous angulatus have been reported
feeding on different spider mites in different parts of the world. Gall midges
(Diptera, Cecidomyiidae) are also reported as effective natural enemy of some spi-
der mites (Gerson 2014).
Considering requirements for effective predation of phytophagous mites, none of
the micro- and macro-predators meet the requirements like shorter developmental
period, high reproductive potential, effective searching capacity at low densities and
same microhabitat preference as that of their prey, seasonal cycle corresponding
with that of their prey, ability to tolerate weather extremes as well as their prey and
ability to tolerate pesticides. However, many phytoseiid mites have comparatively
shorter life cycle than their prey, equivalent reproductive potentials as of their prey
and good searching capacity and also have ability to survive on relatively few prey
and thus are comparatively more effective predators and hold better promise in
management of several phytophagous mites in both greenhouses and under field
conditions. Relative importance of different spider mite predators is, however, clas-
sified tentatively in the following order of descending efficiency: Phytoseiidae
mites, Stethorus spp., thrips and Stigmaeidae mites, Hemiptera and Neuroptera.
20.1.3 Microbial Control of Phytophagous Mites
Pathogens in populations of phytophagous mite species often play a major role in

the regulation of population size and population density in natural habitats. As com-
pared to pathogens of insect pests, knowledge regarding acaropathogens is quite
fragmentary. However, in some situations the populations of mite pests in different
agroecosystems are kept below economic threshold level by the occurrence of a
disease (Bruin and Geest 2009; Bugeme et al. 2008; Fernando et al. 2007).
Sometimes occurrence of acaropathogens in the case of mass rearing of beneficial
mites may be undesirable. Information available regarding different groups of
microorganisms that may cause diseases in phytophagous mites is briefly given
below.
20.1.3.1 Viral Diseases

On citrus red mites (Panonychus citri, CRM), virus diseases in citrus groves in
California and Arizona have been reported by some researchers, but observations
revealed that application of field-collected material of virus-infected CRM showed
better results than if virus suspensions obtained from infected laboratory-cultured
mites. However, promising results reported in small-scale applications only as large
field applications proved less satisfactory. High field temperatures seem to have a
negative effect on the virulence of the pathogen, but the applications of viruses may
prove successful in the future because of their narrow host spectrum as many times
phytoseiid mites are comparatively less infected by the virus. Virus-infected mite of
CRM can be recognized by a darker colour in case of immature stages, but infected
adult mites did not show any colour difference from uninfected adult mites. The
most conspicuous symptom, however, is the presence of birefringent bodies inside
the midgut of mites. These crystal-like structures probably contain guanine. The
virus may cause epizootics in natural populations, but only at high densities of the
spider mites. In case of predatory mite, Metaseiulus occidentalis diseased females
in laboratory studies had a reduced oviposition and often died suddenly with a para-
lyzed appearance. In the alimentary canal of M. occidentalis, different types of virus
particles (of approximately 47 nm, 38 nm and 35 nm in diameter) have been
reported, and these particles seem similar to those reported earlier in the CRM.
20.1.3.2 Bacterial Diseases

In some studies field applications of thuringiensin (β-exotoxin) obtained from
Bacillus thuringiensis reported successful against CRM and Tetranychus pacificus.
Thuringiensin has also been reported as a potential acaricide for T. urticae in some
studies. But some studies also report reduced oviposition in both P. persimilis and T.
urticae after thuringiensin application. So both the virus and P. persimilis applica-
tion cannot be used in combination.
Molecular studies revealed the discovery of Wolbachia in tetranychid as well as
phytoseiid mites. Probably all rickettsiae-like organisms reported associated with
mites in earlier studies are members of the genus Wolbachia belonging to the family
Rickettsiaceae (Breeuwer 1997; Breeuwer and Jacob 1996; Hoerauf and Roy 2007).
These bacteria are present in the reproductive system of mites and insects and cause
many unusual effects in their host such as parthenogenesis (infected females pro-
duce only females), male killing (male embryos die, while female embryos develop
into adults), feminization (infected male embryos develop into females) and cyto-
plasmic incompatibility.
Wolbachia is a bacterium that is vertically transmitted to the next generation
through the females. It modifies host reproduction in several ways in order to
enhance its own spread. This bacterium also induces parthenogenesis in the spider
mite, Bryobia praetiosa, causing males not to be produced. This is to the advantage
of the bacterium, as only females are able to transmit the pathogen to the next gen-
eration. Heat treatment (32 °C) can eliminate Wolbachia from infected mites.
20.1.3.3 Diseases Caused by Fungi

In the fungus phylum, only the order Entomophthorales (meaning insect destroyers)
(class Zygomycetes) contains species that are pathogenic for Arthropoda. These
fungi are mainly obligatory pathogens of insects and many other arthropods, often
with a restricted host spectrum. Entomophthoralean fungi usually penetrate their
host through the cuticle by means of a germ tube that is formed by the conidium
(Gerson et al. 2008; Kenneth et al. 1979; Kumar and Singh 2000). Mite-infecting
species are mainly found in the family Neozygitaceae. After penetration, mycelium
is formed within the body cavity of the host that fragments into a large number of
smaller segments, the hyphal bodies; hyphal bodies are the propagative unit and
multiply by budding and by undergoing fusion. The infected host remains fixed to
the substrate, even after its death. Favourable conditions (high relative humidity)
allow the formation of conidiophores that grow through the cuticle of the host
(Meitkowski et al. 1993). Most entomophthoralean fungi are difficult to culture in
artificial culture media. For mass production, it is often necessary to use live host.
Neozygites tanajoa is specific to cassava green mite, Mononychellus tanajoa; N.
floridana also has restricted host spectrum and is considered useful against spider
mite, T. tumidus, in cotton in the humid subtropical part of the Florida.
Prospectus of Acaropathogens in Mite Pest Management Several cases regard-

ing decimations of population of spider mites by acaropathogens under natural con-
ditions are reported. However, viruses could not be successfully applied against
mite pests as their mass production is very laborious and expensive (Van der Gueest
and Bruin 2008). Moreover, the virus becomes rapidly inactivated by sunlight when
applied in aqueous formulations during high temperature (common in citrus
orchards). Large epidemics of fungal diseases in natural populations of spider mites
and eriophyid mites have been reported frequently, but the major drawback is that
all fungi require a relative humidity near the saturation point for both the spore ger-
mination and spore formation. Fungal pathogen, N. floridana, in some conditions
has been reportedly causing large-scale reduction of population of several species of
spider mite. But such epidemics usually occur later in the season (periods when
relative humidity is near the saturation point) and damage to the crop has already
been inflicted. However, entomophthoralean species are very fastidious; culturing
and sporulation in artificial media are hardly possible which makes mass produc-
tions of the fungi very laborious and expensive because the fungus is grown on liv-
ing mites. Even the infective stages of these fungi are rather short-lived and this
characteristic makes their application difficult. Fungal applications may show good
prospects under greenhouse conditions. Some acaropathogens with active ingredi-
ent Beauveria bassiana, Lecanicillium sp., Metarhizium anisopliae and Hirsutella
spp. have been used from time to time. But only Hirsutella thompsonii have been
reported having high specificity towards mites. H. thompsonii var. synnematosa and
H. thompsonii var. inacea have been introduced as classical biological control
agents in Argentina for the control of Eriophyes sheldoni and Phyllocoptruta oleiv-
ora infesting citrus, but no information about their persistence is available (McCoy
1996).
20.1.4 Biological Control of Weeds by Phytophagous Mites
Weeds are the unwanted plants in fields, in gardens and in open places. They com-
pete with main crops for nutrients, moisture and sunlight and can decrease the crop
quality, raise the production costs due to increased cultivation and hand weeding
and also considerably reduce the crop yields. Weeds also serve as the alternate hosts
for several insects and mite pests and diseases. Most of the worlds’ worst weeds
have been introduced accidentally from other countries leaving behind their natural
enemies. Besides several insects damaging weed plants, some mite species belong-
ing to Acariformes families Tetranychidae (spider mites) and Eriophyidae (worm
mites) have also been reported damaging different weeds and thereby help in their
management to some extent (Cromroy 1983; Coombs et al. 2004; Julien 2001a, b).
20.1.4.1 Role of Spider Mites (Family Tetranychidae)
Prickly Pear
Prickly pear is the general term used to describe cacti of Opuntia species (family
Cactaceae). The term prickly pear is derived from the plants’ spiny and pear-shaped
fruit. The different studies on releases of the prickly pear red spider mite, Tetranychus
desertorum (earlier known as T. opuntiae), were found as an important factor in
controlling prickly pear (Opuntia inermis). Control of prickly pear by Cactoblastis
moth has been reported to be monumental in many countries; however, Cactoblastis
larvae do not thrive in dry conditions (heated houses) or on plants growing in poor
soils and are not effective against all species of prickly pear. But red spider mite, T.
desertorum, thrives in hot and dry conditions on prickly pear species found in dry
areas. Mite infestation results in leaf drop and can cause plant death during high
infestations. Heavy mite infestation restricted fruit formation on the prickly pear
plants (Cordo and Deloach 1976).
T. desertorum is found highly host specific on prickly pear from Southern USA
and Mexico, was introduced in 1922 into Queensland (Australia) for control of
cactus and is still established in this region (declared class 1 pest plant in Queensland).
The mites are reported to decrease by 75% in Opuntia inermis (1.5 metre tall) in 2
years in some studies. Cactoblastis cactorum is invariably credited with a success-
ful control of O. inermis in Australia; T. opuntiae seems to have made a valuable
contribution to the biological control of this weed.
Gorse (Ulex europaeus)

Gorse is a woody leguminous shrub which is commonly found in some parts of
Europe. The weed has established in many other temperate parts of the world and is
a major weed of exotic forests and rangeland in New Zealand, Australia and the
USA. Spider mite, Tetranychus lintearius, is the most damaging species which
attacks gorse in Europe, and its potential as biological control of gorse needs further
investigation. T. lintearius is a colonial, multivoltine species and overwinters as
active females. Mite webbing can completely envelop heavily infested plants. As a
result of continuous mite feeding, severe bronzing of the foliage may occur and
sometimes the death of plants occurs. Females have longer adult period and con-
tinue to reproduce throughout the winter months and seem to have the ability to
cause severe damage to the gorse plants. T. lintearius has been reported only on
Ulex species.
20.1.4.2 Role of Galumnid Mites (Family Galumnidae)
Water Hyacinth (Eichhornia crassipes)

Water hyacinths are problematic aquatic weeds because they interfere with the use
of water by human beings and disrupt native fauna and aquatic flora. Water hya-
cinths are the most damaging aquatic weeds in many parts of the world especially
in tropical regions. Orthogalumna terebrantis has been reported in many cases to
provide good control of water hyacinth. The female mite lays its eggs in small holes
in the surface of leaves and the incubation period lasts 7–8 days. The development
of larvae and nymphs requires about 15 days. Nymphs of the mite form galleries
between the parallel veins of the laminae from which adults emerge. About 200,
000 galleries per square metre (or 10, 000 per plant) caused serious damage to water
hyacinth in the field, but the damage was sporadic and confined to small areas of
plants in midsummer months. From two to three generations of the mite a year
occurred in the field. This mite has many times been reported infesting water hya-
cinth in Argentina, Brazil, Guyana, Jamaica, Zambia and the USA. High popula-
tions of the mite cause leaf discolouration and desiccation. Orthogaluma was first
released in Zambia in 1971 and in India during 1986. Although infestations of dif-
ferent intensities of O. terebrantis on water hyacinth are reported from time to time,
the mite has not been reported to contribute significantly to control water hyacinth;
however, mite releases are reported to establish in Africa and the USA, and the
mites may contribute to control in the presence of other factors.
20.1.4.3 Role of Eriophyid Mites

In some situations, infestation of eriophyid mites can be utilized for the control of
different weeds as they are in many cases host specific (Boczek 1995; Rosenthal
1996; Smith et al. 2010).
Field Bindweed (Convolvulus arvensis, Convolvulaceae)

Bindweed is one of the most widespread and difficult weeds to manage. Vining
plants of bindweed thrive in the arid areas where other plants cannot exist. They are
climbing or creeping herbaceous perennial plants. The bindweed mite, Aceria mal-
herbae, is reported to infest and serve as a biological control agent for field bind-
weed. Mite causes the formation of gall-like growth on leaves; leaves become
thickened and have a fuzzy texture in heavily infested plants. Shoots of infested
plants are mis-shapened and the growth is severely stunted. Newly emerged leaves
appear folded with thickened midribs. Thickened texture and fuzzy appearance of
leaves are good diagnostic characters of the presence of bindweed mite. Eriophyid
mites have the potential to aid in suppression or control of bindweed in many arid
regions. Aceria convolvuli infestation results in curling and distortion of bindweed.
Flowers are not found on mite-infested stems. Thus, mite has many attributes of a
good natural enemy: the mite infestation is very damaging to the weed and has sea-
sonal synchrony with that of the host, breeds and spread rapidly by wind.
Skeleton Weed (Chondrilla juncea, Asteraceae)

Also known as devil’s grass, this is a perennial composite belonging to family
Asteraceae, native to Mediterranean and Central Asia. The weed is a herbaceous
perennial with a deep taproot and interferes with wheat plants regarding nutrients
and moisture and also interferes in using harvesting machinery. Eriophyid mite,
Aceria chondrillae, seriously infests the skeleton weed and destroys flower buds,
suppresses seed production, weakens the plant through gall formation and stunts the
plant by blocking apical development. A Greek strain of the mite was introduced in
1971 into Australia to control Chondrilla weed. Mites are also introduced from Italy
to the USA in 1977. The mites readily attacked the weed but relative humidity of
30% is essential to achieve mite infestation.
Russian Knapweed (Acroptilon repens, Asteraceae)

Russian knapweed belongs to family Asteraceae and is a persistent and perennial
weed. Eriophyid mite, Aceria acroptiloni, attacks flowers and sterilizes A. repens.
Shoots with infested inflorescences are underdeveloped and no more new shoots
form on them. Heavy mite infestation stunts the whole plant and greatly reduces its
foliage.
Other Weeds
Diffuse Knapweed (Centaurea diffusa) Eriophyid mites, Aceria centaurea and A.
thessaloneae, cause pocket-like galls on leaves and the affected weed plant looks
broom-like. This weed is of great economic importance in North America.
Hoary Cress (Cardaria draba, Brassicaceae) Eriophyid mite, Aceria drabae,

attacks the flower head and infested plants do not produce seeds. The weed is very
important in Europe, Asia and America.
Lantana (Lantana camara, Verbenaceae) Lantana is a heavily branched shrub

that grows in compact clumps, dense thickets or as a climbing vine. Eriophyid mite,
Aceria lantanae, is reported infesting lantana plants resulting in disruption of seed
production in Florida.
Poison Ivy (Toxicodendron radicans, Sapindaceae) Infestation of Acullops toxi-

cophagous causes leaf galling and stunted growth of poison ivy in Florida and
Bermuda.
Multiflora Rose (Rosa multiflora, Rosaceae) Eriophyid mite, Phyllocoptes fruc-

tiphilus, transmits virus disease on multiflora rose.
St. John’s Wart (Hypericum perforatum, Hypericaceae) Eriophyid mite has

been reported highly host specific and useful biological control agent of this weed.
Bermuda Grass (Cynodon dactylon, Poaceae) Bermuda grass is commonly

found in many parts of the world in crops, vineyards and plantation crop. Eriophyid
mite, Aceria cynodoniensis, is an important biological control agent of this weed.
Besides the above few examples of eriophyid mites as good control agents suit-
able for classical biological control of weeds, many other stray examples of occur-
rence of other eriophyid mites like A. tamaricis have been considered to have a high
potential for use as classical biological control agents of weeds because they achieve
high population densities and have short generation times, high values of intrinsic
rates of increase and high aerial mobility. However, a detailed life history, thorough
host specificity and evolutionary stability of eriophyid mites need more detailed
investigations.
20.2 Conclusions
Many spider mites (Tetranychidae), false spider mites (Tenuipalpidae), worm mites
(Eriophyidae) and bulb mites (Acaridae) are reported naturally controlled by differ-
ent predaceous mites, mite predaceous insects and acaropathogens (including
viruses, fungi, bacteria, etc.). Predaceous mites (Phytoseiidae, Stigmaeidae,
Cunaxidae and Erythraeidae) are commonly found feeding on different phytopha-
gous mites. Representatives of family Phytoseiidae (Phytoseiulus persimilis) is
known as fast action predator and are commonly used for control and management
of various spider mites under both greenhouse conditions and open field conditions.
Metaseiulus occidentalis, Neoseiulus californicus and Amblyseius fallacis are other
most utilized phytoseiid predators (Schutte and Dicke 2008). Some species of
Stigmaeidae, Tydeidae, Cheyletidae and Erythraeidae families are also reported
actively feeding on spider mites in some situations. Besides predaceous mites, some
representatives of insect families Coccinellidae, Staphylinidae, Thripidae and
Cecidomyiidae are also reported as successful mite predators. Besides different
predators, some microbial agents like viruses, bacteria and fungal agents are also
known to provide good control, but they have problems in mass multiplication
because of their limitation to live only on live mites. Fungal species belonging to
order Entomophthorales and family Neozygitaceae infect mainly mites. But acaro-
fungus species are difficult to culture in artificial culture media and also require a
relative humidity near the saturation point for both spore germination and spore
formation.
Weeds are unwanted plants in fields, in gardens and in open places; they compete
with main crops for moisture and nutrients, and even some act as alternate hosts for
several insects, mite pests and diseases. Besides different insect pests, some of spi-
der mites and worm mites are found selectively feeding and breeding on important
weed plants. Against prickly pear (Opuntia inermis), spider mite Tetranychus deser-
torum is tried with limited success. T. desertorum thrives under hot and dry condi-
tions on prickly pear, and mite infestation results in leaf drop and can cause plant
death in high infestations and restricted fruit formation on the prickly pear plants.
Galumnid mite, Orthogalumna terebrantis, has also been reported in many cases to
provide control of water hyacinth (Eichhornia crassipes). Besides spider mites,
some eriophyid mites like Aceria malherbae against weed (Convolvulus arvensis),
Aceria chondrillae against skeleton weed (Chondrilla juncea) and Phyllocoptes
fructiphilus which transmit viral disease and control multiflora rose (Rosa multi-
flora) have been tried with some success. Besides the above examples of weed con-
trol, some eriophyid mites have been reported successfully against some other
weeds.
Further Reading
Albajes R, Gullino ML, Van Lantern JC, Elad Y (eds) (1999) Integrated, pest and disease manage-
ment. Kluwer Academic Publishers, Dordrecht
Further Reading 395
Biddinger DJ, Weber DD, Hull LA (2009) Coccinellidae as predators of mites: Stethorini in bio-
logical control. Biol Control 51:268–283
Bjornson SE (2008) Natural enemies of mass-reared predatory mites (family Phytoseiidae) used
for biological control. Exp Appl Acarol 46:299–306
Boczek J (1995) Eriophyid mites (Acari: Eriophyoidea) as agents of biological weed control. In:
The Acari (physiological and ecological aspects of Acari-host relationships). DABOR,
Warszava
Breeuwer JA (1997) Wolbachia and cytoplasmic incompatibility in the spider mites, Tetranychus
urticae and T. turkestani. Heredity 79:41–47
Breeuwer JA, Jacobs G (1996) Wolbachia: intracellular manipulators of mite reproduction. Exp
Bruin J, van der Geest LPS (eds) (2009) Diseases of mites and ticks. Exp Appl Acarol 46:1–4,
2009, VI, 350 pp. Hardcover. ISBN: 978-1-4020-9694-5
Bugeme DM, Maniania NK, Knapp M, Boga JJ (2008) Effect of temperature on the virulence of
Beauveria bassiana and Metarhizium anisopliae isolates to Tetranychus evansi. Exp Appl
Acarol 46:275–285
Castillo RC, Verencio R, Narita JPZ (2015) Prospects for biological control of plant feeding mites
and other harmful organisms, Progress in biological control 19. Springer, Cham.
doi:10.1007/978-3-319-15042-0_1
Coombs EM, Clark JK, Piper GL, Confrancesco AF (2004) Biological control of invasive plants in
the United States. 448 pages, OSU Press, Oregon State, Corvallis. ISBN 978-0-87071-029-2
Cordo HA, Deloach CJ (1975) Ovipositional specificity and feeding habits of the water hyacinth
mite, Orthogalumna terebrantis in Argentina. Environ Entomol 4:561–562
Cordo HA, Deloach CJ (1976) Biology of the water hyacinth mite in Argentina. Weed Sci
24:245–249
Cromroy HL (1983) Potential use of mites in biological control of terrestrial and aquatic weeds.
In: Hoy MA, Knutson L, Cunningham GL (eds) Biological control of pests by mites, Special
publication 3304. University of California, Agricultural Experiment Station, Berkeley,
pp 61–66
Dalrymple DD (1993) Controlled environment agriculture. A global review of greenhouse food
production, foreign agriculture economic report no. 89. Economic Research Service, USDA,
Washington, DC
Del Fosse ES, Cromroy HL, Habeck DH (1978) The potential use of eriophyoid mites for control
of weeds. In: Freeman TE (ed) Procceeding of IV international symposium on biological con-
trol of weeds, August 30–September 2, 1976 University of Florida, Gainesville, Florida, USA,
pp 473–475
Fernando LCP, Manoj P, Hapuarachchi DCL, Edgington S (2007) Evaluation of four isolates of
Hirsutella thompsonii against coconut mite (Aceria guerreronis) in Sri Lanka. Crop Prot
26:1062–1066
Gerson U (2014) Pest control by mites (Acari): present and future. Acarologia 54:371–394
Gerson U, Gafni A, Paz Z, Sztejnberg A (2008) A tale of three acaropathogenic fungi in Israel:
Hirsutella, Meira and Acaromyces. Exp Appl Acarol 46:183–194
Gilstrap FE, Oatman ER (1976) The bionomics of Scolothrips sexmaculatus (Pergande)
(Thysanoptera: Thripidae), an insect predator of spider mites. Hilgardia 44:1–59
Hoerauf A, Rao RV (2007) Wolbachia: a bug’s life in another bug, Issues in Infectious diseases, vol
V. Karger Publishers, Basel, 150 pp
Hoy MA, Jeyaprakash A (2008) Symbionts including pathogens, of the predatory mite Metaseiulus
occidentalis: current and future analysis. Exp Appl Acarol 46:329–346
Julien MH (2001a) Biological control of water Hyacinth with arthropods: a review up to (2000).
In: Julian MH, Hill MP, Center TD, Jianqing D (eds) Biological and integrated control of water
Hyacinth, Eichhornia craccipes. ACIAR Proceedings 102. CSIRO Entomology, 120. Meirs
Road, Indooropilay, Queensland, 4068, Australia, pp 8–20
Julien MH (2001b) Determinations of feeding mechanisms of the water hyacinth. Hyacinth
Control J 13:53–55
Kenneth R, Muttath TI, Gerson U (1979) Hirsutella thompsonii, a fungal pathogen of mites.
I. Biology of the fungus in vitro. Ann Appl Biol 91:21–28
Kumar SP, Singh SP (2000) Hirsutella thompsonii: the best biological control option for the man-
agement of the coconut mite in India. Indian Coconut J 31(5):11–17
McCoy CW (1996) Pathogens of eriophyoids. In: Lindquist EE, Sabelis MW, Bruin J (eds)
Eriophyoid mites-their biology, natural enemies and control. Elsevier, Amsterdam,
pp 481–490
Meitkowski R, Belazy S, Van der Gueest LPS (1993) Observations on a mycoses of spider mites
(Acari: Tetranychidae) caused by Neozygites floridana in Poland. J Invertebr Pathol
61:317–319
Poiner G, Poinar R (1998) Parasites and pathogens of mites. Annu Rev Entomol 43:449–469
Rosenthal SS (1996) Biological control of weeds. In: Lindquist EE, Sabelis MW, Bruin J (eds)
Eriophyoid mites. Their biology, natural enemies and control. Elsevier Science BV, Amsterdam,
pp 729–739
Schutte C, Dicke M (2008) Verified and potential pathogens of predatory mites (Acari:
Phytoseiidae). Exp Appl Acarol 46:307–328
Smith L, deLillo E, Amrine JW (2010) Effectiveness of eriophyid mites for biological control of
weedy plants, challenges for future search. Exp Appl Acarol 51:115–149
Stenseth C (1979) The effect of temperature and humidity on the development of Phytoseiulus
persimilis and its ability to regulate populations of Tetranychus urticae (Acarina: Phytoseiidae,
Tetranychidae). Entomophaga 29:311–317
Vala F, Egas M, Breauwer JAJ, Sabelis MW (2004) Wolbachia affects oviposition and mating
behaviour of its spider mite host. J Evol Biol 17:692–700
Van der Gueest LPS, Bruin J (2008) Diseases of mites and ticks: from basic pathology to microbial
control-an introduction. Exp Appl Acarol 46:3–6. doi:10.1007/s10493-008-9222-2
Van der Gueest LPS, Ellict SL, Breeuwer JAJ, Bearling EAM (2002) Diseases of mites. Exp Appl
Acarol 24:497–569
Walter DE, Proctor HC (2013) Mites: ecology, evolution and behaviour (life at a microscale).
Springer, Dordrecht
Welbourn WC (1983) Potential use of trombidioid and erythraeoid mites as biological control of
insect pests. In: Hoy MA, Cunningham GL, Knutson L (eds) Biological control of mites,
Special publication 3304. University of California, Berkeley, pp 103–140 185pp
Werren JH (1997) Biology of Wolbachia. Annu Rev Entomol 42:587–689
Zhang ZQ, Xin JL (1989) Biology of Allothrombium pulvinum Ewing (Acariformes:
Trombidiformes), a potential biological control agent of aphids in China. Exp Appl Acarol
6:101–108
1. https://www.youtube.com/watch?v=pyClfvcSzLA Anystid Mite

2. https://www.youtube.com/watch?v=qre7VrpIYZg Phytoseiulus persimilis [Persimilis Mite]
Consuming Tetranychus urticae [Two-spotted Spider Mite]
3. https://www.youtube.com/watch?v=gt-nW3m8DWc Amblyseius herbicolus feeding on Broad
mite
4. https://www.youtube.com/watch?v=rAM9dfEtV_k Predatory Mite Neoseiulus Predatory Mite
Takes on Damaging Spider Mites
5. https://www.youtube.com/watch?v=QQ8FMX3yWNU Applying Predatory Mites (Mite
brushing Machine also shown)
7. https://www.youtube.com/watch?v=ZvoS0i3o5BY Predatory Mite Bioassays
8. https://www.youtube.com/watch?v=2mFyBxldl-Q Phytoseiulus vs Spider Mite.wmv
Further Reading 397
9. https://www.youtube.com/watch?v=CBEkaemkpSg Stethorus Larva

11. https://www.youtube.com/watch?v=b0mdhQYgtdo Biological control 1.wmv (Includes all
mite predators)
Pesticides
21
Abstract
Pesticides are important pest management tools. Pests may include insects,
mites, fungi and bacteria which cause plant disease. Until mid-twentieth century,
in agroecosystems of low-level productivity, phytophagous mite populations
usually stayed below economic injury levels due to natural regulation by preda-
tory mites and mite predaceous insects. The concept of secondary pest outbreak
was introduced on spider mites (Tetranychidae), the most important plant-
feeding mites as a paradigm. Widespread use of synthetic organic compounds
against insects as target pests destroyed spider mite predators which are gener-
ally more susceptible to pesticides than their prey. Heavy selection pressure by
neuroactive insecticides caused emergence of spider mite population resistant to
these compounds. Earlier wettable sulphur and petroleum oils were sometimes
used as insecticides/miticides. Petroleum oils are used as ‘dormant oils’ and
‘summer oils’ to control mites, scale insects, pear psylla, mealybugs, etc. and
against eggs of spider mites. Acaricides belonging to chemical groups like
bridged diphenyl (chlorbenside and dicofol), dinitrophenol (binapacryl), organo-
chlorine (endosulfan), organophosphate (triazophos), organotin (cyhexatin),
pyrethroid (flucythrinate), thiourea (diafenthiuron) and mite growth regulators
(like clofentezine) were developed. Insecticide Resistance Action Committee
(IRAC) has classified the insecticides and acaricides as the basis for their effec-
tive sustainable management. Role of acaricides in IPM of spider mites is dis-
cussed. Development of acaricide resistance and measures to counteract their
effect are also included.
Keywords
Pesticides • Acaricides • Resistance management

DOI 10.1007/978-981-10-1594-6_21
400 21 Pesticides
A pesticide is any substance or mixture of substances intended for preventing,

destroying, repelling or mitigating any pest. Pesticides are important pest manage-
ment tools; many pesticides have gradually lost this effectiveness due to the devel-
opment of resistance by pest they have earlier controlled. Pest resistance is a
heritable and significant decrease in the sensitivity of a pest population to a pesti-
cide that is shown to reduce the field performance of pesticides. Pests may include
insects, mites, weeds, fungi and bacteria which cause plant disease (Casida and
Quistad 1998; Croft 1990).
In 1942 Paul Hermann Muller of Switzerland discovered the insecticidal proper-
ties of a synthetic chlorinated organic chemical – dichlorodiphenyl trichloroethane
(DDT) – and in 1948 Muller received the Nobel Prize for Physiology of Medicine
for this discovery. DDT belongs to the chemical class of diphenyl aliphatics and is
a persistent pesticide. Methoxychlor, lindane, toxaphene, aldrin, dieldrin and endrin
are all known as chlorinated insecticides. As DDT and their close relatives lead to
their bioconcentration in the soil, water, birds, mammals and in human beings; and
repeated applications of these different pesticides on crop failures like pest resis-
tance, pest resurgence and development of secondary pests resulted (March 1957).
So there has been a decline in the usage of organochlorines, and these pesticides
have been replaced by the other groups of synthetic organic insecticides like organo-
phosphates, carbamates and synthetic pyrethroids. Particularly for control of spider
mites, inorganic sulphur and petroleum oils were earlier mainly used in the same
situations. However, currently for the insect and mite control, an approach com-
monly known as integrated pest management (IPM) is widely applied, taking care
of safety of different compounds to their natural enemies (Innaaya and Degheele
1998).
21.1 Mite Outbreaks
Until the mid-twentieth century, in agroecosystems of low-level productivity, phy-

tophagous mite populations usually stayed below economic injury levels due to
natural regulation by predatory mites and predaceous insects. The concept of sec-
ondary pest outbreak was introduced on spider mites (Tetranychidae), the most
important plant-feeding mites as a paradigm (Collyer and Kirby 1955; Croft and
Van de Baan 1988; Dhooria and Mann 1988; Husseine 1958; Penman and Chapman
1988). Advances in agricultural production after World War II based on the exten-
sive use of pesticides and fertilizers, irrigation and other cultural practices induced
increase in spider mite populations far above economic threshold. Grown under
favourable conditions, host plants became high-quality food sources for the mite
pests which gave rise to outbreaks of their populations and made it possible to com-
pensate for the losses caused by predators’ activity. Moreover, widespread use of
neuroactive insecticides (synthetic organic compounds) against insects as target
pests, but toxic to other nontarget insect and mite species as well, destroyed spider
mite predators which are generally more susceptible than their prey. On the other
hand, heavy selection pressure by neuroactive insecticides caused emergence of
21.3 Development of Specific Acaricidal Compounds 401
spider mite populations resistant to these compounds. Besides the resistance of spi-
der mites and the elimination of their predators as the primary causes, outbreaks are
influenced by sublethal effects of pesticides on behaviour and physiology of pests
or predators. Spider mites, mostly polyphagous species, are appearing commonly as
pests in modern ecosystems worldwide, and some of them are among the most
important crop pests.
21.2 Resurgence
Resurgence of mite pests, in the present context, is defined as the post-treatment

numerical increase in pest populations far above those occurring in conventionally
treated plots. The term outbreak may or may not have been meant by different
researchers to mean resurgences. Resurgence of spider mites becomes evident from
one week to several months post-treatment. Synthetic pyrethroids vary in their
adverse effects on spider mites and also differ in their ability to invoke resurgences
of different spider mite species on diverse plants. Pyrethroids are lethal as well as
repellent to phytoseiids and other predators that feed or prey on spider mites and
affect different phytophagous competitors. Spider mites are likewise repelled by
synthetic pyrethroids, thus becoming more evenly distributed and less web restricted,
with a resultant increase in fecundity. Due to the use of synthetic pyrethroids, spider
mite development is shortened, sex ratios in favour of females, and even onset of
diapause is delayed. Resurgences of spider mites due to application of synthetic
pyrethroids deltamethrin, fenvalerate and flucythrin have been reported frequently
on beans, cotton, soybean, alfalfa, apple, citrus, grape and pear. Different pyrethroid
pesticides have also been reported to modify the physiology, chemistry and even the
nutritive quality of the host plant resulting in increases in spider mite populations
(Gerson and Cohen 1989; Penman and Chapman 1988).
21.3 Development of Specific Acaricidal Compounds
The use of acaricides has increased substantially over the second half of the twenti-
eth century. Since the first serious and widespread outbreaks of spider mites, during
the 1950s, organophosphorus and other neuroactive insecticides were replaced by
specific acaricides, i.e. compounds exclusively or primarily effective against mites.
Several generations of structurally diverse synthetic acaricides, directed against
various biochemical and physiological targets, have been commercialized until
now. Besides specific acaricides, a number of insecticides with considerable acari-
cidal activity (pyrethroids, avermectins, benzoylureas) have also been used, while
some older neuroactive compounds are still available for the control of plant-feeding
mites. Most of the modern acaricides exert their effects through disruption of respi-
ratory processes and by adversely affecting their growth and development. On the
other hand, various natural bioactive products with acaricidal activity (botanical and
microbial pesticides, essential oils, horticultural spray oils and mycopesticides)
402 21 Pesticides
have become important alternatives to synthetic acaricides (Innaaya and Degheele

1998; Knowles 1975).
21.4 Classification of Acaricides
Pesticides are mainly classified based on their chemical structure, mode of action
against host target and sources of their availability.
21.4.1 Inorganic Acaricides
21.4.1.1 Wettable Sulphur

A good acaricide and fungicidal compound. Most of the mites belonging to families
Eriophyidae and Tenuipalpidae are susceptible to sulphur, lime-sulphur sprays or
simply sulphur dusts. Many tarsonemid mites and spider mites have been reported
economically controlled by sulphur application. After application it releases vapours
that are toxic to many mite species particularly bud mites. Sulphur is cheaper, com-
monly available and may be formulated with many other pesticides. It is relatively
specific and generally does not interfere with many predators, but it has been
reported as phytotoxic to many plants when applied during hot weather.
21.4.1.2 Petroleum Oils

Petroleum oils are sometimes used as insecticide–miticides and also used as herbi-
cides and adjutants. Some of their formulations are applied directly to mites, but
usually oils are formulated with the emulsifiers which permit them to be diluted
with water. Petroleum oils are used as ‘dormant oils’ and ‘summer oils’ to control
mites, scale insects, pear psylla, mealybugs, etc. and against eggs of spider mites.
Petroleum oils are of rather low toxicity; however, no resistance to these oils has
been reported. Little or no hazard to birds and beneficial insects is reported; how-
ever, some formulations may be having toxic effects against bees and fish. Depending
on the distillation process and the mineral oil content, they are classified as light
(67–79 %), medium (40–49 %) and heavy (10–15 %). Summer oils have been exten-
sively applied to deciduous and on citrus fruit trees and ornamental plants to control
spider mites and bud mites. However, dormant oil sprays are used to control over-
wintering pests in deciduous trees and vines. Summer oils are used to control pests
during the growing season. Oils kill mites and their eggs by contact. The toxicity
may be due to suffocation of the pest as oils block spiracles (stigmata in mites). Oils
are sometimes used as a repellent to pests. Once the oil dries, it is no longer toxic to
most natural enemies (Croft 1990; Metcalf 1980).
21.5 Synthetic Organic Acaricides 403
21.5 Synthetic Organic Acaricides
21.5.1 Acaricides Based on Their Chemistry
Chemical group Acaricides

Bridged diphenyl Azobenzene, chlorbenside, dicofol, fenson, fluorbenside, tetradifon
and tetrasul
Carbonate Benomyl, carbofuran, methiocarb, aldicarb, thiofanox
Dinitrophenol Binapacryl, dinobuton, dinocap, DNOC
Formamidine Amitraz, chlordimeform, formetanate
Macrocyclic lactone Abamectin, ivermectin, selamectin, milbemectin
Mite growth regulators Clofentezine, fluazuron, flufenoxuron, hexythiazox
Organochlorine Endosulfan, lindane
Organophosphate Amidithion, azinphosmethyl, carbophenothion, demeton-methyl,
diazinon, dimethoate, ethion, parathion, quinalphos, thiometon,
triazophos
Phosphonate Trichlorphon
Organotin Cyhexatin, fenbutatin oxide
Phthalimide Phosmet
Pyrazole Fenpyroximate, tebufenpyrad, fipronil
Pyrethroid Acrinathrin, bifenthrin, cyhalothrin, fenpropathrin, fenvalerate,
flucythrinate, fluvalinate
Pyrrole Chlorfenapyr
Quaternary ammonium Sanguinarine
Quinoxaline Chinomethionat, thioquinox
Sulfite ester Aramite, propargite
Thiazolidine Hexythiazox
Thiourea Diafenthiuron
Unclassified Crotamiton, cycloprate, etoxazole, fenazaflor, fenazaquin
21.5.2 Acaricide Classification Based on Mode of Action
Insecticide Resistance Action Committee (IRAC) has classified the insecticides and
acaricides based on their mode of action (MOA) as the basis for their effective sus-
tainable management. Acaricides are allocated to specific group based on their tar-
get sites (Knowles 1975; Kunz and Kemp 1994). MOA classification list is reviewed
and re-issued periodically, provides insecticides for use in resistance management
strategies to growers, advisors, extension staff, consultants and crop protection pro-
fessionals with a guide to the selection of acaricides/insecticides in resistance man-
agement programmes. Effective Resistance management of this type preserves the
utility and diversity of available acaricides. IRAC is currently encouraging manu-
facturers of pesticides to indicate the IRAC mode of action, group number and
404 21 Pesticides
description on their labels. Such information would be helpful in assisting pesticide

applications in the selection of acaricides and insecticides for use in resistance man-
agement strategies. Relevant MOA groups are selected as per IRAC Mode of Action
Group, March 2012 (Poster ver 1.67 Based on Most classification ver 7.2) (http://
www.irac-online.org/documents/moa-classification/) (Anonymous 2015):
A. Nerve and Muscle Targets Several current acaricides act on nerve and muscle
targets. Acaricides that act on individual targets in this system are generally having
fast action:
Group 1. Acetylcholinesterase (AChE) Inhibitors Inhibit AChE causing hyper-

excitation. AChE is the enzyme that terminates the action of the excitatory neu-
rotransmitter acetylcholine at nerve synapses. 1A. Carbamates (e.g. methomyl). 1B.
Organophosphates (e.g. primiphosmethyl).
Group 2. GABA-gated chloride channel antagonists These pesticides block the

GABA-activated chloride channel causing hyperexcitation and convulsions. GABA
is the major inhibitory neurotransmitter in insects. 2A. Cyclodiene organochlorines
(e.g. endosulfan).
Group 3. Sodium Channel Modulators Keep sodium channels open, causing

hyperexcitation and, in some cases, nerve block. Sodium channels are involved in
the propagation of action potentials along nerve axons. 3A. Pyrethrins, pyrethroids
(e.g. bifenthrin, halfenprox).
Group 6. Chloride Channel Activators Activate glutamate-gated chloride chan-

nels (Glu Cls) causing paralysis. Glutamate is an important inhibitory neurotrans-
mitter in insects. Avermectin, milbemycins (e.g. abamectin, milbemectin).
Group 19. Octopamine Receptor Agonists Activate octopamine receptors, lead-

ing to hyperexcitation. Formamidines (e.g. amitraz).
B. Acaricides for Which the Mode of Action Is Unknown These compounds are
not classified because there is no sufficient information available on their mode of
action. Benzoximate, bifenazate, chinomethionat, dicofol.
C. Respiration Targets Several acaricides are known to interfere with mitochon-

drial respiration by the electron transport and/or oxidative phosphorylation. These
compounds are generally fast to medium fast acting:
Group 12. Inhibitors of Mitochondrial ATP Synthase Inhibit the enzyme that
synthesizes ATP.
12A. Diafenthiuron. 12B. Organotin insecticides (e.g. azocyclotin, fenbutatin
oxide). 12C. Propargite.
21.6 Development of Specific Acaricides 405
Group 13. Uncouplers of Oxidative Phosphorylation via Disruption of the

Proton Gradient Protonophores that short circuit the mitochondrial proton gradi-
ent so that ATP cannot be synthesized (e.g. chlorfenapyr).
Group 20. Mitochondrial Complex III Electron Transport Inhibitors Inhibit

electron transport complex III, preventing the utilization of energy by cells. 20B.
Acequinocyl. 20C. Fluacrypyrim.
Group 21. Mitochondrial Complex I Electron Transport Inhibitors Inhibit

electron transport complex I, preventing the utilization of energy by cells. 21A.
METI acaricides (fenazaquin, pyridaben and tebufenpyrad).
Group 25. Mitochondrial Complex II Electron Transport Inhibitors Inhibit

electron transport complex II. Energy metabolism. Cyenopyrafen and
cyflumetofen.
D. Growth and Development Targets Insect and mite growth regulators act by
mimicking growth hormones, by directly affecting cuticle formation, or lipid bio-
synthesis. Acaricides that act on this system are usually slow acting. The target
proteins are not always known:
Group 10. Mite Growth Inhibitors Incompletely defined mode of action leading
to growth inhibition. 10 A. Clofentezine, hexythiazox. 10 B. Etoxazole.
Group 15. Inhibitors of Chitin Biosynthesis type O. Incompletely defined mode

of action leading to inhibition of chitin biosynthesis. Benzoylureas (e.g. flucycloxu-
ron, flufenoxuron).
Group 23. Inhibitors of Lipid Synthesis Inhibit acetyl-COA carboxylase, part of

the first stop in lipid biosynthesis. Tetronic and tetramic acid derivatives (e.g.
spirodiclofen).
21.6 Development of Specific Acaricides
The first serious and widespread spider mite outbreaks following applications of
neuroactive insecticides were observed at the end of the 1940s, leading to the initia-
tion of work on the research and development of specific acaricides. These com-
pounds, exclusively are primarily effective against mites, were gradually taking
over the organochlorines, organophosphates and carbamates. Bridged diphenyls
(bromopropylate, chloropropylate, chlorobenzilate, chlorfenethol, dicofol, tetradi-
fon) were the first specific acaricides which established themselves in the market in
the 1950s. During the 1960s and early 1970s, the second generation of structurally
rather different specific acaricides emerged, the most important of which were
406 21 Pesticides
propargite, organotins (cyhexatin, fenbutatin oxide) and formamidines (amitraz,

chlordimeform). Most of the first- and second-generation acaricides are not used
any longer. Specific acaricides of the third generation are represented by mite
growth inhibitors (clofentezine and hexythiazox) commercialized in the first half of
the 1980s. In addition to specific acaricides, several structurally diverse synthetic
acaro-fungicides (dinocap, dinobuton, chinomethionate, dichlofluanid) were intro-
duced. On the other hand, the use of sulphur products (that had been exploited as
acaro-fungicides since nineteenth century) was largely displaced by novel synthetic
compounds (Innaaya and Degheele 1998; Croft 1990; Ashton 2013).
Introduction of specific acaricides reduced the adverse impact on beneficial
insects (predators of insect and mite pests, pollinators) to the minimum; at the same
time, specific acaricides proved to be selective, less toxic to predaceous mites than
phytophagous mites. These acaricides effectively control populations of phytopha-
gous mites resistant to neuroactive compounds since these compounds have differ-
ent mode of action, with targets mostly being outside the nervous system. Moreover,
specific acaricides are far more safer for humans, nontarget organisms and the envi-
ronment in comparison to neuroactive compounds in particular organochlorines that
were almost all severely restricted or banned in developed countries in the 1970s.
21.6.1 New Synthetic Acaricides
21.6.1.1 Acaricides Acting on Respiration Targets

In the period between 1991 and 1993, four compounds from different chemical
classes were successively commercialized: fenpyroximate (pyrazole), pyridaben
(pyridazinone), fenazaquin (quinazoline) and tebufenpyrad (pyrazolecarboxamide)
whose mode of action was inhibition of mitochondrial electron transport (MET) at
complex I. These compounds, also known as METI acaricides, quickly gained the
popularity worldwide owing to their high efficiency against both tetranychid and
eriophyid mites, quick knock-down effect and long-lasting impact. In addition,
these substances have low to moderate mammalian toxicity and short to moderate
environmental persistence (Ashton 2013; Mathews 2016).
21.6.1.2 Acaricides Acting on Growth and Developmental Stages

Etoxazole, an oxazoline compound, is an acaricide highly effective against eggs and
immatures of spider mites, nontoxic to adults, but it considerably reduces fertility of
treated females. This insecticide is usually classified among mite growth inhibitors,
together with clofentezine and hexythiazox – older acaricides that cause similar
symptoms. Discovery of spirodiclofen and spiromesifen, tetronic acid derivatives,
broadened the biochemical diversity of acaricides by introducing a completely new
mode of action. These compounds act as inhibitors of acetyl-COA carboxylase, a
key enzyme in fatty acid biosynthesis. Spirodiclofen and spiromesifen are highly
toxic to eggs and immatures of spider mites, while the effects on adult females are
slower with reduced fecundity. Their acaricidal effect is long lasting and stable.
Spirotetramat, a tetramic acid derivate recently introduced, belongs to inhibitors of
21.7 Role of Acaricides in IPM of Spider Mites 407
acetyl-COA carboxylase, although initially developed for control of whiteflies and

aphids, also found as an effective acaricide against spider mites, T. urticae.
21.6.2 Natural Acaricides and Other Alternative Solutions
Azadirachtin The major active ingredient of extracts, oils and other products
derived from the seeds of the Indian neem tree (Azadirachta indica) is reported to
be toxic to various developmental stages. It also acts as antifeedant, reduces fecun-
dity and fertility and shortens the life span of spider mites. Azadirachtin has also
been reported to have effects against some acarid and tarsonemid mites.
Milbemectin A mixture of milbemectin A3 and milbemectin A4 has been isolated

from the fermentations of Streptomyces hygroscopicus subsp. aureolacrimosus.
Milbemectin is a neuroactive acaricide (chloride channel activator), effective against
tetranychid and eriophyid mites.
The most recent example is ‘spinosad’, a mixture of spinosyn A and spinosyn D,

secondary metabolites of Saccharopolyspora spinosa, introduced in 1997 as neuro-
active insecticide, nicotinic acetylcholine receptor agonist. These insecticides also
have significant acaricidal effect. Besides some essential oils, extracted from cara-
way seeds, eucalyptus mint, rosemary, basil, thyme and other plants have shown a
significant acaricidal activity. These oils could be useful as fumigants in the control
of phytophagous mites in greenhouses (Ashton 2013, Mathews 2016).
21.7 Role of Acaricides in IPM of Spider Mites
Biological control of phytophagous mites by predatory mites particularly of family

Phytoseiidae and other predators proved to be a successful alternative to conven-
tional chemical control, especially on greenhouse crops. In modern crop protection,
acaricides should be used as biorational compounds: highly effective against mite
pests and relatively safer to their predators (i.e. selective), with low risk to human
health and the environment. Biorational acaricides are important element of IPM of
spider mites. So it is very important to know the effects of acaricides and other pes-
ticides on phytoseiid mites. Besides lethal effects (mortality), pesticides also cause
a variety of sublethal effects, by changing the biological parameters and/or behav-
iour of survivors (Chandler et al. 2000; Chapman and Martin 2005).
Organochlorines, organophosphates, carbamates, pyrethroids and other broad-
spectrum insecto-acaricides are toxic to a large number of insect and mite species,
including phytoseiid mites and majority of other arthropods. However, abamectin
and milbemectin, which are also broad-spectrum insecto-acaricides, are considered
safer to beneficial arthropods under field conditions due to their short environmental
persistence, rapid uptake into treated plants and fast degradation of surface residues.
408 21 Pesticides
Specific acaricides are considered harmless to majority of predatory insects. So

from the standpoint of selectivity, it is essential to be aware of the comparative tox-
icity of acaricides to phytophagous mites and predatory mites. Compounds like
organotins, mite growth inhibitors and regulators, diafenthiuron, some METI acari-
cides, bifenazate, spirodiclofen and spiromesifen are usually graded as selective
acaricides. Spinosad and azadirachtin seem very much compatible with predatory
mites. Non-selective compounds can be made safer for use by special application
technology by releasing the predators so that they would be exposed to older resi-
dues by using the strain of predators with developed resistance to acaricides and
other predators. So transition from conventional pest control to IPM actually
changes the role of pesticides (acaricides) in modern crop protection: within the
principles of IPM, pesticides are applied highly rationally and in interaction with
other control tactics.
21.8 Acaricide Resistance
Resistance refers to an inheritable change in the sensitivity of a pest population that

is reflected in the repeated failure of a product to achieve the expected level of con-
trol when used according to the label recommendation for that species. Resistance
does not always occur, but resistance has been reported to many acaricides and
insecticides. Complicating the understanding and management of resistance is the
problem of knowing which type of resistance is present in a given pest population.
For example, some pest populations are known to have cross-resistance, i.e. they are
not effectively controlled with pesticides having the same mode of action that gen-
erally targets the same site within the pest. Both the carbamate and organophosphate
insecticides target acetylcholinesterase although each group of insecticides is chem-
ically different from one another. However, the greatest resistance concern arises
when multiple resistance is confirmed. Multiple resistance is the most difficult type
of resistance to manage because the number of management options is reduced
(Coles and Dujden 2014).
The two-spotted spider mite, Tetranychus urticae, is one of the most important
pests in a wide range of outdoor and protected crops worldwide, and its control is
still largely based on the use of acaricides/insecticides. Because of its short life
cycle, abundant progeny and arrhenotokous reproduction, it is able to develop resis-
tance very rapidly to many compounds of different chemical groups. As a conse-
quence, it has the dubious reputation to be the ‘most resistant species’ in terms of
the total number of pesticides to which populations have become resistant, and its
control has become problematic in many areas worldwide. From year 1910 to 2010,
arthropod insecticide resistance have been reported in 553 species against 331 com-
pounds; among the top 20 most resistant arthropods, T. urticae have been rated at
the top of the list. Another spider mite species, Panonychus ulmi (pest of fruit trees),
has been placed at ninth rank. Acarid mite, Rhizoglyphus robini (also known as bulb
mite) – a pest of ornamental plants and stored onions – has been ranked at 19th posi-
tion among 20 most resistant pests.
21.9 Management of Acaricide Resistance
Biological, biochemical and genetic characterization of resistance is one of the

essential elements in defining the strategy for management of acaricide resistance in
phytophagous mites (Knowles 1975; Ashton 2013; Mathew 2016; Metcalf 1980).
An effective acaricide resistance management programme could be based on gen-
eral resistance management principles endorsed by IRAC. The reduction of the
selection for resistance is possible to attain only if there were available as many as
possible acaricides with different modes of action. Resistance management requires
access to a diversity of chemistries with different modes of action. Whole genome
sequencing of TSSM and other spider mites will provide opportunities to move
from knowledge and understanding of resistance mechanisms to the practical appli-
cation in the field and might deliver novel control tools (Roush and Mckenzie 1987;
Roush and Tabashik 1990; Whalon et al. 2008).
Strategies to manage resistance are aimed at reducing the selection pressure from
pesticide to a minimum while still achieving control. Recommended strategies
involve the use of tactics designed to increase the useful life of a pesticide and
decrease the interval of time required for a pest to become susceptible once more to
a given pesticide. Adoption of IPM programme using predators, avoidance of exces-
sive repeated applications of an acaricide, usage of miticides from different chemi-
cal groups, monitoring mite population at different stages of growth of plant and
correct timings of miticides reaching even the underside of leaves are the guidelines
for reducing the risk of miticide resistance.
21.10 Conclusions
Until the mid-twentieth century, phytophagous mite populations in agroecosystems

of low-level productivity usually stayed below economic injury levels due to natural
regulation by predatory mites and insects. But advances in agricultural production
through the extensive use of pesticides and fertilizers, irrigation and cultural prac-
tices resulted in outbreaks of different spider mites on many vegetable crops, fruit
trees, ornamental plants and fibre crops. The use of many different pesticides for
controlling different other major insect pests resulted in elimination of mite preda-
tors resulting in mite outbreaks. The use of synthetic pyrethroids against different
insect pests further aggravated the mite problem as they were highly toxic and had
repellent effect on phytoseiid mite predators. So to tackle mite problem on eco-
nomic crops, frequent use of different groups of synthetic pesticides resulted in
development of resistance to different pesticides rendering chemical controls uneco-
nomic against mite pests. The use of sulphur dusts which was cheaper, easily avail-
able and did not interfere with the mite predators was promoted, but it too had
phytotoxicity problems on some crops. Petroleum oils (both summer oils and winter
oils) also used to control mites, scale insects and psylla on fruit trees, but they had
comparatively low toxicity (Cranham and Helle 1985; Georghiou 1986). But with
the extensive use of pesticides belonging to different chemical groups, resistance
410 21 Pesticides
against them developed very quickly because the spider mites had high fecundity
and low developmental rates, and many generations developed quickly in a shorter
time resulting in uneconomic control of mites.
Two-spotted spider mites, Tetranychus urticae, developed very high level of
resistance and are ranked as number one most resistant among twenty most resistant
pests. Spider mite, Panonychus citri – a pest of citrus – also ranked at number nine.
Bulb mite, Rhizoglyphus robini, ranked at number nineteen. Insecticide Resistance
Action Committee (IRAC) classified different pesticides based on their mode of
action (MOA) which facilitated extension staff and other plant protection profes-
sionals as a guide for selection of appropriate pesticides for use to delay resistance
against them. Different measures for management of acaricide resistance have been
recommended. Modern acaricides include etoxazole – an oxazoline compound
which is highly effective against eggs and immatures of spider mites. Discovery of
spirodiclofen which has new mode of action broadened biochemical diversity of
acaricides. Some novel natural acaricides like azadirachtin, milbemectin and spi-
nosad are also developed and being used.
Further Reading
Anonymous (2015) IRAC MoA classification version 7.2. http://www.irac-online.org/documents/
moa-classification/
Ashton AQ (2013) Pesticides – advances in research and application. Scholarly Edition, Atlanta.
ISBN 078-1-481-67892-6
Casida JE, Quistad GB (1998) Golden age of insecticide research: past, present or future? Annu
Rev Entomol 43:1–16
Chandler D, Davidson G, Pell JK, Bell BV, Shaw K, Sunderland KD (2000) Fungal biocontrol of
Acari. Biocontrol Sci Tech 10:357–384
Chapman RB, Martin NA (2005) Spider mite resistance management strategy. In: Pesticide resis-
tance: preventions and management strategies. New Zealand Plant Protection Society Inc,
Hastings, pp 61–70
Coles TB, Dujden MB (2014) Review: insecticide/acaricide resistance in fleas and ticks infesting
dogs and cats. Parasite Vec 7(8):9 http://www.parasitesandvectors.com/content/7/1/8
Collyer E, Kirby AHM (1955) Some factors affecting the balance of phytophagous and predaceous
mites on apple in South East England. J Hortic Sci 30:97–110
Cranham JE, Helle W (1985) Pesticide resistance in Tetranychidae. In: Spider mites: their biology,
natural enemies and control, vol 1B. Elsevier, Dordrecht, pp 405–421
Croft BA (1990) Arthropod biological control agents and pesticides. Wiley Interscience, New York
Croft BA, Brown AWA (1975) Responses of arthropod natural enemies to insecticides. Annu Rev
Entomol 20:285–325
Croft BA, Van de Baan HE (1988) Ecological and genetic factors influencing evolution of pesticide
resistance in tetranychid and phytoseiid mites. Exp Appl Acarol 4:277–300
Dabrowski T (1968) The successive action of fenitrothion and carbaryl on the spider mites
(Tetranychidae) and predatory mites (Phytoseiidae) in the apple orchards. Biul Kwarant Ochr
Rose 40:537–552
DeBach P, Bartlett BR (1951) Effects of insecticides on biological control of insect pests of citrus.
Dhooria MS, Mann GS (1988) Role of insecticides in inducing mite outbreaks. In: Mishra PC (ed)
Soil pollution and soil organisms. Ashish Publishing House, New Delhi, pp 245–262
Further Reading 411
Dhooria MS, Sagar P (1995) An outbreak of Tetranychus cinnabarinus (Boisd) (Acarina:

Tetranychidae) on squash-melon and its control. Int J Acarol 1:6–9
Dondale CD (1968) A model outbreak of the mite, Tetranychus mcdanieli McGregor in Ontario.
Proc Entomol Soc Ont 100:29–45
Georghiou GP (1986) The magnitude of the resistance problem. In: Pesticides resistance strategies
and tactics for management. National Academy Press, Washington, DC, p 14
Gerson U, Cohen E (1989) Resurgences of spider mites (Acari : Tetranychidae) induced by syn-
thetic pyrethroids. Exp Appl Acarol 6:29–46
Gerson U, Veintraub PG (2007) Mites for control of pests in protected cultivation. Pest Manag Sci
63:658–676
Harris FH (1996) Reproduction and mortality of the two-spotted spider mite on fruit seedling
treated with chemicals. J Econ Entomol 59:501–506
Helle W (1985a) Aspects of pesticides resistance in mites. In: Griffthis DA, Bowman CE (eds)
Acarology VI, vol 1. Ellis Harwood Ltd, Chichester, pp 122–131
Helle W (1985b) Resistance in Acarina: mites. Adv Acarol 2:71–93
Hennebery TJ, Taylor EA (1962) The effect of acaricide insecticide combinations on two-spotted
mite and aphid populations on outdoor rose. J Econ Entomol 55:332–334
Hueck HL, Kuenen DJ, DeBoer PJ, Draefsel J (1952) The increase of egg production of the fruit
tree red spider mite (Metatetranychus ulmi Koch) under the influence of DDT. Physiologia
Comp Oeco 2:371–377
Hull LA, Starner VR (1983) Impact of four synthetic pyrethroids on major natural enemies and
pests of apple in Pennsylvania. J Econ Entomol 26:122–130
Husseine KK (1958) The effect of insecticides on outbreaks of spider mites on cotton. FAO Plant
Prot Bull 6:155–157
Innaaya I, Degheele D (1998) Insecticides with novel mode of action. Narosa Publishing House,
New Delhi
Knowles CO (1975) Basis for selectivity of acaricides. In: Street JC (ed) Pesticide selectivity (her-
bicide, fungicide, insecticides, acaricide). Marcel Dekker & Co./Utah State University,
New York, pp 155–173
Kunz SE, Kemp DH (1994) Insecticides and acaricides: resistance and environmental impact. Rev
Sci Tech 13:1249–1286
March RB (1957) The chemistry and action of acaricides. Annu Rev Entomol 3:355–376
Mathews G (2016) Pesticides: health, safety and the environment, 2nd edn. Blackwell, Wiley
Metcalf RL (1980) Changing role of insecticides in crop protection. Annu Rev Entomol
25:219–256
Penman DR, Chapman RB (1988) Pesticides induced mite outbreaks. Pyrethroids and spider
mites. Appl Acarol 45:265–276
Ramasubramaniam T, Ramaraju R, Regupathy A (2005) Acaricide resistance in Tetranychus urti-
cae Koch (Acari: Tetranychidae): global scenario. J Entomol 2:33–39
Roush RT, Mckenzie J (1987) Ecological genetics of insecticide and acaricide resistance. Rev
Entomol 32:361–380
Roush RT, Tabashik BE (1990) Pesticides resistance in arthopods. Chapman and Hall, New York
Sandhu SS, Chander P, Singh J, Sidhu AS (1987) Effects of insecticidal sprays on the plants and
secondary pest inductions in Hirsutum cotton in Punjab. Agric Ecosyst Environ 19:169–176
Van de Vrie M (1962) The influence of spray chemicals on predatory and phytophagous mites on
apple trees in laboratory and field trials in the Netherlands. Entomophaga 7:243–350
Whalon ME, Croft BA, Mowry TM (1982) Establishment of a permethrin resistant predatory mite,
Amblyseius fallacis, in a Michigan apple orchard. Environ Entomol 11:1096–1099
Whalon ME, Mota-Sanchez D, Holling RM (2008) Global pesticide resistance in arthropods.
CABI, Wallingford/Cambridge, MA
Yu SJ (2008) Insecticide resistance. In: The toxicology and behaviour of insecticides. CRC Press/
Taylor and Francis, Boca Raton, pp 201–230
Parasitic Mites on Honeybees
22
Abstract
Honeybee, Apis mellifera which is native to Europe and Africa, has been trans-
ported from all areas where men live. Production of many agricultural crops is
dependent on bees for their pollination. More than 100 species of mites have
been reported with honeybees. Out of different mite species associated with hon-
eybees, only Acarapis woodi, Tropilaelaps clareae, Varroa jacobsoni and Varroa
destructor are economic pests of honeybees, and their infestation may lead to
destruction of the beekeeping industry in many cases. Morphology, develop-
ment, symptoms and diagnosis and dispersal of these parasitic species along with
measures to manage them in a befitting way are discussed. Methods to detect
these mites and check their dispersal to new colonies are discussed. Control mea-
sures include manipulative methods, use of soft chemicals and acaricides and use
of various non-chemical treatments against parasitic mites. Varroa destructor
described as a separate species seems to be most important among mite pests
responsible for widespread colony losses of Western honeybee. Details about its
morphology, life history, economic importance, measures to manage them on
bee colonies and steps to check their entry into uninvaded areas are discussed.
Keywords
Parasitic mites • Apis mellifera • Varroa destructor • Quarantine
There are four species of honeybees on earth – Apis mellifera is native to Europe
and Africa; A. cerana, A. dorsata and A. florea are native to Asia. A. mellifera has
been transported from all areas where men live. Honeybees have been successfully
established everywhere except in tropical Asia where parasitic mites that parasitize
on some of the three Asian species are apparently responsible for their demise and
are a great threat to the survival of beekeeping industry, both managed and feral
honeybees (Bailey and Ball 1991; Coffey 2007; DeJong et al. 1982). So the future
production of many agricultural crops dependent on bees for their pollination relies

DOI 10.1007/978-981-10-1594-6_22
414 22 Parasitic Mites on Honeybees
on keeping the good health of bees through management of their different insect and
mite pests. To understand properly the implications of parasitic mites responsible
for reduced yield of honey, reduced pollination of crops and maintenance of healthy
population of honeybees in different seasons, it is most pertinent to know the hosts
and their behaviour. The honeybee, Apis mellifera, is a social insect and lives in
colonies. Each colony has two female castes: the single queen and the workers who
may number 20,000 to 50,000 per colony. An array of pheromones regulates the
colony life. Each colony also has males called drones, who serve mainly to insemi-
nate the queen during her mating flight (Delfinado 1963; Garg and Kashyap 2002).
In literature more than 100 species of mites have been reported associated with
honeybees, but most of the mite species are not of any great importance. Out of the
different species associated with honeybees, only Acarapis woodi (Rennie),
Tropilaelaps clareae (Delfinado and Baker), Varroa jacobsoni (Oudemans) and
Varroa destructor (Anderson and Trueman) are economic pests of honeybees, and
if these mites are not properly managed, they may lead to destruction of the bee-
keeping industry in many cases (Gupta 1993; Sammataro et al. 2000).
22.1 Parasitic Mites
As stated above, four species of parasitic mites are of great economic importance in
apiculture due to their capability of causing destruction of honeybee colonies world-
wide. These species are discussed below as regard their morphology, development,
symptoms and diagnosis, dispersal and measures to manage them in a befitting way
(Matheson 1993; Sammataro et al. 2000).
22.1.1 Tracheal Mite (Acarapis woodi; Tarsonemidae)
These mites live inside the tracheae and air sacs of adult honeybees. It was origi-
nally described as Tarsonemus woodi. Mite infestation causes symptoms which are
known as ‘acarine disease’ but also referred to as ‘acariose’. Earlier the disease was
also known as ‘Isle of Wight’ disease. Tracheal mites are reported from all the major
continents of the world. They are found in the prothoracic spiracles and occasion-
ally in the abdominal and other thoracic air sacs of bees (Bailey and Perry 2001;
Komeili and Ambrose 1999). Earlier this mite was considered as the major pests of
honeybees. Female mite measures 120–190 μm in length and 77–80 μm in width;
males measure 125–136 μm in length and 60–77 μm in width (ER1).
22.1.1.1 Development
The female mite enters the body of the honeybee through the large first thoracic
spiracles; queens, workers and drones are equally susceptible to attack. Entrance
into the young bees must be made within 5 days of its emergence or it will not be
attacked by the mite. After the female mite lays her eggs in the trachea, upon hatch-
ing the young mites presumably feed on the haemolymph of the bee often piercing
22.1 Parasitic Mites 415
the tracheal wall with their mouth parts. These punctures cause the injured tissue to
become melanized, which result in a dark brown spotting of the tracheal wall – a
characteristic of heavy mite infestation.
Female mite collects within the trachea of worker bees within 24 h after the bees
emerge from their cells, and the ratio of male to female mites found within tracheae
ranges from 1:3. A female may start egg laying after 3–4 days of its emergence.
Incubation period varies from 3 to 4 days; males emerge after 11–12 days, while the
female emerges in 14–15 days.
Only adult bees under 7 days old are susceptible to infestation, and their suscep-
tibility diminishes rapidly from the moment they are newly emerged from their
brood cells. Some researchers emphasize that hairs of bees stiffen with ageing, and
the dense barriers of hairs at the entrance of the first thoracic spiracles thus became
impossible; mites can leave the spiracle when migrating, supposedly because the
hair barrier, like a valve, acts only one way. Old bees and queens are comparatively
resistant to mite infestation. The spread of mites within the colony depends upon
intimate contact between old-infested bees and young susceptible bees; mites can
only migrate in this way.
22.1.1.2 Symptoms and Diagnosis

The wings of infested bee get disjointed and become K shaped. The infested bees
become dysenteric and sluggish; and their bodies appear shiny and the abdomen
distended. Even the infested bees make small clusters in and around the outside of
the hive. There are no outward symptoms of infestation by A. woodi. It can only be
detected by dissection to expose the trachea leading from the anterior thoracic spir-
acles. When ‘Congo red’ is injected into haemolymph of infested bees, both adults
and larval mites quickly turn red. Irregular dark stains develop within the infested
trachea, the whole of which eventually blacken in infestations of long duration;
however, in a healthy bee, the trachea is recognizable as a spiral tube and is pearly
white. If the forelegs and prothorax of a bee are teased from the rest of the body with
a scalpel, the prothoracic trachea is exposed to view, and those which are darkened
are visible to the naked eyes. The disease affects the flying ability of the bee, and the
bee begins to crawl. This disease also shortened the life of a bee. Disease is notice-
able in early spring and autumn when hundred of bees are seen crawling about
around the hives, and the diseased bees are uneasy and roam about aimlessly within
the hive.
22.1.1.3 Detection of Mite Infestation

For early detection of mites and the symptoms of mite infestation in bees, sample at
least 50% of the colonies in an apiary. Collect only old bees as they are most likely
to have an infestation and are the easiest to diagnose. A total of 50 bees/apiary are
collected, and place them in a 90% ethanol (absolute alcohol solution) or freeze
them in a small vial. Dissection of bee is required for positive diagnosis of mite
infestation.
22.1.1.4 Dispersal
Acarine disease may be present in one hive without spreading to others in the apiary,
for the mites cannot live long outside the bee and can only be transferred to young
bees. It is generally considered that the most prolific cause of spreading is ‘drifting’.
It is also notable when hives are in a long row, those at the leeward end become
stronger because of their tendency to drift in that direction. Drones, in particular, are
constantly changing hive. Swarming also seems to be a means of transfer since
when a swarm issues, any bee (winged) may join it, irrespective of the hive from
which they came. Apart from the means for dissemination within the apiary, acarine
disease can readily be imported in swarms, stocks and nuclei as any other disease.
Other Acarapis spp. Found on Bees In addition to Acarapis woodi (which are
endoparasites), some other related species like Acarapis externus (Morgenthaler)
and Acarapis dorsalis (Morgenthaler) are also found associated with bees as ecto-
parasites in some areas where apiculture is practised. A. externus commonly called
the ‘neck mite’ lives in a localized area behind the head capsule of adult bees on the
ventral side of the neck. A. dorsalis lives in a V-shaped groove between the mesos-
cutum and mesoscutellum. Another ectoparasitic species, Acarapis vagans
(Schneider), is found on the wings and first abdominal segment. All these external
or ectoparasitic species of Acarapis (externus, dorsalis and vagans) are widespread
in distribution but appear to pose no serious threat to apiculture (Delfinado 1963).
Some researchers have also reported doubtful role of A. woodi mites in causing
symptoms referred to as acarine or Isle of Wight disease, but subsequently many
publications (particularly of Bailey and Perry 2001) concluded that the disease was
almost certainly caused by bee paralysis virus, which does cause the symptoms
independent of the mite, and similar associations with large densities of bee colo-
nies were found. So A. woodi does not seem to be a serious pest of honeybees. So
before initiating random control of these mites, more studies on its economic impor-
tance as pests of bees are needed under different situations.
22.1.1.5 Control
A. woodi has little effect on strong colonies and does not usually require treatment.
As brood is not infested, it may be separated, when sealed, from infested colonies
and used to create new uninfested ones by hatching in incubators and providing new
infested queens or it may be added to uninfested colonies.
Relatively specific, usually synthetic acaricides for controlling, A. woodi have
been reported in literature. Some are best applied as a smoke from smouldering
paper strips impregnated with the compounds, others produce vapours from impreg-
nated strips of wood or plastic, and a few are systemic in action and can be fed in
syrup to bees. But some of these pesticides leave unacceptable persistent residues in
honey. This has lead to the use of less specific materials like menthol and formic
acid. Menthol is usually applied as 25 g crystals placed on the top of the combs of a
colony. Formic acid can be applied as a liquid (70% industrial formic acid) soaked
on absorbent cardboard placed on the floor of the colony.
22.1.2 Tropilaelaps clareae
Tropilaelaps clareae is first described in 1961 from the material collected from the
Philippines. This mite is less widely distributed than the parasitic mite, V. jacobsoni,
but is reported in association with bees throughout tropical Asia (Webster and
Delapane 2001; Wogke 1985). Many research workers believe that T. clareae is also
a serious pest of honeybees, and the situation becomes complicated and at times
serious when both T. clareae and V. jacobsoni infest the honeybee colonies simulta-
neously (ER2).
Mites of both the sexes are elliptical or oval in shape. The females are reddish
brown in colour having palpus directed anteriorly. The males are light brown in
colour and are somewhat smaller than the females. Average length of females is 990
μm, with an average width of 510 μm. The male had a mean length of 917 μm and
a width of about 418 μm. In both the males and females, gnathosoma seems com-
pletely hidden from above by the dorsal shield. Eggs are almost round in shape and
measure about 277 μm in length and 238 μm in width and appear milky white to the
naked eye.
22.1.2.1 Life Cycle

The duration of egg stage is about 1–1.25 days. Eggs hatched into a six-legged
larva, and males looked milky white in colour with somewhat swollen idiosoma.
Larval stage lasted 0.67–0.83 days and transformed into a protonymphal stage
which too was milky white in colour and has swollen idiosoma. It attaches with the
body of worker pupa and feeds on the vital fluids. Protonymphal stage lasted for
2.5–2.8 days and then transformed into a deutonymphal stage which was also oval
in shape. Freshly formed deutonymphs are white in colour but later turned into a
light yellow. The male deutonymph is whitish in colour having elliptical body and
is somewhat smaller than the female deutonymphs. The total developmental period
for female mite is 7–8 days and 6–7 days for males. After completing the life cycle
inside the cell, newly emerged mites seek a fresh late instar larva, and cycle goes on
(Koeniger and Muzaffar 1988).
22.1.2.2 Symptoms
Brood pattern in an infested colony is scattered and gives irregular and spotty
appearance as against healthy brood which is uniform and compact. Infested brood
cells have depressed/sunken cappings. Cell with infested pupae are uncapped and in
some pupae partly eaten by bees can be seen. Hive debris contains a number of dead
mites. Partly eaten pupae and worker bees with deformed/mutilated wings and
stunted abdomens can be seen in front of the hive which are discarded by nurse
bees. In heavily infested colonies, crawlers can be seen crawling on the ground.
Such crawlers have normal wings and abdomen, but their longevity has been
reduced due to mite infestation. In such colonies, queen often stops egg laying, and
colony becomes broodless. Food reserves are more quickly consumed in T. clareae-
infested colonies as compared to healthy ones.
22.1.2.3 Diagnosis
A number of diagnostic procedures are available for detecting T. clareae infestation.
Most reliable method is to open large number of sealed worker brood cells and to
remove the pupae with forceps. Another species, Tropilaelaps koenigerum
(Delfinado and Baker), have also been reported parasitic on honeybees, Apis dor-
sata from Sri Lanka:
(i) Examination of hive debris. A thick sheet of white paper is placed on the bot-
tom board, and after some days, mites that naturally die and fall on the bottom
board are collected and examined. A wooden board can also be fitted with a
wire gauze of 8–12 mesh size over the white sheet to protect debris and paper
from bees. The debris can also be examined by placing it in a jar filled with
95% alcohol. The mites will float over the surface, collected and examined.
(ii) Examination of brood. Sealed brood is uncapped, and each pupa is removed
with forceps and examined. In case of heavy infestation, at least 200 cells
should be examined.
(iii) Chemical diagnosis. For observing infestation of T. clareae infestation, an
acaricide should be used for diagnosis, and debris on the bottom board is
examined for dead mites. To avoid contamination this type of diagnosis should
not be followed when the hive contains the honey.
22.1.2.4 Control Measures

Sulphur dusting is reported to control T. clareae infestation. Fumigation with 5 ml
of 35% formic acid/day reported to be highly effective and needs application after
weekly intervals till infestation is eliminated. But its dose needs to be standardized
considering its deleterious effect on the brood and worker bees. Amitraz applica-
tions also reported successful against mites. Apistan and fluvalinate fumigation also
reported effective against mites and had no deleterious effect on brood and also did
not reduce the longevity of adult bees.
Non-Chemical Control Measures Brood-rearing cycle of the infested colonies

should be manipulated in a way that the mites are deprived of sealed and unsealed
brood for at least three days. Removal of brood from the bee colonies resulted in
mortality of all T. clareae mites within 48 h. In beekeeping, there are several means
of creating broodless situation in an infested colony. Sometimes both deprivation
and chemical control (like fumigation) of mites are combined in an intelligent way.
22.1.3 Varroa jacobsoni
This mite is a native ectoparasite of Apis cerana bees throughout Asia and is widely
distributed on Apis mellifera and A. cerana in both the Old and New Worlds and in
Asia and the Far East (Bhaskar and Putatunda 1989; Ramirez and Otis 1986). The
mite infestation caused serious concern for apiculture industry throughout the world
(ER3, ER4 and ER6).
22.1.3.1 Symptoms
The mite reproduces mainly on the drone pupae of A. cerana and A. mellifera. Live
mites usually are found within the sealed brood cells or are partly hidden between
the abdominal segments of adult bees, so in spite of their comparatively bigger size
than other bee parasitic mites, they are not easily noticed by the beekeepers.
However, dead mites that fall from adult bees or are cleaned from brood cells can be
easily seen in debris from the hive floor. Severely infested colonies of A. mellifera
in Europe often show symptoms of brood disease as the mite can activate and trans-
mit honeybee viruses which may lead to colony mortality (DeJong 1997; Gupta
1993). Mite parasitized pupae sometimes produce stunted or deformed newly
emerged bees. Emerging bees from infested cells are comparatively underweight
and may also have reduced longevity. Severe mite infestation of A. cerana colonies
may cause absconding.
22.1.3.2 Diagnosis
The mite is easily seen as brown or whitish spots with the naked eye against the
white body of developing bee pupae when observed after uncapping with uncapping
fork, or else the pupae may be lifted from cells and examined. Sampler of bees can
be placed in a large screw-topped glass jar containing a cotton wool pad moistened
with diethyl ether and rotated for about 10 s. Both bees and mites are anaesthetized;
the mites will fall from the bees and stick to the sides of the jar.
Through Sticky Board A white paper or plastic sheet covered with petroleum
jelly or another sticky agent is placed on the bottom board of a colony, and the hive
is stroked with the pipe tobacco in a smoker. After closing the hive for 10–20 min,
the board is removed, and the mites are counted. Alternatively, a sticky board by
itself can be left in place for 1–3 days. Hive debris can be examined for finding out
mites fallen from bees.
22.1.3.3 Spread of Mites

Varroa jacobsoni mites have been introducing into the new areas of the world on A.
mellifera through importation of bee stocks and also through migratory beekeeping
practices. In Asian countries, V. jacobsoni may be a limiting factor in the success of
beekeeping operations with A. mellifera. The transfer of the mite from A. cerana to
A. mellifera is inevitable in all regions where the two bee species occur together.
These mites may also spread from one to other area by transfer of infested brood
combs/colonies, drifting and robbing.
22.1.3.4 Biology
The gravid mite enters a brood cell shortly before it is capped, and in the capped
cell, the mite starts laying eggs as soon as the larva has finished cocoon spinning
(ER5, ER7, ER8 and ER9). Female mite must feed on larval blood before she can
lay eggs. Mites can live on the blood of adult bees but in this way cannot lay eggs.
The female mite lays 2–5 eggs at varying intervals, and the eggs hatch after 24 h.
The complete development of the female takes 8–10 days and that of the male for
6–7 days. The males mate with the females in the capped cells. The chelicerae of the
male are transformed for sperm transfer, and the male therefore cannot take food
and dies soon after mating. The mother female and young female (now mated)
remain in the brood cell until the adult bee emerges when they leave the cell sticking
to the body of the bee. The adult bee is usually an intermediate host and means of
transport for the mite. The most serious parasitization always occurs on the older
larvae. Drone larvae are preferred to worker larvae. Ideal temperatures for varroa
development correspond to the ideal temperatures of drone bees. In case of heavy
infestation, queens’ larvae may also be infested by the mite. An infested drone may
have as high as 12 mites, but up to 5 mites per worker bee may be found, and a
number of mites in a colony may exceed 10,000 individuals. Mite populations in an
individual brood cell are reported as high as 12 per worker brood, but up to 20 mites
may be found per drone cell. The young female mites live for about two months in
summer but may live for 3–8 months during winter. The number of mites is small in
spring, increases during the build-up period and is largest in autumn. The mite pop-
ulation is correlated to brood rearing. The infestation may be very heavy during
fourth year of mite infestation. The colony becomes weak and may finally perish.
22.1.3.5 Control Measures
Manipulative Methods These methods exploit the biological necessity of V.

jacobsoni to enter honeybee brood cells to reproduce. Mites have a strong prefer-
ence for drone brood and a proportion of the mite population may be removed by
the systematic rearing of the drone brood that is destroyed after the cells are sealed.
An alternative method is to confine the queen on a single empty comb for three suc-
cessive periods of 8–9 days. Each comb is removed after the cells are sealed and
may be destroyed or returned to the colony after mites have been killed by treatment
with formic acid.
Soft Chemicals Formic acid, lactic acid and volatile essential oils from plant
extracts have been approved for mite control in some countries. Formic acid (60%)
put on a plate of pulp board placed beneath the brood nest of a colony kills most
mites even those within sealed brood. But lactic acid (15% solution) is sprayed on
to adult bees on every comb will kill adhering mites but not those within the brood
cells. Thymol, eucalyptus oil and menthol applications are also effective against
mite.
Acaricidal Control Acaricides are mainly applied through fumigant strips which
are burnt in the colony, plastic carriers impregnated with synthetic pyrethroids and
application of systemic acaricides through food exchange. If properly applied these
acaricide applications are very effective against mites. However, repeated applica-
tions of acaricides may result in accumulation of residues in the combs. Generally
phenothiazine, dicofol, bromopropylate, chlordimeform hydrochloride, fluvalinate
(Apistan or Mavrik strip) and amitraz have been used in different formulations.
Non-Chemical Treatment The Varroa mite depends on bee brood to complete its
developmental cycle. Since the mite prefers drone to worker brood, frames of drone
cells are given to the colonies, which will rear drone brood in them. When the cells
are sealed, the frames containing the mites trapped inside the cells can be removed
and destroyed.
Another method involves caging the queen for 21 days to make the colony brood-
less. Thereafter the activities of the queen are restricted to a couple of comb using
queen excluder. The queen lays in these combs, and the adult Varroa mite will enter
the brood cells to deposit their eggs. When most of the brood is sealed, these frames
are removed and destroyed.
22.1.4 Varroa destructor Anderson and Trueman
Until 2000, it was believed that V. jacobsoni is the mite species responsible for
widespread honeybee colony losses. But taxonomic work published in 2000
(Anderson and Trueman 2000) indicated that a previously unidentified species
Varroa destructor was responsible for the damage, while V. jacobsoni was shown to
be only moderately harmful to Western honeybees; the following details are only for
V. destructor, which has been reported the most devastating pest of Western honey-
bees, Apis mellifera. Although the Varroa complex includes multiple species, V.
destructor is the species responsible for the vast majority of the damage attributed
to mites. In Asian countries including India, V. destructor is well established and
causing significant reduction in honey yield (Akratnakul 1987; Baggio et al. 2004;
Hoffmann and Curry 2005).
22.1.4.1 Morphology
Adult female mites are reddish brown to dark brown and are oval in shape, measur-
ing 1.00–1.77 mm in length and 1.50–1.99 mm in width. Adult males are yellowish
with lightly tanned legs and spherical body (0.75–0.98 mm in length; 0.70–0.88 mm
width).
22.1.4.2 Life History

Adult females undergo two phases in their life cycle, the phoretic and reproductive
phases. During the phoretic phase, female varroa feed on adult bees and are passed
from bee to bee as bees walk past one another in the colony. During phoresy the
female mites live on adult bees and usually can be found between the abdominal
segments of the bees. Varroa mites puncture the soft tissue between the segments
and feed on bee haemolymph through the punctures. Phoretic mites pass between
colonies of bees when infested bees drift into another colony. The varroa infestation
also spreads when beekeepers transport colonies from one area to another; through
swarming and moving to a new location, the spread of varroa is simultaneously
done. During robbing activities, spread of varroa from colony to colony also occurs.
The phoretic period may last 5–11 days when brood is present in the hive or as long
as 5–11 days when brood is present in the hive or as long as 5–6 months during the
winter when no brood is present in the hive. Female mites in presence of brood have
an average life expectancy of 27 days, yet in the absence of brood may live for many
months. An adult worker, drone or queen bee emerges from the cell, respectively, in
21, 24 or 16 days after an egg is deposited.
22.1.4.3 Economic Importance

Accurate estimates of the effect of varroa on the apiculture industry are hard to find,
but it is safer to assume that the mite invades hundreds of thousands of colonies
worldwide, resulting in billions of dollars of economic losses. Varroa mite problems
result in increases in the cost of beekeeping and lowering the margins of beekeep-
ers. Varroa mites rarely kill adult bees but definitely shorten their life span, but var-
roa mites can kill immature bees. Certain viruses are also associated with varroa
mite presence and their levels in a colony. In the infested colony, occurrence of
newly emerged adult bees with misshapen wings is commonly observed.
22.1.4.4 Management
Early detection of mites on adult and immature bees can help in their proper man-
agement in the hive. Keeping sticky paper trap under the honeybee nest can help to
find both live and dead mites falling from their host bees. Ether rolls and powdered
sugar technique can also be used for knowing the presence of varroa mites. Minimum
of 300 adult bees are sampled in these techniques. For knowing economic threshold
levels, sampling devices are of permanent importance and helpful in devising IPM
to varroa mite control. Some authors have suggested an economic threshold (ET)
for varroa mites around 3000 mites/colony, although development of ET level is
dependent on several other factors. Several pyrethroids, organophosphates, essential
oils and organic acids have been used in many countries to control Varroa mites
(Rice et al. 2004).
Other Varroa Species on Bees Varroa underwoodi (Delfinado-Baker and

Aggarwal) reported in 1987 from Apis cerana colonies in Nepal. This species is
very similar to V. jacobsoni but is much smaller in size (780μm long and 1170 μm
wide) and is light chestnut brown in colour. The body of male is rounded, weakly
sclerotized with light tanning on legs and setae. Euvarroa jacobsoni was first
described in 1974 from Thailand and thereafter from India and many other places.
This mite parasitized only the drone brood. E. sinhai is quite large (1040 μm in
length and 1000 μm in width) and brown in colour and is broadly pear shaped lack-
ing a fixed cheliceral digit. Its life cycle is similar to that of V. jacobsoni except that
the mites attack only drone brood. The adult female and the protonymphal and
deutonymphal stages of both sexes feed on drone brood. Only adult females leave
the cells, and they are phoretic on adult drones.
Further Reading 423
22.1.5 Other Mites Found on Bees
Several solitary and social bee species are also found to host a great diversity of mite
species. Associations of different mite species on bees may vary from Locustacarus
buchneri (Stammer) (family Podapolipidae) reported as tracheal parasite on various
species of Bombus bees. Acarapis woodi also reported endoparasites in the tracheae
of Bombus bees. Chaetodactylus spp. (family Chaetodactylidae) reported as clepto-
parasites with megachilid bees and Hosstia spp., Chaetodactylus krombieni (Baker),
have been reported as parasitic or may have phoretic role on leaf or mason bees
(family Megachilidae). The family Acaridae has been reported to be associated with
carpenter bees, which feed on the provisions after destroying the bees’ eggs. Many
mite species have been found as commensals in nests particularly of social honey-
bees, bumble bees and stingless bees. Tarsonemid mite, Tarsonemus indoapis, has
also been reported from neck region of workers of Apis species.
22.2 Conclusions
There are four species of honeybees on earth. Apis mellifera is native to Europe and
Africa. But A. cerana, A. dorsata and A. florea are native to Asia. Out of more than
100 species of mites reported associated with honeybees, only 4 species, namely,
Acarapis woodi, Tropilaelaps clareae, Varroa jacobsoni and V. destructor, are eco-
nomic pests and, if uncontrolled, may lead to destruction of beekeeping industry in
many cases. Previously A. woodi (endoparasite) was considered as a very serious
pest of bees, but reports state that acarine disease symptoms are because of bee
paralysis virus which does cause the symptoms independent of the mite. T. clareae
is reported as an important pest of honeybees particularly in tropical Asia and some-
times proves serious when it appears in association with V. jacobsoni, but is only
moderately harmful to Western honeybees. However, V. destructor is the species
responsible for the vast majority of the damage attributed to parasitic mites in
Western honeybees, A mellifera. In Asian countries too, V. destructor is well estab-
lished and is causing significant reduction in honey yield and losses on account of
pollination of crops resulting from bad bee health.
Further Reading
Akratnakul P (1987) Amitraz for Varroa control. In: Honey bee diseases and enemies in Asia: a
practical guide. FAO Rome Publication, Rome, pp 1–51
Anderson DL, Trueman JHW (2000) Varroa jacobsoni (Acari: Varroidae) is more than one species.
Baggio A, Arculeo P, Naneti A, Morinelli E, Mutinelli F (2004) Field trial with different thymol-
based products for the control of Varrosis. Am Bee J 144:395–400
Bailey L, Ball BV (1991) Honey bee pathology. Academic, London
Bailey L, Perry JN (2001) The natural control of the tracheal mite of honey bees. Exp Appl Acarol
25:745–749
Bhaskar S, Putatunda RN (1989) Mesostigmatid mites associated with bees in India. In: Progress
in acarology vol 2. Oxford and IBH Publ. Co, New Delhi, pp 287–289
Coffey MF (2007) Parasites of the honey bee. Teagase Crops Research Centre, Oak Park, 47 pp
DeJong D (1997) Mites: Varroa and other parasites of brood. In: Honeybee pests, predators and
diseases, 3rd edn. The A.I. Root Company, Medina, pp 278–327
DeJong D, Morse RA, Eickwort GC (1982) Mite pests of honeybees. Annu Rev Entomol
27:229–252
Delfinado MD (1963) Mites of the honeybees in South-East Asia. J Apic Res 2:113–114
Garg A, Kashyap NP (2002) Mites and other enemies of honey bees in India. In: Perspectives in
Indian apiculture. Agrobios, Bikaner, pp 264–303
Gupta M (1993) Honey bee mites: an overview. Indian Bee J 55:1–10
Hoffmann GD, Curry R (2005) The population dynamic of Varroa mites in honey bee colonies
Part-I – the Varroa programe. Am Bee J 145:592–595
Koeniger N, Muzaffar N (1988) Life span of the parasitic bee mite, Tropilaelaps clareae on Apis
cerana, A. dorsata and A. mellifera. J Apic Res 27:207–212
Komeili AB, Ambrose JT (1999) Biology, ecology and damage of tracheal mites in honey bees
(Apis mellifera). Am Bee J 130:253–257
Matheson A (1993) World bee health report. Bee World 74:176–212
Ramirez WB, Otis GW (1986) Developmental phases in the life cycle of Varroa jacobsoni –an
ectoparasitic mite on honey bees. Bee World 67:92–97
Rice ND, Winston ML, Higo MA (2004) Integrated pest management for the parasitic mite, Varroa
destructor (Anderson and Trueman) in colonies of honey bees (Apis mellifera). Am Bee
J 144:791–795
Sammataro D, Gerson U, Needham G (2000) Parasitic mites of honey bees: life history, implica-
tions, and impact. Annu Rev Entomol 45:519–548
Webster TC, Delaplane KS (2001) Mites of the honey bee. Dadant and Sons, Inc., Hamilton,
280 pp
Wogke J (1985) Tropilaelaps clarea, a serious pest of Apis melliferae in the tropics, but not danger-
ous for apiculture in tropic zones. Am Bee J 125:497–499. Parasitic Mites on Honeybees
1. https://www.youtube.com/watch?v=6wy2PG_MB4Y Episode 1 : Tracheal Mites

2. https://www.youtube.com/watch?v=_W2DsqT3I2o Varroa jacobsoni
3. https://www.youtube.com/watch?v=tJO0epbVGXg Varroa
4. https://www.youtube.com/watch?v=R5LxTf-EYIY Drone frame full of Varroa mites
5. https://www.youtube.com/watch?v=G9-FGA3bwEw&list=PL72NAHApLyjlTHXyRrzT0Zn
QRpBuz00bD&index=27 Biology of the Varroa Mite
6. https://www.youtube.com/watch?v=WyKhmvv4HHA&list=PL72NAHApLyjlTHXyRrzT0Zn
QRpBuz00bD&index=26 Varroa destructor Under Microscope X100 X400 Pest of Honeybees
7. https://www.youtube.com/watch?v=u3nZQ2kiYHs Varroa Mite History, Distribution and
Biology
8. https://www.youtube.com/watch?v=G9-FGA3bwEw Biology of the Varroa Mite
9. https://www.youtube.com/watch?v=a2vg59Snt6c Life Cycle of the Honeybee and Varroa Mite
10. https://www.google.co.in/search?q=Varroa+destructor&espv=2&biw=1024&bih=640&sourc
e=lnms&tbm=isch&sa Varroa destructor (Images)
Medical and Veterinary Acarology
23
Abstract
More than 250 species of mites are recognized as the cause of health-related
problems for humans and domestic animals. Types of problems include tempo-
rary irritation of the skin due to bites or feeding on host skin, fur or feathers;
persistent dermatitis; mite-induced allergies; transmission of pathogenic micro-
bial agents; and invasions of respiratory passages, ear canals and occasionally
internal organs. Chicken mite (Dermanyssus gallinae) is considered as one of the
most serious pests of poultry birds and may also serve as vectors for diseases
such as ‘salmonellosis’ and avian ‘spirochaetosis’. Mite feeding in poultry birds
results in pain, irritation, restlessness and decreased egg production. Straw itch
mite (Pyemotes ventricosus), chiggers (Eutrombicula alfredugesi) and burrow-
ing mites (Sarcoptes spp., Notoedres spp. and Knemidocoptes spp.) are other
important parasitic mites. Non-burrowing mites (Psoroptes spp., Chorioptes spp.
and Otodectes spp.) also have great economic importance as parasites of veteri-
nary animals. Nasal mites, belonging to families Halarachnidae and
Macronyssidae, infest nasal cavity of dogs and snakes. House dust mites
(Dermatophagoides farinae and D. pteronyssinus) live in association with man
and are responsible for house dust mite allergy. Places with damper climates are
more favoured by mites than dry ones. Ticks are exclusively bloodsucking ecto-
parasites and disperse pathogenic organisms including protozoan, viral, bacterial
and even fungal pathogens. Tick-borne diseases of livestock include babesiosis,
theileriosis, tularaemia, Lyme disease and Rocky Mountain fever.
Keywords
Chicken mite • Straw itch mite • House dust mite • Ticks
Mites attacking humans with some frequency originate in a variety of habitats.

Some species migrate to man from birds, including the chicken mite and the tropical
fowl mites. Others such as grain mites and mushroom mites are found in food

DOI 10.1007/978-981-10-1594-6_23
426 23 Medical and Veterinary Acarology
materials or stored products. The straw itch mite and furniture mite come from plant
material, and the chigger mite is found in lawns and open woodlands. From rodents
come the tropical red mite and the mouse mite, whereas the itch mite and follicle
mites are permanent residents on man. Infestation with either mites or ticks is
referred to as ‘acariasis’ (Burgess 1994; Evans 1992; Wooley 1988). Important spe-
cies of mites and ticks reported in literature in relevance to human beings and other
animals including both under wild and kept as pet and poultry and in zoos are dis-
cussed briefly in this chapter.
23.1 Mites
More than 250 species of mites are recognized as the cause of health-related prob-
lems for humans and domestic animals. Types of problems include temporary irrita-
tion of the skin due to bites or feeding on host skin, fur or feathers; persistent
dermatitis; mite-induced allergies; transmission of pathogenic microbial agents;
invasions of respiratory passages, ear canals and occasionally internal organs; and
abnormal fear of mites (delusory acariasis).
23.1.1 Chicken Mite (Dermanyssus gallinae (DeGeer);

Dermanyssidae)
Dermanyssus gallinae is also commonly known as red mite, roost mite and poultry
mites. They are comparatively large sized (about 1.5 mm) with long legs. Body
colour is white to greyish black, becoming red when engorged with blood. The mite
spends much of its time away from its host, the nymphs and adults only visiting
birds to feed mainly at night. They are parasites of birds (including poultry birds)
and also feed on humans and many other mammals resulting in dermatitis. Chicken
mites spend most of their time in the environment and drop off the host after feed-
ing. Poultry houses with wooden construction are the favoured habitats of these
mites; they lay their eggs in the crevices. Life cycle is completed in one week. The
adults can survive for several months without feeding so that a reservoir population
can persist in unoccupied poultry houses and aviaries. This mite is especially a
threat to fowls housed in old buildings (Axtel and Arends 1990; Kettle 1990).
D. gallinae is considered one of the most serious pests of poultry. Mites infest
hens, affect both directly and indirectly and may also serve as vectors for diseases
such as ‘salmonellosis’ and avian ‘spirochaetosis’. Chicken mites also feed on live-
stock and human beings causing dermatitis and skin lesions in some people. Mite
feeding in poultry birds may result in pain, irritation, restlessness and decrease in
egg production. When mite infestation is quite high, they may cause anaemia in
hens with consequent weakness and allergy. Blood spotting of eggs (reddish-black
drops of squashed mites and or mite faeces) may be due to D. gallinae infestation
(ER6).
23.1 Mites 427
23.1.2 Northern Fowl Mite (Ornithonyssus sylviarum (Canestrini

and Fanzago); Macronyssidae)
Ornithonyssus sylviarum is found worldwide and is very similar looking to the

chicken mite but differs in that it breeds among the feathers of the host and may
complete development without leaving the host. O. sylviarum have been frequently
reported in Australia, New Zealand and temperate parts of the Northern Hemisphere.
They can survive for about 2 weeks away from their host. Northern fowl mite may
bite man causing annoyance and even dermatitis. The problem most frequently is
irritation from the occasional bites by wandering mites which are otherwise a gen-
eral parasite of birds particularly the domestic fowls, sparrows, swallows and many
other bird species. In the absence of the normal host birds, human annoyance is
frequently observed. The mite is almost a permanent parasite, and infection is by
contact or by placing in accommodation recently vacated by infected stock (ER7).
In case of heavy mite infestation, birds are restless and lose weight from irritation;
egg production may be reduced, and there may be severe anaemia. Infested birds
may have thickened crusty skin and soiled feathers around the vent (Kettle 1990).
23.1.3 Tropical Rat Mite (Ornithonyssus bacoti (Hirst);

Macronyssidae)
Tropical rat mite, also known as Liponyssus bacoti, commonly found associated
with rats but also feeds on many warm-blooded animals including man. Mites lay
eggs in the burrows, nests and cages of rodents and small marsupials. The mite bite
is painful causing intense itching and a skin irritation symptoms commonly known
as rat mite dermatitis (ER7 and ER 25). O. bacoti has not been reported transmitting
typhus or any other disease, although the habits of the mite may suit for such a role.
Attack on man almost always associated with areas which may be infested with rats
in buildings. Complaints of rat mite dermatitis are common in rat-affected areas like
in warehouses, stores and theatres. Rat control may intensify the mite attack on
man. After each feeding mite will drop from their hosts and may be found on a
variety of surfaces near rat-infested areas. Mites can survive for up to two months
without the blood meal. O. bacoti is found on birds including chicken, turkeys,
ducks, pigeons, sparrows, starlings and mynah. It also feeds on mammals if birds
are not available. Mite bites can be painful and pruritic and may cause irritation and
localized dermatitis (Mullen 2002).
23.1.4 House Mouse Mite (Liponyssoides sanguineus (Hirst);

Dermanyssidae)
In earlier literature, Liponyssoides sanguineus is also known as Allodermanyssus

sanguineus. This mite is primarily a parasite of small rodents such as mice (Mus
musculus), but when it leaves the mice host and wanders around or inside buildings,
then it bites man. The bites can be painful and pruritic and may cause irritation and
localized dermatitis. House mouse mite is known as a vector of rickettsial pox – a
mild and nonfatal disease of man. Life cycle of L. sanguineus is similar to that of D.
gallinae. The adults can survive up to two months without host in the environment
(ER1).
23.1.5 Grain Mites (Glycyphagus domesticus (DeGeer); Acaridae)
Stored grain mites are also sometimes known as furniture mite and mushroom mite.
Mites are found commonly infesting all types of grain flour, stored foods, cheese
and in mushroom beds. Stored grain mites prefer moist locations and develop rap-
idly under favourable conditions. Life cycle is completed in about 17 days and is
considerably extended under adverse conditions producing ‘hypopus’ which is
transported from one place to the other through insects and mice (ER1). On avail-
ability of favourable conditions, hypopus sheds its skin and resumes normal growth
and development. So control measures should be planned keeping in view the for-
mation of hypopus stage as this stage is resistant to many adverse weather condi-
tions and pesticides. Grain mite infestations have been reported to cause the mild
dermatitis in personnels handling mite-infested grains and other stored products.
Such dermatitis is also referred in literature as ‘grocer’s itch’ and ‘copra itch’. Grain
mites are not bloodsuckers but are a cause of only mild irritations which can easily
be controlled if a source of exposure is identified and managed properly (Fuller
1956).
23.1.6 Straw Itch Mites (Pyemotes ventricosus (Newport);

Pyemotidae)
This mite species live on other arthropods like larvae of several insects like
Angoumois grain moth, granary weevils (on rice, bean and pea) and the pink boll-
worm. Reproduction of mite is quite rapid; this mite has unusual development as the
eggs are laid inside the body, hatches and develop inside the females’ swollen body
(ER10). Only the sexually mature adults emerge from the female. About 200 adult
mites emerge from a single female. Under domestic conditions, mite bites originate
from beetle-infested cereals and beans. Personnels engaged in threshold straw or
handling grains or other material infested with insect hosts often are overrun by
these mites. Mite bites produce a rash-like dermatitis which may cover large areas
of the body; the rash appears about 12 h after the attack and is accompanied by
severe itching. Larvae of trombiculid mites are known as chiggers. These mites are
parasitic only.
23.1 Mites 429
23.1.7 Chiggers (Eutrombicula alfredugesi (Oudemans);

Trombiculidae)
Chiggers are also known as red bugs during larval stages, but both adults and
nymphs are free-living. Chiggers infest a variety of areas ranging from those over-
grown bush to well-kept lawns (ER1). They are also frequently found in orchards,
woodlands, pine straw and along lakes and stream margins. Adults overwinter in
earthen cells in the soil, scavenge on decaying matter and emerge from soil in the
spring to lay their eggs. Orange-coloured larvae emerge from the eggs and normally
feed on snakes, turtles, rabbits, birds and other wildlife (ER8). Chiggers are very
active but are barely visible to the naked eyes. Whenever man comes in contact with
infested vegetation, chiggers swarm over the entire body and may take several hours
before they settle down to feed. Chiggers attack seems to be concentrated at points
where the clothing is pressed against the skin as under belts or garters.
Egg laying begins when soil surface temperatures are regularly above 16 °C, and
after hatching chigger larvae climb onto grass blade, twigs or other objects from
which they can move readily and snag a passing host. After feeding for 2–4 days,
the larvae drop from the host and become nymph after moulting. A complete life
cycle requires from 7 to 10 weeks. Nymphs and adults are not parasitic. They are
predators on small insects, other mites and their eggs. Chiggers do not burrow into
the skin, but larval stage firmly attaches to the skin. After feeding, the larvae dis-
lodge and drop to the ground. An allergic reaction to saliva injected by larval chig-
gers causes reddish welts which itch. Though the bites are primarily annoying, the
hosts’ scratching may break the skin and permits infection. A large infestation on
poultry may kill some of the birds, whenever possible avoid moving in areas of the
tall grass or weeds where little sunlight penetrates to the soil. Such areas harbour
most of the chiggers. Avoid sitting or living in such areas or in shaded areas even
with short grass or thatch. Protection from chiggers uses two approaches. The first
approach is the use of a repellent which can discourage chiggers from attacking.
The most effective repellents are ‘Deet’ and ‘permethrin’. Both are applied to cloth-
ing. The second approach is to keep down mite population through regularly lawn
mowings. But large populations may require the use of acaricides like bifenthrin,
cyfluthrin and carbaryl.
23.1.8 Burrowing Mites
Sarcoptes, Notoedres and Knemidocoptes are the three genera of family Sarcoptidae
which have burrowing mites. However, family Demodicidae also have one burrow-
ing mite genera, namely, Demodex.
Sarcoptes scabiei (DeGeer) These mites occur on humans and more than 100 spe-
cies of mammals and marsupials. But by biological adaptation, different strains
have evolved which are largely host specific. Sarcoptid mites in veterinary animals
and on humans are well known as a cause of ‘mange’ (ER4 and ER5). The disease
symptoms caused by the mite feeding on human beings are generally known as
‘scabies’.
Scabies Sarcoptes scabies var. hominis is found on humans and can only develop
and reproduce on a human host. Scabies mites tend to prefer areas of folded skin
(e.g. web between fingers, under buttocks, elbows, wrist areas, around genitals, etc.)
for burrowing. They are generally white or colourless. Scabies mites do not transmit
disease; however, their burrowing and feeding activities create the irritation, and
allergen requires symptomatic of the infestation. The mite activities and the associ-
ated itching and scratching may also lead to secondary bacterial infections. The
primary mode of transmission of the human scabies mite is direct skin contact
between two individuals. Application of 5 % permethrin cream (Elimite) and iver-
mectin (Stromectol) administration under medical supervision only is recom-
mended. Application of 10 % crotamiton (Eurax) has been reported effective in
treatment of the scabies (Arlian 1989b; Kettle 1990).
Sarcoptic Mange of Cattle In many cases, mild symptoms because of mite infes-
tation may be found, but sarcoptic mange is potentially the most severe of the cattle
manges. Mite is mainly found on neck and tail regions, but it may be found in any
part of the body. Mild infection show merely scaly skin with little hair loss, in severe
cases the skin becomes thickened. There is marked loss of hairs, and crusts form on
the less well-haired parts of the body such as the escutcheon of cows. There is
intense pruritus leading to loss of meat and milk production (Mullen 2002). Value
of hides is also decreased because of damage by scratching and rubbing. Ivermectin
injections have been reported effective against cattle mange.
Sarcoptic Mange of Goat and Sheep Mite prefers regions without wool such as
the face, ears and groin. Affected areas are first erythematous and scurfy. Intense
pruritus is present, and sheep scratch and rub the head. Because of the itch, sheep
are almost continuously restless and are unable to graze, so there is progressive
emaciation. In case of haired sheep the whole body may be affected. In goats, the
condition of mange is often chronic. Irritation with encrustations, loss of hair and
excoriation from rubbing and scratching is found.
Mange of Rabbits Burrowing mites, S. scabiei and Notoedres cati (Hering), pres-
ent a zoonotic danger. These mites also affect dogs and cats, causing a transient
itching and dermatosis (ER9). The burrowing mange mite, N. cati, is a c osmopolitan
parasite of domestic cats, but it also infests several wild cats (e.g. cheetah, snow
leopard), mongoose and domestic rabbits. These mites spread rapidly from one rab-
bit to another through nymphs and larvae that can be on the surface of the skin.
Wounds appear on the lips and nose, later around the head, neck and sometimes
around the genitalia. Scratching by the rabbits leads to alopecia (loss of fur). Watery
stuff comes out from affected portions that form crusts upon drying. Self-mutilation
leads to wounds and secondary bacterial infection. The mange-affected rabbits
23.1 Mites 431
become lethargic and can die within a few weeks. Mange in rabbits has been treated
by avermectin injections. Wounds can also be treated by benzyl benzoate.
Knemidocoptes (Family Knemidocoptidae) Knemidocoptes mutans (Robin and

Lanquetin) infests domestic birds. K. mutans affects poultry birds and causes ‘scaly
leg’ symptoms (also called ‘tassel foots’). The entire life cycle is spent burrowing in
the unfeathered scales, skin of a bird’s feet and shanks or occasionally its beaks and
cere (the fleshy area around its nostrils). Individual birds contract the mites through
prolonged direct contact with other infested surroundings. Older birds are the most
commonly affected; it is uncommon to find these mites in birds in a commercial
operation: Commercial poultry are generally young, and they rarely have direct
contact with older birds that may be infested with this mite. First indication of para-
sitism is a brittle, flaky or powdery appearance to the birds’ legs. The mite burrow-
ing and feeding activity causes inflammation and disfigurement of the skin with
crusts, scale and scab formation with swollen legs and feet. The skin hardens and
exudes fluid. Sometimes comb and neck region are also infested. Heavily infested
birds may die because of hypersensitive allergic reactions. K. gallinae, K. laevis and
Neocnemidocoptes gallinae (Raillet) infest primarily the birds. K. pilae causes scaly
leg or tassel foot in caged birds. K. jamaicensis and K. intermedius also reported
infesting canaries causing ‘scaly leg’ symptoms. Procnemidicoptesjansseni Fain
and N. gallinae are infesting pheasants, chickens, pigeons and geese, resulting in
depluming. As a preventive measure, isolate or cull birds that are infested with scaly
leg mites. But if a small number of birds are affected, then treat scaly leg mites by
directly applying an oil-based product such as petroleum jelly and a mixture of
50:50 kerosene and cooking oil mix or Blue Ribbon (a commercially available mix-
ture of plant oils and camphor in a canola oil extract).
Demodex (Family Demodicidae) These mites are burrowing mites but are quite
different in form and behaviour from the members of family Sarcoptidae. They
infest all domestic mammals and even man. Mites infest the hair follicles and seba-
ceous glands. Demodex folliculorum (Simon) infests man, D. phylloides (Croker)
on pigs, D. caprae (Raillet) on goats, D. canis on dogs, D. bovis on cattle and D.
equi on horses (ER21). Most species spend the entire life cycle in the follicles or
glands (ER21). The mange by these mites is commonly found in the dogs, and the
symptoms are known as ‘demodectic mange’. It is most uncommon for a host spe-
cies to harbour two to four different Demodex spp. Only prolonged contact with the
mange-affected animals can transmit the mite to the healthy animals. Absence of
pruritus is a notable feature of all types of demodectic mange. For confirmatory
diagnosis, deep scrappings are necessary to take mites deep from the follicles and
glands. Healthy feral animals almost never suffer from demodectic mange; labora-
tory or domesticated hosts are the usual victims. Normally the mites are not readily
accessible to topical application of acaricides, but repeated treatment is necessary.
23.1.9 Non-Burrowing Mites
Families Psoroptidae, Cheyletidae and Dermanyssidae have representatives which

have great importance as regards veterinary animals. Their members do not burrow
into the dermis, but some representatives also suck fluid from the skin. Some other
members are also bloodsuckers. As regards their behaviour, some species live per-
manently on the skin surface, and others live mainly in the hair or fur. But some
other representatives visit the host jointly to feed. In some of the species, adult stage
is free-living but must feed on animals during their larval phase (Kettle 1990;
Steelman 1976).
Psoroptes Members of these genera are typically non-burrowing mites. Females

during its lifetime of 4–6 weeks lay about 90 eggs; development from egg to adult
takes about 10 days. Psoroptes ovis (Womersley) and P. bovis (Hering) infest mainly
sheep, camel and cattle; P. equi (Raspail) infests horses; and P. cuniculi (Delafond)
mainly infests rabbits but may also infest horses, cattle, sheep and goats. Symptoms
produced by feeding of mites are known as ‘psoroptic mange’. Initial symptoms of
mite feeding on sheep are areas of inflammation with small vesicles, but as the
lesion spreads, the border areas of lesion have moist areas (containing mites), while
the central region is dry covered with a yellow crust. Intense itching associated by
rubbing and scratching of the skin in sheep is found. The wools from infested sheep
become ragged and stained and are shed from large areas. Several mite-infested
sheep are restless and cease to feed; adult sheep may lose weight and are even
debilitated. In livestock, the main effect of the mite infestation is a pruritus caused
by biting and sucking activities of mites which result in the formation of vesicles,
and the exudates dry on the skin to form a crust. Infested cattle are restless, and their
food intake is significantly decreased.
Chorioptes (Family Psoroptidae) These mites are found in cattle, sheep, goats
and equines, sometimes causes serious conditions in the infested animals. Chorioptes
bovis (Hering) is commonly found on these hosts worldwide. Sometimes these
mites have been reported as a separate species: C. bovis, C. ovis and C. equi, mainly
on the bases of hosts infested. But now all these species are considered as C. bovis
only. These mites feed only on the skin surface. In cattle, condition is known as
‘chorioptic mange’ and mainly occurs in the housed animals on the neck, tail head,
udder and leg regions. Lesions caused by mite feeding are localized and spread
slowly. Pruritus is caused by mites resulting in rubbing and scratching. In sheep,
these mites are mainly found on the legs, but cause little harm. In equine hosts,
chorioptic mange occurs as crusty lesions with thickened skin on the legs. Superficial
movements of mites on hosts particularly during the night cause irritation and
restlessness.
Otodectes cynotis (Family Psoroptidae) O. cynotis (Hering) commonly cause

mange in cats and dogs and other small mammals (ferrets and red fox). Mites are
mainly found in the external ear of their hosts, feed superficially and take about 3
weeks for the completion of its life cycle. Signs of irritation in the hosts appear only
23.1 Mites 433
sporadically with the transient activity of the mites. Scratching activity of the mite-
infested cats may cause excoriation of the posterior surface of the ear pinna, and this
with head shaking may result in a haematoma of the ear flap. In dogs, O. cynotis
infestation is a common cause of ‘otitis externa’.
Psorergates ovis (Womersley) (Family Cheyletidae) Psorergates ovis is com-

monly known as ‘itch mite’ in the case of sheep and feeds superficially on the skin.
Mite infestation is common in fine wool breeds such as merino. It is transmitted
through physical contact, but when the fleece lengthens, it serves as a barrier for the
mite transfer. Itch mite mainly attacks young ones of sheep. It takes about 3–4 years
before the whole fleece area is affected. The mite infestation causes chronic irrita-
tion and skin thickening. In severe cases the whole fleece, which is difficult to shear
because of its matted consistency, will have to be discarded. Sometimes in labora-
tory mouse, too much incidence of mite Psorergates simplex is noticed. Other psor-
ergatid mites like P. muricola on rodent (Mus musculus) and P. rattus on Rattus
norvegicus have been reported.
Cheyletiella spp. (Family Cheyletiellidae) C. yasguri (Smiley) on dogs, C. blakei

(Smiley) on cats and C. parasitivorax (Megnin) on domestic rabbits are commonly
reported. These mites are found mainly on the hairs and fur and feed on the skin.
Mite causes mild mange on their hosts but can spread rapidly through kennels. It
also causes characteristic dermatitis which results in shedding of the skin scales into
the hair or fur giving the host a powdery or mealy appearance; presence of moving
mites among this debris has given the common name of ‘walking dandruff’. There
is very little skin reaction or pruritus symptoms.
23.1.10 P
neumonyssus caninum (Chander and Rube) (Family
Halarachnidae)
Pneumonyssus caninum is commonly known as nasal mites. They are parasites of

the nasal cavity and sinuses of dogs. The infection is probably transmitted by direct
nose contact between animals. Infection of mite is associated with head shaking and
inverted sneezing as well as with chronic rhinitis and sinusitis. Treatment with iver-
mectin has been reported effective. Canary lung mite, Sternostoma tracheacolum
(Lawrence) (family Rhinonyssidae), also reported feeding in nasal passages and
tracheal tissues of canaries and domestic birds.
nake Mite (Ophionyssus natricis (Gervais);

23.1.11 S
Macronyssidae)
Snake mites attach to the skin under the scales and feed on blood and fluid. Mite-
infested snake may have severe itchiness, reduced feeding, dysecdysis (abnormal
shed) or the presence of ‘mite dust’ which looks like dandruff or dust on the skin
and scales. Some mite-infested snakes will soak themselves in water bowls to
relieve itchy skin. Snake mites can be seen in the enclosure or hidden under scales,
especially under the eyes and mouth. Mites can crawl from one snake to another if
they are housed together and can also be dispersed accidentally by people handling
snakes in enclosures or zoos (ER 24). These mites also reported on captive lizards,
turtles and crocodiles. Snake mites have spread from worldwide herpetoculture, and
the mite is a major pest of captive snakes in zoos. These mites have also been shown
to act as vectors for Inclusion Body Disease (IBD). Mite infestation in humans has
also been reported. Life cycle of snake mite takes about 13–19 days at 24°–29 °C
and 70–90 % relative humidity. A female mite lays 60–80 eggs in her life. So any
natural sticks, logs and previously infected enclosures that have not been kept
snake-free for 7 weeks can occasionally harbour mites or their eggs from the wild
and then infect the new reptiles. For controlling snake mites, pyrethrin sprays or
washes have been reported effective. Administration of ivermectin under supervi-
sion of trained veterinary doctor can be quite effective in controlling mites.
23.1.12 House Dust Mites
Dust mites are found in almost every home where these live in dust which accumu-
lates in carpets, bedding, fabrics and furniture. The principal dust mite species are
Dermatophagoides pteronyssinus (Troussart), D. farinae (Hughes) and Euroglyphus
maynei (Cooreman), among the top three species in terms of global frequency and
abundance (all belonging to family Pyroglyphidae). But in tropics and subtropics,
Blomia tropicalis (van Bronswijk, de Cock and Oshima) belonging to family
Echimyopodidae also has emerged as an important dust mite. Glycyphagus spp. and
Lepidoglyphus spp. may also be commonly found in the house dust (Arlian 1989a,
b; Revsbech and Dueholm 1990). There can be as many as 1000 mites in one
gramme of house dust. These mites feed on dead skin and are a major problem
because they can cause wheezing, itching, other allergic reactions and possibly
asthma (ER11 and ER22).
The life cycle for a male house dust mite is 10–19 days, and for a mated female,
it may be up to 70 days. Each female lays 60–100 eggs in her life. Optimum condi-
tions under laboratory conditions for development of dust mites are found to be
25°–30 °C and 75–80 % relative humidity. In northern Europe, dust mite population
is highest in late summer and autumn and least during winter months. Places with
damper climates are more favoured by mites than dry ones; thus, allergy to mites
tends to be rarer among people living in continental interiors and mountainous
regions.
Allergens from house dust mites are the most common (besides allergens from
domestic pets and cockroaches) ubiquitous allergens to which people are exposed
and become sensitized. Dusting, vacuuming, bed making or any other activity at
homes causes settled dust to become airborne, and the faecal pellets and smaller
allergen-bearing particles become temporarily suspended in the air. The airborne
faecal pellets may be inhaled, and people who are atopic, i.e. are genetically
23.2 Ticks 435
predisposed to develop allergic reactions to common allergens from pollens, dust

mites and animal skin scales, respond to this exposure either by antibodies which
then bind with immunologically active cells to mediate the release of mediators
such as histamine and the development of localized inflammation. The allergic reac-
tions are manifested as asthma, eczema, rhinitis and conjunctivitis. Roughly speak-
ing, 1–2 % of the world population suffers from allergy to house dust mites. Allergen
exposure in bedrooms can be reduced by using a mattress cover, replacing old pil-
lows and, if possible, by removing the carpet.
23.1.13 Other Important Mites
Parasitic mites, Myobia musculi (Schrank) and Radfordia affinis (Poppe) (both of
family Myobiidae), occur on the laboratory mouse and its wild progenitor –the
house mouse– and may parasitize other laboratory animals. R. ensifera (Poppe)
attacks laboratory pets. Eggs of these mites are usually laid attached to the bases of
the hosts’ hairs. Life cycle of myobiids is generally quite brief (about 2 weeks), and
the mites freely move between host individuals. Mite infestations are usually low in
intensity in wild mammal hosts and of little or no consequence, but on laboratory
rodents, they are frequently found in high intensity and may result in tense pruritus
and hair loss known as myobic mange (Mullen 2002). Hair-clasping mites of super-
family Listrophoroidea have legs specialized for clasping a hair, for example,
Chirodiscoides caviae (Fain) – a parasite of guinea pigs; and Myocoptes musculinus
(Koch) is a parasite of rodents. Lynxacarus radovskyi (Fain) is a hair-clasping mite
of domestic cat. Trixacarus caviae (Fain et al., Hovell and Hyatt), which resembles
Sarcoptes scabiei, is a parasite of the guinea pigs and causes pruritus so intense that
guinea pigs are subject to fits and seizures because of vigorous scratching or manip-
ulation of the skin. Some members of genera Cosarcoptes, Prosarcoptes, and
Pithesarcoptes (family Cercopithecidae) are parasitic on Old World monkeys.
These mites resemble Sarcoptes morphologically, biologically and pathogeneti-
cally. Pneumonyssus simicola (Banks) have been reported in lung parenchyma of
monkeys, Macaca mulatta, causing lesions scattered throughout the lungs. Monkeys
can be reared free of P. simicola mites if they are separated from birth and reared in
isolation from adult monkeys (Kettle 1990).
23.2 Ticks
Ticks comprise a distinct group of exclusively blood-feeding ectoparasites and are

very familiar to most people in virtually all regions of the world (ER12, ER13 and
ER18). Ticks transmit a greater variety of disease-causing pathogenic agents includ-
ing protozoan, viral, bacterial and even fungal pathogens. Tick-borne diseases of
livestock such as babesiosis, theileriosis, and heartwater have made it difficult or
impossible to raise domestic animals for food or animal products (ER17 and ER15).
Infestation and large-scale feeding by ticks can also cause irritating or even fatal
injury to humans and animals because of paralysis, toxicity or severe allergic reac-
tions to their bites (ER14, ER20 and ER23). In view of the enormous literature
available on different ticks and the variety of diseases caused by tick-borne patho-
gens and injurious substances injected, only general account of important tick-borne
diseases is given below.
Babesiosis Tick species, Ixodes scapularis (Say) and I. ricinus (Linnaeus), are
vectored by the protozoans, Babesia microti, B. divergens and B. gibsoni, in human
beings. In cattle, tick species Boophilus annulatus (Say) and B. microplus
(Canestrini) cause bovine babesiosis (Angus 1966).
Tularaemia Tularaemia is caused by bacteria, Francisella tularensis, and is vec-

tored by tick Haemaphysalis leporispalustris (Packard) in humans and various
other animals.
Lyme Disease Lyme disease is caused by bacterium, Borrelia burgdorferi, and is

vectored by tick Ixodes scapularis and I. ricinus and other species in humans, dogs,
cats and other animals.
Avian Spirochaetosis Avian spirochaetosis is caused by bacterium, Borrelia anse-

rina, and vectored by tick Argas persicus (Oken) in birds.
Rocky Mountain Fever Rocky Mountain fever is caused by bacterium, Rickettsia

rickettsii, in humans and is vectored by ticks, Dermacentor variabilis (Say), D.
andersoni (Stiles) and other tick species.
Canine Ehrlichiosis In dogs this disease is caused by bacterium, Ehrlichia canis,

and vectored by tick species, Rhipicephalus sanguineus, I. ricinus and Amblyomma
americanum (Linnaeus).
Heartwater Heartwater is caused by bacterium, Cowdria ruminantium, in rumi-

nants, and is vectored by ticks Amblyomma herbraeum (Koch), A. variegatum
(Fabricius) and others.
Q Fever Q fever is caused by bacterium, Coxiella burnetii, in humans and large

livestock. This disease is vectored by many tick species.
Colorado Tick Fever In rodents, domestic animals and humans, this disease is
caused by arboviruses which are vectored by tick species, I. ricinus.
African Swine Fever African swine fever is caused by Iridovirus and is transmit-
ted by ticks, Ornithodorus moubata (Murray), in domestic pigs and wild bears.
Ticks in addition to feeding injuries also inject tick proteins into the body of the
victims resulting in symptoms known as ‘tick paralysis’, tick-bite allergies and
sweating sickness and some other tick toxicosis symptoms in human, cattle, sheep
and some other mammals. Ixodes holocyclus, Dermacentor variabilis, D. ander-
soni, and Argas persicus are important tick species. Complex of problems related to
ticks and tick borne diseases of cattle and human beings have created a demand for
methods to control ticks and reduce their loss in human beings and cattle. Controlling
of tick infestations and the transmission of tick-borne diseases will remain a great
challenge in the future for the cattle industry in tropical and subtropical areas of the
world (Sonenshine and Roe 2013). So tick control will remain as a priority in these
regions considering the quantum of losses caused in these belts, and losses by ticks
can be cut considerably by adopting effective control measures.
23.3 Conclusions
More than 250 species of mites and ticks are recognized as the cause of health-
related problems for humans and domestic animals. Feeding by mites and ticks are
known to cause problems ranging from temporary irritation of the skin to persistent
dermatitis, mite-induced allergies and transmission of pathogenic microbial agents
to invasions of respiratory passages and ear canals. Chicken mite (Dermanyssus
gallinae) and northern fowl mite (Ornithonyssus sylviarum) are parasites of poultry
and other birds resulting in severe dermatitis in some cases. Some mites
(Ornithonyssus bacoti and Liponyssoides sanguineus) infest particularly rats and
mice and also are a nuisance to humans. Grain mites (Acarus siro) also serve as a
nuisance to personnels handling grains and result in symptoms like ‘copra itch’ or
‘grocer’s itch’; straw itch mites (Pyemotes ventricosus) infest insect larva and some-
times wipe out insect cultures. Burrowing mite (Sarcoptes scabiei, Notoedres spp.
and Knemidocoptes spp.) infestation results in scabies in human beings and sarcop-
tic mange in cattle, goat, sheep, rabbits and poultry birds. Non-burrowing mite
(Psoroptes spp., Chorioptes spp. and Otodectes spp.) infestations are commonly
found on the skin of cattle, cats, dogs and foxes resulting in many types of skin
irritations and reducing the quality of fur. House dust mites (Dermatophagoides
pteronyssinus and D. farina) which live in dusts in carpets, beddings, furniture, etc.
cause various types of allergic reactions and asthma (Willadren 2001).
Ticks comprise an exclusively blood-feeding ectoparasites of humans and other
mammals and transmit a greater variety of disease causing pathogenic agents.
Babesiosis, theileriosis and heartwater are important tick-borne diseases of cattle
resulting in economic losses in production of food/other products from cattle, sheep,
goats, horses, etc. Some disease caused by a bacterium Borrelia burgdorferi is vec-
tored by ticks on humans, dogs, cattle and other animals. Rocky Mountain fever
caused by bacterium Rickettsia rickettsii is also vectored by ticks Dermacentor
variabilis and D. andersoni in humans. African swine fever caused by Iridovirus is
also transmitted by ticks, Ornithodorus moubata, in domestic pigs and wild bears
(Oliver 1989; Sonenshine and Roe 2013; Tatchell 1992).
Different measures for management of various complications involved with par-

asitism by different ticks and mites on humans, pets, cattle, poultry birds, goat,
sheep and pig rearing are suggested in the account of mites given (Solomon 1983).
Further Reading
Angus BM (1966) The history of the cattle tick, Boophilus microplus in Australia and achieve-
ments in its control. Aust Soc Parasitol 26:1341–1355
Arlian LG (1989a) Biology and ecology of house dust mites, Dermatophagoides spp. and
Euroglyphus spp. Immunol Allergy Clin North Am 9:339–356
Arlian LG (1989b) Biology, host relations, and epidemiology of Sarcoptes scabiei. Annu Rev
Entomol 34:139–161
Axtel RC, Arends JJ (1990) Ecology and management of arthropod pests of poultry. Annu Rev
Entomol 135:101–121
Burgess I (1994) Sarcoptes scabiei and scabies. Adv Parasitol 33:235–292
Evans GO (1992) Principles of acarology. CABI, Walligford
Fuller HS (1956) Veterinary and medical acarology. Annu Rev Entomol 1:347–366
Kettle DS (1990) Medical and veterinary entomology. CAB Internatioal, Wallingford
Mullen G (2002) Mites (Acari). In: Medical and veterinary entomology. Elsevier, New York,
pp 449–474
Oliver JH (1989) Biology and systematic of ticks. Annu Rev Ecol Syst 20:397–420
Revsbech P, Anderson G (1987) Storage mite allergy among grain elevator workers. Allergy
42:423–429
Revsbech P, Dueholm M (1990) Storage mite allergy among bakers. Allergy 45:204–208
Solomon KR (1983) Acaricide resistance in ticks. Adv Vet Sci Comp Med 27:273–296
Sonenshine DE, Roe RM (2013) Biology of ticks. Oxford University Press, New York
Steelman CD (1976) Effects of external and internal arthropod parasites on domestic livestock
production. Annu Rev Entomol 21:155–178
Tatchell RJ (1992) Ecology in relation to integrated tick management. Insect Sci Appl
13:511–561
Willadren P (2001) The molecular revolution in the development of vaccines against ectoparasites.
Vet Parisitol 101:353–367
Wooley TA (1988) Acarology: mites and human welfare. Wiley, New York
1. http://www.cdc.gov/nceh/ehs/Docs/Pictorial_Keys/Acarina.pdf Acarina: illustrated key to

some common adult female mites and adult
2. http://www.cdc.gov/nceh/ehs/Docs/Pictorial_Keys/Ticks.pdf Ticks: key to Genera in United
States
3.
h t t p : / / d i g i t a l c o m m o n s . g e o r g i a s o u t h e r n . e d u / c g i / v i e w c o n t e n t .
cgi?article=1119&context=honors-theses Identification guide to larval stages of ticks of medi-
cal importance in the USA (Many SEMs of ticks)
4. https://www.youtube.com/watch?v=EJvfDMUnQU8 Human parasites; scabies! (Norwegian
scabies, stories, news, and more)
5. https://www.youtube.com/watch?v=h5LCwiEsR68 The human itch mite (Scabies mite)
6. https://www.youtube.com/watch?v=GUjU3T55-co Dermanyssus gallinae (Red mite, chicken
mite)
7. https://www.youtube.com/watch?v=a6wT8hOcYwQ Rodent mites (Ornithonyssus bacoti)
Further Reading 439
8. https://www.youtube.com/watch?v=lc5DaJnM01w Red velvet mites

9. https://www.youtube.com/watch?v=KXFa2q-h2L4 Feline Notoedric mange mites
10. https://www.youtube.com/watch?v=QFgcyns95TE Grain itch mite Pediculoides ventricosus
11. https://www.youtube.com/watch?v=hyP6czmysXI Cannibal mites (Dust mites)
12. https://www.youtube.com/watch?v=xDSwXsKhOMo Tick on tick
13. h ttps://www.youtube.com/watch?v=WpOyWesQtgc&list=PLlRvaTnNBxia9NlhGfdI5-
zgytIv75qbG&index=36 Baxter healthcare tick animation
14. https://www.youtube.com/watch?v=27McsguL2Og How to properly remove a tick
15. https://www.youtube.com/watch?v=0g_lt0FcQag Tick Lifecycle.mp4
16. https://www.youtube.com/watch?v=hlleWRp_13I The Worst Tick infestations ever
17. https://www.youtube.com/watch?v=QM0kBwSU3hc&index=145&list=PLlRvaTnNBxia9Nl
hGfdI5-zgytIv75qbG Tick borne diseases of cattle
18. https://www.youtube.com/watch?v=p7KpmrQ9DYM&index=176&list=PLlRvaTnNBxia9Nl
hGfdI5-zgytIv75qbG Tick meat allergy
19. https://www.youtube.com/watch?v=m9H3lnRaiXA 3 Reasons why ticks suck
20. https://www.youtube.com/watch?v=eATpttq6DfA&index=198&list=PLlRvaTnNBxia9NlhG
fdI5-zgytIv75qbG The Tick key
21. https://www.youtube.com/watch?v=sgav_kZ_Hi4 Demodex blepharitis – is it for real?
22. https://www.youtube.com/watch?v=qEvW3ZGAAg8 Dust mite
23. http://www.afrivip.org/sites/default/files/identification_complete_1.pdf Ticks: tick identifica
tion
24. http://www.reptilesmagazine.com/Snakes/Snake-Health-101/Causes-Prevention-and-Treatme
nt-of-Snake-Mites-Acariasis/ Causes, prevention, and treatment of snake mites, acariasis
25. http://www.dgvd.org/media/news/publikationen/2009/ddg_09094_eng.pdf Tropical rat mites
(Ornithonyssus bacoti) – serious ectoparasites
Forensic Acarology
24
Abstract
Mites can be found pretty much everywhere in our houses and furnishings, on
our clothes and even in the pores of our skin. Forensic acarology is the name
given to the study of mites (including ticks) that forms part of the evidence in
legal cases, but primarily associated with death enquiries. As mites are so wide-
spread, there are few situations in which people and different objects associated
with the crime will not be exposed to them, and they could serve as important
trace evidence in forensic investigations. Flies along with their phoretic mites are
the first scavengers colonizing a dead body. During butyric fermentation of
corpse and advanced stages of decay, large numbers of mites are attracted to the
corpse and may be visible to the naked eye. Large quantities of mites give a fluffy
appearance to the decomposing body. More than 100 species of mites from over
60 families have been reported on animal carcases, but 75 mite species distrib-
uted in 20 families are also reported on human corpses. Representatives of fami-
lies Parasitidae, Macrochelidae, Ascidae, Demodicidae, Acaridae,
Histiostomatidae and Uropodidae are dominantly reported on dead bodies.
Besides Mesostigmata, sometimes members of Astigmata and Prostigmata are
also reported on carcases.
Keywords
Forensic acarology • Butyric fermentation • Corpse • Criminolegal • Carcases
Mites can be found pretty much everywhere in our houses and furnishings, on our
clothes and even in the pores of our skin. They are microhabitat specific and might
provide evidential data on the movement or relocation of bodies or locating a sus-
pect at the scene of a crime. Forensic acarology is the name given to the study of
mites (including ticks) that forms part of the evidence in legal cases, but is primarily
associated with death enquiries. Because of their high diversity, wide occurrence
and abundance, mites may be of great value in the analysis of trace evidence (ER2

DOI 10.1007/978-981-10-1594-6_24
442 24 Forensic Acarology
and ER3). So a suspect can leave behind not only his or her own skin mites but also
soil mites attached to the shoes, socks and the lower parts of trousers (Baker 1999;
Coloff 2009; Hughes 1976).
Most criminology investigations that explore trace evidence and analyze post-
mortem intervals 48 h after death include the study of arthropods. Usually larger
insects are mostly reported in and around a body or at the scene of a crime but usu-
ally mites which are also found are not preserved. Because mites are so widespread,
there are few situations in which people and different objects associated with the
crime will not be exposed to them, and they could serve as important trace evidence
in forensic investigations (Coloff 2009; OConnor 2009; Smith 1986).
The first arthropod scavengers colonizing a dead body are flies along with their
phoretic mites. The flies complete their life cycle in and around the corpse, while
the mites may feed on the immature stages of the flies (Perotti 2009a, b). Considering
the role of mites in forensic investigations, a special issue of the journal Experimental
and Applied Acarology was published in 2009 which was devoted entirely to the
forensic acarology and included 11 review articles on different aspects. Review
articles included – Introduction of forensic acarology; Early post-mortem changes
and stages of decomposition in exposed cadavers; Megnin re-analyzed: The case of
new-born baby girl, Paris, 1818; Carcasses and mites; Phoretic mites associated
with animal and human decomposition; Astigmatid mites of forensic interest;
Indoor mites and forensic acarology; Human hair follicle mites and forensic acarol-
ogy; Acari in Archaeology; and Can fresh water mites act as forensic tools? – were
published including a lot of earlier published references in literature. Most of the
salient findings reported in above publications are discussed in this chapter.
24.1 Decomposition of Cadaver
Decomposition of an exposed cadaver is a continuous process beginning at the

moment of death and ending when the body is reduced to a dried skeleton. Traditional
estimates of the period of time since death or post-mortem interval have been based
on a series of grossly observable changes to the body including lower mortis, algor
mortis, rigor mortis and similar phenomenon. But in recent years, insects, mites and
other arthropods have been increasingly used by the law enforcement agencies of
different countries to provide an estimate of the post-mortem interval. Although the
process of decomposition of exposed cadaver is continuous, it is useful to divide this
into a series of five stages, namely, fresh, bloated, decay, post-decay and skeletal
(Goff 2009).
Typically there are two points known just before beginning the task of estimating
a period of time since death: the last time the individual was reliably known to be
alive and the time at which the body was discovered. The death occurred between
these two points and the aim is to estimate when it is generally accepted that there
is actually no scientific way to precisely determine the exact period of time since
death. In forensic entomology and acarology, an estimation of the period of arthro-
pod activity on the body is studied. The period of activity will reflect the minimum
24.2 Decomposition Stages of Corpse 443
period of the time since death or post-mortem interval (PMI), but it will not pre-
cisely determine in most of the cases; the latter point is more accurately known than
the former (Perotti 2009a, b; Rasmy 2009).
24.2 Decomposition Stages of Corpse
In the current literature, the state of the carcase is described by a stage of decompo-
sition rather than by a wave of arthropod colonization. Generally five states of decay
have been recognized for exposed and concealed carcases (ER1):
(i) Initial decay, fresh state

(ii) Putrefaction, bloated stage
(iii) Black putrefaction, active decay, decay stage
(iv) Butyric fermentation, advanced decay
(v) Dry decay, dry decomposition, skeletal or remains stage
During butyric fermentation and advanced decay stages, large numbers of mites
are attracted to the corpse, and mites at these stages become visible to the naked eye.
Sometimes they are mistaken for mould which is also present at this stage. Large
quantities of the mites give a fluffy appearance to the decomposition of pigs, and in
cases of dogs, mites have sometimes been found in large numbers on the upper
surface of the carcases. In case of guinea pigs where any skin is left by the skin feed-
ers of the previous stage, an immense number of tyroglyphid mites consumed the
remainder skin leaving nothing but bones. Numerous species of mites are found on
both exposed and buried corpses with coffin in a grave buried three feet deep. But
mites were not found in large numbers on the submerged carcases (Megnin 1894;
Perotti et al. 2009; Turner 2009).
Mites have been reported during decomposition of corpses (active and advanced
decomposition, dry remains) of lizards, toads, guinea pigs and chickens during dif-
ferent stages of decomposition. Water mites are also reported associated with sub-
merged pigs, wild bear, alligators and deer. Mites have also been reported at crime
scenes or associated with the human corpses.
Some mite species will end up at a carcase as incidentals, as species that use the
corpse as a concentrated resource extension of their habitat; however, mites as inci-
dentals might be a minority group. Many mite species arrive at a carcase through
phoresy on a necrophagous or necrophilous insects. The phoresy is often highly
taxon specific. Many mite species arriving by phoresy are likely the product of evo-
lutionary adaptation to a specialized food source and habitat, the opposite of inci-
dental. But if mites are incidental, they might become the centre point of trace
analysis in a forensic setting. In some oligospecific infestations, only mites of the
family Acaridae (Tyroglyphidae) have been reported (OConnor 2009; Perotti et al.
2009).
Symbiotic mites, Demodex brevis (Akbulatova) and Demodex folliculorum
(Simon) (family Demodicidae), feeding on sebaceous or fat glands and hair
follicles, have also been reported on human corpses. Within the decomposition
stages, several mite species belonging to different families of Astigmata,
Mesostigmata and Prostigmata are reported on human corpses.
24.3 Arthropods and Decomposing Body
Four basic relationships between a decomposing body and different arthropods are
established:
(i) Necrophagous species. Feeds actually on the corpse and are the early invaders.
Blow flies (Calliphoridae) and flesh flies (Sarcophagidae) of insect order
Diptera are the early invaders. Acari (Acaridae) are typically not found playing
any role at this stage.
(ii) Predators and parasites of necrophagous species. Many of the beetles
(Silphidae, Staphylinidae and Histeridae), and dipterous family Calliphoridae,
are parasitic on fly larvae and pupae. The Acari, primarily Acaridae, are not
active but were present at this stage.
(iii) Omnivorous species. Ants, wasps, some beetles and some associated arthro-
pods that feed on the corpse.
(iv) Adventive species. Some springtails, spiders, centipedes and millipedes may
also be commonly feeding on the corpse.
(v) Accidentals. Some insect species are accidentally falling on the corpse from
the surrounding vegetation.
24.4 Megnin and Forensic Acarology
At the end of the nineteenth century, the work of Jean Pierre Megnin in Paris
(France) proved to be the starting point of forensic acarology. Megnin documented
his observations in La Faune des Cadavres (The Fauna of Carcasses). He was the
first to list eight distinct waves of arthropods colonizing human carcases. His first
wave included flies and mites, and his sixth wave was composed of mites exclu-
sively when the desiccation of exposed human corpse had started (5–12 months
after death). Megnin reported Mesostigmata (Uropodidae and Trachytidae) and
Astigmata (Acaridae, Histiostomatidae and Glycyphagidae) mites at this stage.
Megnin did not report any mites in the seventh wave (Megnin 1894; Perotti
2009a, b).
24.5 Mite Diversity Found on Carcases
More than 100 species of mites from over 60 mite families have been reported on
different animal carcases, but 75 mite species distributed in 20 families are also
reported on human corpses. Large populations of phoretic mites are sometimes
24.5 Mite Diversity Found on Carcases 445
found in abundance on carcases and corpses and may outnumber their scavenger
carrier in both number and diversity. Many phoretic mites travel on scavenger
insects and are highly specific (Desch 2009). They are usually found in a specialized
transitional transport or dispersal stage, often moulting and transforming to adults
shortly after arrival on a corpse or carcase. Many species are having faster develop-
ment and generation cycles than their carriers; more than 200 phoretic mite species
are reported associated with corpses in literature (Perotti 2009a, b; Perotti et al.
2009; Rasmy 2009).
More than 12 mite species belonging to Mesostigmata (mostly Parasitidae and
Macrochelidae) are dominantly reported than mites in other groups. Majority of the
mesostigmatans seems to be associated with particular species of fly or carrion bee-
tles. Prostigmatans (Demodex spp., Demodicidae) are reported during the first stage
or during early stages of decomposition. But no mite species are found during putre-
faction and bloating stage and putrefaction stages of terrestrial and freshwater con-
ditions. However, during butyric fermentation or advanced stages, mites belonging
to mesostigmatid families Ascidae (mainly Proctolaelaps spp., Hypoaspis spp.,
Zerconopsis spp.), Macrochelidae (Glypholaspis spp., Macrocheles spp.),
Parasitidae (Pergamasus spp., Poecilochirus spp.) and Uropodidae (Urobovella
spp.), Astigmata families Histiostomatidae and Acaridae (Acarus spp., Sancassania
spp.) and Oribatida families Caneisiidae and Mycobatidae were commonly found
on human corpses. On human corpses and animal carcases mainly representatives
of Astigmata, Mesostigmata and Prostigmata are found. But during dry decomposi-
tion of the human corpse, stage members of mesostigmatid mites (Dinychidae,
Laelapidae, Macrochelidae, Uropodidae); members of astigmatid mites, Acaridae
(Acarus spp., Tyrophagus spp., Sancassania spp.,), Glycyphagidae, Histiostomatidae
and Lardoglyphidae; and members of prostigmatid family Cheyletidae (Cheyletus
spp.) have been reported.
During dry decays about 52 species of the Astigmata are also reported as pho-
retic on scavengers, the majority travelling on late-arriving scavengers such as hide
beetles, skin beetles and moths. Further several species of carrion beetles may visit
a corpse simultaneously, and each beetle may carry 1–10 species of phoretic mites.
The composition of the phoretic mite assemblage on a carcase might provide valu-
able information about the conditions of and time elapsed since death.
Studies on many mite species belonging to different families, their abundance,
hosts, location, habitat and season in which have been found on birds (chickens,
crows, sparrows, etc.), mammals (pigs, cats, dogs, rats and mice, rabbits, fox, guinea
pig, bear, deer), fishes, frogs, alligator and turtles have been reported in literature.
Follicle mites may occur on canine and primate bodies. Phoretic and parasitic mites
of 26 families have been reported from adult trogid beetles and piophilid flies and
on dung beetles.
24.6 Role of Freshwater Mites
Most freshwater mites are predators, detritivores or algivores and are hence unlikely
candidates as tools for ageing or determining original placement of corpses or other
bodily remains. However, some aquatic Astigmata have occasionally been reported
feeding on the tissues of moribund aquatic animals (Proctor 2009; Walter and
Proctor 1999). Based on their biology, there is little chance that freshwater mites
may be of great value as forensic tools. Representatives of four major lineages of
mites are found in or on the surface of freshwater. Mesostigmata (more than 25 spe-
cies); Prostigmata, including ‘true’ water mites (Hydrachnidia) with more than
5000 species, and Halacaridae with about 150 freshwater species; Oribatida with
about 100 species; and Astigmata with about 100 species are known. Semiaquatic
microstigmatans of family Ascidae (Cheiroseius spp. and Platyseius spp.) are resi-
dents of standing or slowly moving water. These mites are non-swimming and usu-
ally occupy the surface of film rather than deeply submerged substrates.
Semiaquatic Mesostigmata are mainly predators of nematodes and small arthro-
pods and can potentially be attracted to saprophagous invertebrates that themselves
are feeding on corpse tissues near the water surface. Water mites also known as
Hydrachnidia or Hydracarina are more ecologically diverse than aquatic mesostig-
matans. They are present in both standing and running water, shallow and deep, and
there are swimming and non-swimming species. Water mites are parasitic on insects
as larvae and also predatory as deutonymphs and adults. There are no records of
water mites consuming dead vertebrate flesh that complete their life cycle within the
carrier, and it seems very unlikely that they would feed directly on a corpse.
However, like the mesostigmatans, it is possible that they can be attracted to
saprophagous invertebrates.
24.7 Conclusions
Mites can be found everywhere in our houses and furnishings, on our clothes and
even in the pores of the skin; they are microhabitat specific. Because of their high
diversity, wide occurrence and abundance, mites may be of great value in the analy-
sis of trace evidence the suspect can leave behind, and that information can be used
to determine time of death or provide vital clues about a suspect and the circum-
stance (Solarz 2009; Turner 2009). No detailed studies on critical relationship of
phoretic mites found on corpses are available, but such studies can provide traces of
information useful for crime investigations. Molecular biology studies on different
mites found on corpses may be very useful for knowing exactly the identification
and their characterizations. However, for producing any suggestions/evidence, in
this direction there should be close cooperation among entomologists, acarologists
and forensic experts dealing with the cases under different situations. Priority should
be given for production of user-friendly aids like picture-based keys of corpse-
associated mites covering all stages of decomposition of corpses in different situa-
tions. The European Association for Forensic Entomology (EAFE) – an international
Further Reading 447
body of physicians and scientists – compiled a list of best practice standards in

forensic entomology. More awakening should be created among entomologists and
acarologists about mite diversity found, and their role regarding decomposition of
corpses at different stages or waves by offering teaching courses in forensic ento-
mology and acarology, and researches in this respect be strengthened at postgradu-
ate levels. Special symposia on forensic entomology and acarology need to be
organized in different countries for dissemination of recent knowledge in this
direction.
Further Reading
Amendt J, Campobosso CP, Gaudry E, Roitr C, LeBlank HN (2006) Pests practiced in forensic
entomology-standards and guidelines (Electronic Version). Int J Leg Med (online version).
http://www.springerlink.com/content/1726y387131190683
Arlian LG (2009) Arthropod allergens and human health. Annu Rev Entomol 47:395–433
Baker AS (1999) Mites and ticks of domestic animals: an identification guide and information
source. Natural History Museum, London
Braig HR, Perotti MA (2009) Carcases and mites. Exp Appl Acarol 49:45–84.
doi:10.1007/510493-009-9287-6
Coloff MJ (2009) Dust mites. CSIRO Publishing, Collingwood, 600pp. ISBN 9780643065895
Desch CE (2009) Human hair follicle mites and forensic acarology. Exp Appl Acarol 49:143–145.
doi:10.1007/510493-009-9272-0
Goff ML (2009) Early post-mortem changes and stages of decomposition in exposed cadavers.
Exp Appl Acarol 49:21–36. doi:10.1007/s10493-009-9284-9
Hughes AM (1976) The mites of stored food and Houses, Technical Bulletin. Ministry of
Agriculture, Fisheries and Food, London, 400 pp
Megnin P (1894) The fauna of corpses. Application of entomology to forensic medicine. G. Masson
and Gauther-Villars et Fils, Paris
Merritt RW, Wallace JR (2001) The role of aquatic insects in forensic investigations. In: Byrd JH,
Carter JL (eds) Forensic entomology. CRC Press, Boca Raton, pp 177–215
OConnor BM (2009) Astigmatid mites (Acari: Sarcoptiformes) of forensic interest. Exp Appl
Acarol 49:125–133
Perotti MA (2009a) Forensic Acarology. Exp Appl Acarol 49:1–2
Perotti MA (2009b) Megnin re-analysed: the case of newborn baby girl, Paris 1878. Exp Appl
Acarol 49:37–44
Perotti MA, Braig HR (2009) Phoretic mites associated with animal and human decomposition.
Perotti MA, Braig HR, Goff ML (2009) Phoretic mites and carcases. In: Amenth J, Gof ML,
Compobosso CP (eds) Current concepts in forensic entomology. Springer, Dordrecht
Proctor HC (2009) Can freshwater mites act as forensic tools. Exp Appl Acarol 49:161–165.
doi:10.1007/s10493-009-9274-Y
Rasmy A (2009) Forensic Acarology. A new area for forensic investigation. Acarines 3:1–3
Smith RGV (1986) A manual of forensic entomology. British Museum (Natural History), London,
205 pp
Solarz K (2009) Indoor mites and forensic acarology. Exp Appl Acarol 49:135–142
Turner B (2009) Forensic entomology: a template for forensic acarology? Exp Appl Acarol 49:15–
20. doi:10.1007/s10493-009-9274-Y
Walter DE, Proctor HC (1999) Mites: ecology, evolution and behaviour. University of New
Southwales Press/CAB International, Sydney/Wallingford

1. https://www.youtube.com/watch?v=OFJrow7yaec The fascinating process of human
decomposition
2. https://zoolology.wordpress.com/2012/12/31/forensic-acarology/ Forensic acarology
3. http://dergipark.ulakbim.gov.tr/entoteb/article/view/1014000067 A new branch in acarology;
Forensic acarology
Accidental Acarophagy
25
Abstract
Considering the great abundance of mites in almost all habitats and their minute
size, acarophagy must be common and even humans must be eating lots of mites
accidentally through different ways (foods and food products). The highest num-
ber of mites (both active and dormant forms) is found on fruits and vegetables
when they are brought into markets for wholesale and retail sales. Plant-feeding
mites belonging to families Tetranychidae, Tenuipalpidae, Tarsonemidae and
Eriophyidae are many times ingested by eating mite-infested vegetables, fruits
and fodder crops. Pancake syndrome or oral mite anaphylaxis (OMA) is charac-
terized by severe allergic manifestations found in atopic patients, shortly after
the intake of foods made with mite-contaminated wheat flour. OMA is found
more frequently in tropical and subtropical environments and is more often trig-
gered by pancakes. House dust mites and mites infesting grains are commonly
present in the flours producing OMA. Sometimes pulmonary and intestinal aca-
riasis is often reported in human beings and animals after ingestion of mite-
infested stored grains and other stored products including animal feed and
through too much inhalation of mites in persons handling such foods for longer
durations.
Keywords
Acarophagy • Pancake syndrome • House dust mites • Stored products
Considering the great abundance of mites in almost all habitats and their minute
dimensions, acarophagy must be common, and even humans must be eating lots of
mites accidentally through different ways (foods and food products). The highest
number of mites (both active and dormant forms) is found on fruits and vegetables
when they are brought into markets for wholesale and retail sales (Skubala et al.
2006; Thind and Clarke 2001; Walter and Proctor 1999). On examination under
magnifying glass (10x or more) or under microscopes, one can see different groups

DOI 10.1007/978-981-10-1594-6_25
450 25 Accidental Acarophagy
of mites representing both the superorders Parasitiformes and Acariformes. In many

scattered reports, various intestinal disorders have been reported in human beings
and animals after ingestion of mite-infested fruits and vegetables, mite-infested
stored wheat flour, processed foods and several animal feeds, mushrooms and
cheese (Balance et al. 1997; Demerrel et al. 2004; Szilman et al. 2006). Salient
available information regarding the above aspects is presented below.
25.1 Plant-Associated Mites
Mite infestations of many plant-feeding mites belonging to families Tetranychidae,

Tenuipalpidae, Tarsonemidae, Eriophyidae and Tuckerellidae are many times found
on different vegetable crops, fruit trees and fodder crops. Besides many other repre-
sentatives of plant-associated mites belonging mainly to families Cheyletidae,
Cunaxidae, Phytoseiidae and Stigmaeidae are also commonly associated with plant-
feeding mites (Skubala et al. 2006). Whenever human beings or animals eat fruits,
vegetables and fodder crops directly after picking from plants, they might consume
the mites along with the fruits and vegetables. Direct consumption of guava, rasp-
berries, strawberries, redcurrants, leafy vegetables (like spinach), okra fruits, musk-
melon (cantaloupes), long melon and cucumber may also result in too much
ingestion of mites along with the fruits. However, reports have revealed that about
50 % of mites are removed if the mite-infested fruits/vegetables are washed in run-
ning water, but eggs (glued to the surface) and dormant mites are not properly
removed even after washing. But peeling off of outer skin definitely helps in remov-
ing mites and reducing risks involved by their ingestion. Fruits which have uneven
surfaces on them are more preferred for the development and multiplication of
mites. Even the sepals and leaves which accompany the fruits at marketing stages
are also highly favourable for multiplication of mites.
From Poland, in one study 90 different food samples representing different spe-
cies of fruits, vegetables and mushrooms that were purchased from different city
markets and examined under microscopes and magnifying glasses revealed the exis-
tence of 53 different species of mites on them. Leafy vegetables like spinach, ama-
ranth, mustard and fenugreek leaves which are commonly infested by different mite
species are eaten in some cases directly without cooking, and in this way mites may
be ingested along with the leaves (ER4 and ER5). Animals which graze from fields
on various fodder crops (sorghum, maize, berseem, etc.) may also ingest several
mites from infested plants. Even the fruits like apple stored in refrigerators/cold
storages, when taken from big stores and supermarkets, revealed several plant-
associated mites. One of the study conducted in Poland revealed 22 species of mites
belonging to different families on stored apples alone (Chmielewski 1998).
However, the open and intact apple skin has normally been found free of mites.
25.2 Stored Food 451
25.2 Stored Food
25.2.1 Stored Wheat Flour
25.2.1.1 Pancake Syndrome or Oral Mite Anaphylaxis (OMA)

This is a new syndrome characterized by severe allergic manifestations occurring in
atopic patients shortly after the intake of foods made with mite-contaminated wheat
flour. The clinical entity, observed more frequently in tropical/subtropical environ-
ments, is more often triggered by pancakes, and for that reason it has been desig-
nated as ‘pancake syndrome’. Because cooked foods are able to induce the
symptoms, it has been proposed that thermo-resistant allergens are involved in its
production. A novel variety of this syndrome occurs during physical exercise and is
therefore named dust mite ingestion-associated exercise-induced anaphylaxis
(Miller and Hamanaivary 2007; Sanchez-Machin et al. 2010; Sanchez- Borger et al.
2013; We et al. 2005).To prevent mite proliferation and the production of anaphy-
laxis, it has been recommended that wheat flour be stored at low temperatures in the
refrigerator.
One of the studies published in 1963 reported the case of a 56-year-old female
patient who died after a massive ingestion of mites contained in a pap made with
milk and wheat flour. The autopsy revealed intense intestinal irritation, disseminated
granulomas in multiple organs and the presence of many mites of species, Acarus
siro L., in stools and guts. OMA have also been reported from the USA, Japan,
Brazil, Taiwan, Spain and Venezuela. In general, there is a predominance of reports
from countries located in the intertropical area, probably because in these regions
environmental conditions especially higher temperature and relative humidity which
are more favourable for mite reproduction are found for longer periods of time.
25.2.1.2 Mite Species Involved in OMA

Different mite species are present in the flours producing OMA. Among them are
house dust mites (Dermatophagoides pteronyssinus (Trouessart) and D. farinae
Hughes), and storage mites such as Suidasia spp., Aleuroglyphus ovatus Troupeau,
Lepidoglyphus destructor (Schrank), Tyrophagus putrescentiae (Schrank),
Thyreophagus entomophagus Laboulbene, Blomia tropicalis and B. freemani are
commonly reported in flours producing OMA. Flour mites contaminate grain and
flour by producing allergens and transferring pathogenic microorganisms (Demerrel
et al. 2004; Spiegal et al. 1994; Thind and Clarke 2001). Mite-infested foodstuffs
acquire a sticky sweet smell and an unpalatable taste. Dust mites are not just limited
to mattresses, sofas and rugs but there are chances that they have been in your con-
sumables. Dust mites infest around 7 % of all wheat flour in Singaporean houses as
the climate of Singapore is a perfect breeding ground for these little critters. By
ingestion of contaminated flour, acute urticaria is prompted. But for those persons,
who are highly susceptible to house dust mites (HDMs), this could also trigger
severe asthma attacks as well as anaphylaxis. Severe infestation of flour by HDMs
results in a brownish tinge throughout the flour called ‘mist dust’. Mite dust gives
off misty odour and if crushed, the flour will become sticky and at this point will
acquire a honey-like smell.
25.2.1.3 Intestinal Acariasis

Acariasis is a condition in which live mites are deemed to be living in the lungs, guts
or the urinogenital system. Usually mites are found during pathological examina-
tion of samples of sputum, faeces or urine. With pulmonary acariasis, mites may
have been accidentally inhaled following high occupational exposure involving
stored products, whereas intestinal acariasis is likely due to ingestion of contami-
nated food. Intestinal acariasis may not cause injury of the intestinal mucosa and
invade the intestinal muscles but may stimulate the immune system by the dejecta,
metabolites and cleaned body of the mites itself.
25.2.1.4 Stored Animal Feed

Flour mite, Acarus siro, is a pest of stored grains and flour. The flour mite is pale
greyish white in colour with pink legs. Flour mite contaminates grain and flour
(stored as animal feed) and produces allergens and even transfers pathogenic micro-
organisms. Infested foodstuffs acquire a sickly sweet smell and an unpalatable taste.
When mite-infested feed is taken by animals, they show reduced food intake, diar-
rhoea, inflammation of the small intestine and impaired growth. The mites com-
monly infest mouldy animal feed such as baled hay and stored grain products (Dunn
et al. 2002; Heranz 1963; Hubert et al. 2003; Hubert et al. 2015; Hughes 1976). The
mites themselves are not considered a health hazard to animals feeding on infested
products; however, moulds on which the mites are feeding may produce toxins
which can poison animals that ingest the contaminated food.
25.2.2 Prophylactic Measures
Oral mite anaphylaxis (OMA) is a severe, potentially lethal allergic condition that
occurs in many countries and is frequently undiagnosed. Early recognition may lead
to the implementation of simple prophylactic measures in at-risk populations of
allergic patients. Observations revealed that most of the mite species in stored con-
ditions prefer moist areas and are commonly found in cracks and crevices near their
food source. The mites are barely visible to the naked eye, but may be present in
such huge numbers that the source of the infested materials appears to move. A tan
coating or pile of the ‘mite dust’ comprised of living and dead mites and their waste
products may appear on surfaces under or near infested foods. Prolonged contact
with mites can also be a problem in other storage areas such as barns or
outbuildings.
For the proper management of these stored mites, thorough cleaning of food
debris and spills even in pet food shelves and filling of cracks and crevices where
pests hide is a big must. Contaminated human or animal feed should preferably be
discarded so that further contamination of new lot may be checked. Always store
pet foods away from the general pantry area, and store them in tightly sealed
25.3 Dried Fruits 453
containers with tight lids. Improve air circulation among stored foods to reduce
humidity and prevent mouldy growth. Do not store at one place susceptible products
such as flour, baking mixes, cereals, animal feeds, etc. Try to buy animal pet food in
small packings and use them promptly.
The mites are sensitive to magnesium phosphide treatment. However, the limita-
tion of the fumigation is due to Carpoglyphus lactis (Linnaeus) mites’ ability to
form hypopus stadia with almost zero physiological and breathing activity which
enables them to survive fumigation. Manipulation of temperature and relative
humidity does not seem practical as hypopus developmental stage can survive under
unfavourable moisture and temperature conditions. However, some studies reported
that exposure to very low temperature (−15 °C) or low barometric pressure (95 mm
Hg) for 96 h completely kills the mites. However, small quantities of wheat flour
needed for human consumption may be invariably stored in refrigerators.
25.3 Dried Fruits
Dried fruit mite, Carpoglyphus lactis, has been found in most of the samples of
dried fruits like raisins, plums, figs, pineapple and banana collected from different
supermarkets. Dried apricots and plums are also reported highly susceptible to the
attack of different mites. In some species of dried fruits, there has been very fast
reproduction of mites at 25 °C and 85 % relative humidity (Chmielewski 1998;
Cobanoglu 2009; Hubert et al. 2011). Studies have revealed that these mites are able
to enter every dried fruit packaging material including polypropylene and alumin-
ium foils indicating that mites can move from package to package in supermarkets.
These mites are also known as allergen producers and are vectors of mycotoxin-
producing fungi. From Czech Republic, studies revealed about 13 % of 180 samples
of dried fruits collected from different supermarkets had infestation of C. lactis.
Mite contamination levels ranged from 0 to 660 per g of dried fruits. Mite contami-
nation is reported in dried apricots, figs, plums and raisins. Accidental consumption
of Tyrophagus longior (Gervais)-infested dried fruits resulted in diarrhoea due to
intestinal acariasis. Similar symptoms are also reported in persons who accidentally
consumed Acarus siro-infested dried seeds, grains, cereals and cheese. Such cases
have been mainly found in intertropical areas because in these regions environmen-
tal conditions are favourable, i.e. higher temperature and relative humidity occur-
ring for a longer period.
Simulation studies revealed that the hypothetical increase of mites was 20,000
mites on dried figs, followed by 50,000 contaminants on dried pineapple; 8000 on
dried apricots, plums and dates; and 3000 on raisins. In some studies mite infesta-
tions have been reported even in ready-to-use cereal-based foods including baby
food, biscuits and breakfast cereals. Commercially packaged popcorn, dried fruits
and peas are normally free of insects and mites at the time of purchase. Mite infesta-
tions usually take place after the package has been opened and the products are
stored for a relatively long period. So such foods should be stored in glass, metal or
other sealable containers. Mite infestations, if noticed, can be killed by placing the
material in a deep fridge or at lower temperatures in refrigerator for a minimum of

7–8 days. In case of beans and peas, mite infestations can be eliminated by heating
in the oven at 68 °C for about 30 min. But heat is not recommended for popcorn.
Even the seeds kept for planting should not be subjected to heating or cooling as
these treatments may reduce their germination rate. In Czech Republic, under labo-
ratory conditions, starting from a hypothetical level of ten mites of C. lactis per g of
dried fruits, the risk level of 1000 mites per g of dried fruits is reached in 42 days for
dried figs, dates and plums and 490 days for dried pineapple at 25 °C and 85 % rela-
tive humidity.
Mites are known as allergen producers and are vectors of mycotoxin-producing
fungi. The risks associated with C. lactis can disseminate the fungal stores. The
association of mites and mycotoxin-producing fungi such as Aspergillus spp. and
Penicillium spp. is noticed, and during their migration mites can disseminate the
fungal stores. The contamination of dried fruit is directly correlated with the risk of
direct consumption of hazardous mites (Arlian 2002; Aucamp 1969; Trucksess and
Scott 2008).
25.4 Mushrooms
Astigmatid mite, Histiostoma feroniarum (Dufour); Histiostomatidae is a cosmo-

politan species having a wide range of habitats including stored food, mushroom
houses and some agricultural plants like alfalfa, seeds and onions. Sometimes these
mites are also found on some insect cultures under laboratory conditions.
Contamination of stored food by H. feroniarum, which may seriously endanger
human health because of allergies, has been reported in samples from many super-
markets. The highest number of mites is also found on unwashed mushrooms
(Suillus luteus). This is not surprising because mushrooms are usually heavily pol-
luted with leaf litter and soils which are extremely rich in H. feroniarum and some
other oribatid mites. So more mites from these sources are transferred to mush-
rooms and breed there. However, as a storage mite, they can cause less significant
damage of grains and oilseeds. Contamination of stored food by arthropods includ-
ing this mite can cause respiratory syndrome like asthma and allergy (ER 3).
25.5 Soil Mites
Oribatid mites are found commonly in soil and leaf litter. These mites usually play
a beneficial role in soil formation but sometimes may cause damage to roots of cul-
tivated crops and may also contaminate foods by acting as harbourers of edible parts
like leaves, stems and fruits. Sometimes fodder crops like alfalfa, sorghum, maize
and timothy are particularly abundantly harbouring these mites, and in cases of
grazing by animals or through cut fodders, these mites are ingested directly by
them. However, these mites reproduce slowly and rarely may cause significant
25.6 Cheese 455
damage to the crops, but contaminate edible parts of crops, and foraging animals
can ingest them directly.
Oribatid mites Atropacarus striculus, Ceratozetes gracilis, Oribatula tibialis,
Scheloribates laevigatus and Suctobelbella falcata are commonly reported on low
growing crops. A high number of mites are found from unwashed leaves, vegetables
and fruits. Oribatid mites, S. laevigatus, found commonly as coprophage, are inter-
mediate hosts of the tapeworms, Moniezia expansa (family Anoplocephalidae),
which are parasitic on mammals including sheep, goat and cattle.
25.6 Cheese
Sometimes grain mite (Acarus siro), mould mite (Tyrophagus putrescentiae) and
cheese mite (Tyrolichus casei) infest stored foods and other organic debris such as
grain, flour, cereals, dried fruits, vegetables, pet foods, cheese, dried milk, dried
bananas, dried copra, ham, sugar, etc. These mites often prefer a moist and damp
location. Sometimes the infested materials appear to move due to the enormous
numbers of mites. Heavy infestations of grain mites have a sweet ‘minty’ odour,
best detected when mites are crushed between the thumb and forefinger. Coating or
piles of brownish ‘mist dust’ may appear on open shelving around the base of flour,
stacks on the surface of cheese or in other foods. Such piles consist of dead and liv-
ing mites and their cast off skins and faeces. Stomach disorders may result if mites
are taken internally with infested foods. However, acarid mites are responsible for
the sharp flavour of a famous German cheese: Altenburger ‘Milkenkase’. Any one
ingesting this cheese (with its thousands of mites) for the first time may have gas-
trointestinal disturbances. Mites can survive temperatures near freezing and may
become more prevalent during colder months. Symptoms like diarrhoea due to
‘intestinal acariasis’ produced by infestation with Tyrophagus longior have been
reported, which are frequently found on cheese and dried fruits.
‘Mimolette’ cheese is a hardball of ripened cows’ milk and is popular in the
northern city of Lille (France). In France it is also known as Boule de lille after its
city of origin. Cheese balls weigh about 2 kg (approximately 4.5 pounds). Its name
comes from the French word molle, meaning soft. This cheese has a similar appear-
ance to a cantaloupe melon. The greyish crust of aged mimolette is the result of
cheese mites intentionally introduced to add flavour by their action on the surface of
the cheese. Mimolette lovers love to eat this cheese at different stages of ageing.
Maturing the cheeses involves storing them in damp cellars and turning them every
week. At the same time, the surface of the cheese is brushed to remove cheese mites
which feast on its surface. As the cheese ages, evidence of mites can be seen in the
pitted and moonlike surface on the cheese. But the FDA is turning away imports of
mimolette cheese in the USA because the FDA seems concerned that cheeses with
too many mites could trigger allergic reactions and so has rejected shipments of
mimolette cheese with comments like filthy, putrid or decomposed substance that is
‘unfit for food’ (ER2).
25.7 Meat (Including Cured Fish)
Very tiny mites are reported occurring on the cured meat. In the surroundings the
mites may also be found infesting grain, cheese and flour. Heavily infested material
may have a sweet or ‘minty’ odour and the surface may appear to move if extremely
numerous mites are found. Over time, infested areas develop a powdery appearance
from the build up of dead mites and their shed skins. Since mite infestations are
limited to the surface, it may be possible to brush off most of the mites. A light coat
of vegetable oil can be rubbed thoroughly over the surface to kill the remaining
mites and eggs. Repeat the treatment in 7–10 days. Improve ventilation as much as
possible.
If the cured fish has a dusty rough-textured appearance, the dust in fact may be
mites. On cured fish, the common acarid mites found are Lardoglyphus spp. (family
Acaridae). Feeding by the adults and nymphs of Lardoglyphus spp. will cause quan-
titative loss of dried cured fish. Quality loss may result from the contamination by
live and dead bodies of mites, which can be very numerous. L. konoi has been found
infesting salted fish products as well as unsalted dried fish. Optimum conditions for
the development of L. konoi are 30–35 °C and 87 % relative humidity and below, but
population of mites increases at humidities of 75 % and above (up to 98 %).
Lardoglyphus spp. can become pests in many tropical and subtropical situations,
especially where ambient conditions are humid or the fish is not well dried. Home-
cured meats may also be infested by mites, and if a large number of mites are found,
dermatitis can develop from handling infested foods. Mites can be controlled with a
thorough cleanup at the infested area. In addition to L. konoi, L. angelina and L.
zacheri have also been reported on cured fish.
The development of Lardoglyphus spp. can be very rapid at 23 °C and 87 % rela-
tive humidity. The life cycles of L. konoi and L. zacheri take only 9–11 and 10–11
days, respectively. Both the species produce hypopus stage which do not feed but
are mainly a dispersal stage. Hypopi can climb onto the larvae and adults of beetle
pests and use their suckers to attach themselves to the insects’ cuticle and remain
there on the upper surface of the elytra for several days and are therefore often car-
ried to new infestation sites before they detach themselves and change into
tritonymph (Nunen 2015; Haines and Rees 1989).
25.8 Conclusions
Mites are well-known allergen producers. House dust mites are of a primary con-
cern owing to their proximity to humans. But recently many reports are appearing
regarding sensitizations of humans to many other stored product mites besides
Dermatophagoides spp. The consumption of pest mites along with infested food
may cause allergic reactions. The second risk is associated with the vectoring of
mycotoxin-producing fungi like Aspergillus spp. and Penicillium spp. The mites
may cause diseases inside the body through production of allergen, cutaneous der-
matitis and also in most parasitic diseases where they act as vector. Ingestion of
Further Reading 457
house dust mites may result in respiratory acariasis (taking place in the lungs), and
higher ingestion of stored product mites may result in intestinal acariasis and
patients may suffer from diarrhoea and may complain of abdominal pain. In human
acariasis, mites are visible in the stools. Sometimes mites are visible in the sputum
and the degree and extent of the disease is estimated by taking into account the total
number of mites seen in the sputum.
Various species of mites often infest stored foodstuffs, meat, baby food, dried
fruits, mushrooms, cheese, different drugs and pet foods and may cause economic
losses in terms of food quality and quantity. The losses are especially high under
humid and warm area. So such food items should be properly stored at appropriate
conditions depending on a quantity of material being stored. In case of dried fruits
like apricots, figs, plums and raisins, mites develop very fast at 25 °C and 85 % rela-
tive humidity. So such items should be stored in refrigerators near freezing.
Sometimes plant-feeding mites are found in large numbers on edible parts of
plants like leaves and fruits. So there are chances of ingestion of higher number of
mites along with the leaves and fruits when taken directly from the plants without
washing and cooking. In many underdeveloped societies, fruits of musk melon,
cantaloupe, sapota and papaya are taken without peeling and washing resulting in
ingestion of mites and the intestinal disorders. Most of the mites are not removed
even by washing the fruits in running water. So thorough washing of fruits and peel-
ing before consumption are strongly recommended. Even in case of vegetables like
long melon, cucumber and tomatoes, fruits should be consumed only after thorough
washing/peeling off.
Oral mite anaphylaxis (OMA) or ‘pancake syndrome’ is a severe, potentially
lethal allergic condition which occurs in many countries, but frequently remains
undiagnosed. Early detection of OMA may lead to implementation of simple pro-
phylactic measures in at-risk populations of allergic patients. OMA is a new syn-
drome characterized by severe allergic manifestations occurring in atopic patients
shortly after intake of foods made with contaminated wheat flour. Because cooked
foods are able to induce the symptoms, it has been proposed that thermo-resistant
allergens are involved in its production. A novel variety of this syndrome occurs
during physical exercise-induced anaphylaxis. To prevent mite proliferation and the
production of anaphylaxis, wheat flour should be stored at low temperatures, and
animal or pet feed should only be stored in properly sealed small packings. If
cheese, ham and salami are stored for long periods at room temperatures, then they
need to be examined at frequent intervals for mite infestations.
Further Reading
Arlian LG (2002) Arthropod allergens and human health. Annu Rev Entomol 47:395–433
Aucamp JL (1969) The role of mite vectors in the development of aflatoxin in groundnut. J Stored
Prod Res 5:245–249
Balance C, Quiraltge J, Castgillo R, Delgado J, Arteaga C, Berter D (1997) Anaphylaxis after
ingestion of wheat flour contaminated with the mites. J Allergy Clin Immunol 99:305–313
Chmielewski W (1998) Mites (Acarina) collected from stored apples. J Fruit Ornam Plant Res
6:33–40
Cobanoglu S (2009) Mite population density analysis of stored dried apricots in Turkey. Int
J Acarol 35:67–75
Demerrel DG, Olmos CE, El-Dohr JM (2004) Mites in the mix: dust mite contamination of a flour
product. J Allergy Clin Immunol 113(Suppl):32–35
Dunn JA, Thind BB, Banks J, Chambers J (2002) The development of ELISAs for the detection of
storage mites in grain. Food Agric Immunol 14:323–327
Haines CP, Rees DP (1989) A field guide to the types of insects and mites infesting cured-fish. FAO
corporate document repositary, produced by Fisheries and Agriculture Department (http://
www.fao.org/docrep/003/t0146e/T0146E02.htm)
Heranz G (1963) Hypersensitivity reaction to the ingestion of mites (Tyrophagus farina), patho-
logic study of a fatal case. Rev Med Univ Navarra 7:137–149
Hubert J, Stejskal V, Kubatova A, Munzburgeva Z, Vacova M, Oarkova E (2003) Mites as selective
fungal carriers in stored grain habitats. Exp Appl Acarol 29:69–87
Hubert J, Erban T, Nesvorna M, Steskal V (2011) Emerging risk of infestation and contamination
of dried fruits by mites in the Czech Republic. Food Addit Contam 28:1129–1135
Hubert J, Nesvorna M, Volek V (2015) Stored product mites (Acari: Astigmata) infesting food on
various types of packaging. Exp Appl Acarol 65:137–142
Hughes AM (1976) The mites of stored food and houses. Tech Bull Minist Agric Fish Food 9,
400 pp
Miller JD, Hamanaivary PJ (2007) The pancake syndrome. Allergy Asthma Proc 28:251–252
Nunen SV (2015) Tick-induced allergies: mammalian meat allergy, tick anaphylaxis and their
significance. Asia Pac Allergy 5:3–16
Sanchez-Borger M, Raul-Suarez C, Arnaldo CV, Fernan CF, Enrique FC (2013) Anaphylaxis from
ingestion of mites: Pancake anaphylaxis. J Allergy Clin Immunol 131:31–35
Sanchez-Machin I, Glez-Paloma PR, Iglesias-Souto J, Iraola V, Matghem V (2010) Oral mite ana-
phylaxis. Allergy 65:1345–1347
Skubala P, Marzec A, Skolowska M (2006) Accidental Acarophagy: mites found on foods, vegeta-
bles and mushrooms. Biol Lett 43:249–255
Spiegal WA, Anolok R, Jakabovice E, Arlian LG (1994) Anaphylaxis associated with dust mite
ingestion. Annu Allergy 72:56
Szilman P, Szilman D, Szilman M, Meszynska E, Maniurka H, Solar K, Sieron AL (2006)
Occupational exposure to allergenic mites among workers of the Silesian Zoo. Biol Lett
43:375–380
Thind BB, Clarke PG (2001) The occurrence of mites in cereal-based foods destined for human
consumption and possible consequences of infestation. Exp Appl Acarol 25:203–215
Trucksess MW, Scott P (2008) Mycotoxins in botanicals and dried fruits: a review. Food Addit
Contam 35:181–192
Walter DE, Proctor HC (1999) Mites, evolution and behaviour. CABI Publishing, New York,
322 pp
We DC, Shyur SD, Ho CM (2005) Systemic anaphylaxis after the ingestion of pancake contami-
nated with the storage mite, Blomia freemani. Ann Allergy Asthma Immunol 95:612–614
1. http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3651046/#__sec2title Pancake syndrome (Oral

mite anaphylaxis)
2. https://www.youtube.com/watch?v=MEGCH8W-EEU Cirons de la Mimolette Vieille, Cheese
Mites, acariens de fromage, Macro 100mm f2,4 1:1 × 2
3. http://en.agaricus.ru/cultivation/diseases/mites Mites
Further Reading 459
4. http://www.esciencecentral.org/journals/accidental-acarophagy--is-it-safe-for-us-2375-4397-
1000e110.pdf Accidental acarophagy – is it safe for us?
5. https://www.researchgate.net/publication/237457967_Accidental_acarophagy_mites_found_
on_fruits_vegetables_and_mushrooms Accidental acarophagy: mites found on fruits, vegeta-
bles and mushrooms
Glossary
26
Acari A very large varied subclass of class Arachnida; representatives are com-
monly called mites and ticks. In some old classifications, mites and ticks are
placed in order Acarina or as order Acari. They are usually small; their length
ranges from 0.08 to 16 mm in mites and 2–30 mm in ticks. In modern classifica-
tory accounts, in subclass Acari, two superorders Acariformes and Parasitiformes
are recognized. Their representatives are predaceous, parasitic, scavengers or
phytophagous.
Acaricide Alternatively known as miticide. A pesticide used to control mites and
ticks (e.g. chlorobenzilate, dicofol, avermectin, fenpyroximate), but is relatively
non-toxic to insects. The acaricides include a diverse array of chemical struc-
tures. A number of insecticides also display acaricidal activity.
Acaridomatium Dwelling spots for mites, often provided by certain plants which
in turn benefit from the presence of mites.
Acariformes A superorder of subclass Acari – representatives are without visible
stigmata, and their tactile and chemosensory hairs contain actinochitin – an
Abstract
About 115 terms pertaining to information given in 25 different chapters, included in the present
book ‘Fundamentals of Applied Acarology’ are given. Terms concerning knowledge on different
body regions, leg regions, and different structures particularly regarding various tactile and nerve
setae found on gnathosomal and idiosomal regions of body are presented in an authentic way.
Many terms pertaining to biology, ecology, and control techniques employed against these
creatures are included. Meaning of different plant parts on which mites are found feeding are also
included in this section. Terminology concerning Integrated Pest Management (IPM) are included
for better understanding the impact of mite pests on different field crops, vegetable crops and
different fruit plants. For obtaining additional information on these and more other terms concern-
ing applied acarology, readers are advised to see references listed under both Further reading, and
Electronic References (ER) at the end of this chapter.

DOI 10.1007/978-981-10-1594-6_26
462 26 Glossary
o ptically active material which exhibits birefringence in polarized light. The

Acariformes contain three suborders: Acaridida (Astigmata), Oribatida
(Cryptostigmata) and Actinedida (Prostigmata).
Acarine Disease Also known as Isle of Wight disease. A disease of adult bees
caused by the parasitic mite, Acarapis woodi, which enters from the tracheae and
feed. Mite infestation causes some debilities in bees; however, no outward signs
of mite infestation in bees may be found.
Acarocecidium A gall caused by gall mites (e.g. Eriophyidae).
Acarology A branch of zoology dealing with the scientific study of mites and ticks.
Acarophily A symbiotic relationship between plants and mites.
Acarophobia Abnormal fear of mites.
Actinopilin Optically active component of the core of birefringent setae in acari-
form mites. Actinopilin may occur in true setae (typical mechanoreceptors and
trichobothria) and the eupathidia and famulus. Solenidia do not have
actinopilin.
Agrochemicals The term agrochemicals includes plant nutrients like fertilizers as
well as other alternate chemicals like plant growth regulators, pheromones, hor-
mones, attractants, repellants, chemosterilants and bioorganisms.
Agroecosystem An agroecosystem is basically the ecosystem of an area as modi-
fied by the practice of agriculture, horticulture or animal rearing. It is composed
of cultivated land, the plants contained or grown thereon and the animals associ-
ated with these plants.
Ambulacrum The ambulatory appendage at the distal end of the tarsus, usually
consists of a pair of lateral claws and an empodium.
Apotele Terminal segment of appendages attached to the tarsus.
Berlese Funnel A sampling device that uses heat to drive away small insects and
mites from a sample of soil, vegetation or litter. The material is placed on a
coarse screen platform which has been inserted into a jar containing 70 % alco-
hol. Over the funnel is lighted a lamp. As the material dries, the insects and mites
which evade the light and heat are driven away from the material in 6–8 h and are
collected in the jar containing the alcohol.
Biodiversity The natural diversity of living organisms, the range of types and spe-
cies of plants and animals in an area. This also includes diversity within species,
between species and of ecosystem.
Biological Control The action of parasites, predators or pathogens on a host or
prey population which produces a lower generation equilibrium position than
would prevail in the absence of these agents. Biological control has been most
26 Glossary 463
successful against pests of perennial crops especially orchards than in annual

cropping systems.
Bursa Copulatrix Genital pouches in females of Astigmata with a copulatory
opening for accepting sperm (its opening is not genital opening).
Camerostome A recess under the rostora tectum that allows retraction of the
chelicerae and palps of oribatid mites.
Cannibalism Phenomenon of feeding on other individuals of the same kind.
Chela In mites distal pincer-like part of chelicerae. It is composed of two jaws: the
fixed jaw (distal part of the chelicerae body) and the movable jaw (the apotele).
Chemical Control The use of chemicals to kill, deter or in any way suppress pest
populations. The use of chemical control agents is useful as they act quickly and
result in rapid reductions in pest populations. But too much reliance on pesti-
cides results in problems like pollution of water and human dwellings, delays
occur in pest management attempts and development of resistance to pesticides
occurs frequently.
Chigger The parasitic larva of a trombiculid mite. Chiggers damage by feeding as
well as by serving as vectors of scrub typhus disease.
Claw A hooked structure on the distal end of the pretarsus, usually paired.
Crook Stage That stage of plant growth as it emerges from the soil.
Crop Rotation A technique of pest control which involves the deliberate planting
of specific crop sequences to make the year-to-year survival of soil-dwelling
pests difficult or impossible.
Crown The point where the stem and root join in a seed plant. This term is also
used to describe the foliage and branches of trees.
Cultural Control Manipulation of cultural practices to provide control of a pest.
Manipulation of cultural practices like sanitation, manuring, soil cultivation,
water and humidity management, strip farming, intercropping, crop rotation and
isolation, cultivation of trap crops and keeping variations in sowing date helps in
disrupting the life cycle of pests thus achieving their maximum control.
Degree Hours or Degree Days Degree days are also known as day degrees. An
accumulation of heat units above some threshold temperature for a 24-h period.
The units are used to estimate the total heat required to complete one generation
of a mite or insect.
Deutogyne A structurally different form of female of an eriophyoid mite species
that is found in the winter. Females of the same species in the summer (protogy-
nes) may have been described as different species, causing taxonomic
confusion.
464 26 Glossary
Deutonymph A developmental stage of mites; the deutonymph is the second

nymphal stage; often it is the last stage before moulting to the adult. Some
deutonymphs can be highly modified as hypopi, allowing the mites to survive
unfavourable conditions and perhaps to disperse to more favourable environ-
ments via phoresy.
Deutosternum A groove or gutter-like structure on the subcapitulum in parasiti-
form mites, often with transverse rows of anteriorly directed denticles.
Dockage Foreign material in harvested grain such as weed seeds, chaff and dusts;
value depreciation of a product owing to the presence of insects, mites or other
foreign material.
Domatia Plant chambers produced specially to house certain arthropods, espe-
cially mites and ants. Domatia may be hollow stems, swollen petioles or thorns,
which are used by mites and ants either for feeding or as nest sites or both.
Duplex Setae The pair of setae found on tarsus I and tarsus II, often with joint
bases. One of the pair is usually very long (solenidion) and the other a small
tactile seta.
Economic Injury Level (EIL) The lowest density of pest population that does
economically significant damage. The concept of EIL has marginal utility to
pests of public health importance wherein the most critical factor is the presence
or even the potential of a pathogen within the vector or host population.
Economic Threshold (ET) The pest density at which control measures should be
applied to prevent an increasing pest population from reaching the economic
injury level. It is sometimes also known as action threshold.
Exuviae The undigested exocuticle and epicuticle of nymphs and larvae shed dur-
ing ecdysis.
Festoons The ornamentation in some ticks that resembles a series of loops around
the margin of the body.
Flag Stage Stage of growth in cereals and other grasses at which the sheath and
leaf have been produced from which the head will emerge.
Forensic Acarology The use of acarological information in criminal investiga-
tions. Knowledge of the behaviour and development rates of carrion mites inhab-
iting corpses allows estimates of egg-laying time and, therefore, the time of death
of the victim. The police sometimes utilizes such information to investigate cases
of homicides.
Haller’s Organ A sensory field on the tarsus of legs I of ticks that contains
olfactory receptors along with heat and humidity receptors.
Hypopus Nonfeeding deutonymph in the Astigmata adapted for dispersal and
resisting adverse environmental conditions. They have a large ventral plate and
use insects for phoretic dispersal.
26 Glossary 465
Hypostome The anterior part of the subcapitalum in parasitiform mites; a barbed

piercing organ in ticks that contains retrorse teeth for attaching to the host.
Integrated Pest Management (IPM) A management system that uses all the suit-
able techniques and methods in as compatible a manner as possible to maintain
pest problems at levels below those causing economic injury. Cultural methods,
crop rotation, pest-resistant varieties, time of sowing or harvesting, trap crops
and chemical methods are the major components (tools) of IPM.
Key Pests The serious and the persistent pests which attack a crop and cause major
damage every season unless controlled is regarded as key pest.
Larva The second instar in acariform mites having a complete ontogenetic
sequence.
Life Cycle Habit and changes undergone by an organism from the egg stage to its
death; the sequence of events in the lifetime of an organism.
Life Table A description of the age-specific survival of cohorts of individuals in
relation to their age or stage of the development. Life table provides age-specific
data which are ideal for use in population management.
Lint Fibre surrounding the seed of unginned cotton.
Metastriata Hard ticks (Ixodidae) are divided into two groups, the Metastriata and
the Prostriata. The metastriatid ticks have an anal groove posterior to the anus.
Naso Unpaired protuberance on digital prodorsum in some acariform mites. Also
known as nasus.
Notogaster The dorsal hysterosoma or, in a more specific sense, the shield cover-
ing the dorsolateral aspects of hysterosoma in many oribatid mites.
Nymphochrysalis The protonymphal stage in the Parasitengona; a calyptostatic
cuticle formed within the larval cuticle in which the deutonymph stage
develops.
Occasional Pest A mite species which reaches significant levels only occasionally
and sporadically exceed the economic injury level.
Organic Farming A means of food production which discourages use of the
organic fertilizers and the plant protection chemicals. Pest control within organic
production systems is largely based upon the cultural techniques of crop rotation,
host plant resistance and various agronomic practices that are designed to
improve plant health.
Otoacariasis Invasion of the ear canal by mites or ticks.
Ovipositor An extrusible organ for laying the eggs. Female oribatid mites have a
well-developed ovipositor that ends in three fingerlike projections. Ovipositors
are less well developed or absent in other mites.
466 26 Glossary
Parasitiformes A superorder of the subclass Acari. Their representatives possess

one to four pairs of lateral stigmata posterior to the coxae of the second pair of
legs. Actinochitin is lacking in their tactile and chemosensory hairs. The
Parasitiformes have four suborders, of which the Gamasida (Mesostigmata) and
Ixodida (Metastigmata) are of great medical and veterinary importance. Location,
size and other structural aspects are used in taxonomic descriptions.
Peritreme Small plate perforated by spiracle opening in ticks, mites and some
insects. Peritreme is associated with a stigma. It is either a taenidium or a closed
(or partly closed) canal connected with the stigma or, a more or less, concave
plate surrounding the stigma.
Pesticide Treadmill The situation where the number of pesticide applications that
has to be made increases year after year has been called the pesticide treadmill.
In some of the cases, it has led to the complete collapse of agroecosystem.
Phoresy A type of migration where mites board larger animals (usually insects)
and cease or otherwise alter normal behaviours (e.g. feeding, reproduction,
movement, etc.) until some cue elicits the departure from the animal and the
resumption of normal behaviour. Phoresy usually results in the dispersal of
populations.
Phoretomorph Forms adapted for phoresy.
Phytosanitation The measures requiring the removal of or destruction of the
infected or infested plant material likely to form a source of reinfection or rein-
festation. This is done by disposal of unwanted crops and crop residues and the
elimination of weeds or volunteers that can harbour pests.
Phytotoxemia Also known as ‘toxicosis’. It is a group of diseases that are caused
by the toxins produced by insects or mites while feeding. An insect or mite
whose feeding produces symptoms of disease is said to be toxicogenic and the
condition is spoken of as ‘phytotoxemia’.
Prelarva The first instar in acariform mites having a complete ontogenetic
sequence, but absent in derived Prostigmata and Parasitiformes (except
Opilioacarida); incompletely formed, without functional mouthparts, often
retained within the egg shell or just extruding from it.
Prostriata Ixodid ticks are divided into Prostriata and Metastriata. Prostriata ticks
have an anal groove anterior to the anus and enclosing the anus.
Protogyne The normal feeding form, as opposed to the overwintering or aestivat-
ing form (deutogyne) in case of eriophyoid mites.
Protonymph The first stage in the life cycle in which the mite or tick has four pairs
of legs, coming after the larval stage in which only three pairs of legs are present
(except in the Eriophyoidea).
26 Glossary 467
Pteromorph Wing-like to shelf-like lateral extensions of the body in oribatid and

tarsonemid mites.
Ptychoidy The ability of some oribatid mites to withdraw the legs between two
body regions like a penknife being closed or a box being closed and resulting in
seedlike appearance. A derived form of dichoidy.
Pulvillus Cushion-like structure located ventral of and between the paired claw
and forming part of the ambulacrum.
Quarantine Regulations designed to prevent the spread of agricultural pests
through the commerce involving certain agricultural products and the restrictions
of the movement of infected plant material comprise what is referred as quaran-
tine; place where the people or animals are kept for the inspection.
Quiescence State of inactivity that is induced directly by adverse environmental
conditions in mites and ticks. Typically normal activity is restored immediately
upon a return to favourable environmental conditions.
Ratoon New tillers which grow from the stubbles of harvested plants. These new
tillers constitute the ratoon crop.
Resurgence The term resurgence is used to express a sudden increase in popula-
tion number of insects or mites. One type occurs when the target species which
was initially suppressed by the insecticidal treatment undergoes rapid recovery
after the decline of the treatment effect. Resurgence can be managed by avoiding
hormoligosis, by avoiding natural enemy destruction and by intelligent use of
pesticides considering their physiological and ecological selectivity.
Retrorse Teeth Turned or bent backward; this type of teeth is found on the
hypostome of ticks and makes it difficult to remove the tick from the host during
feeding.
Rhagidial Organs Recumbent solenidia in shallow depressions on the distal seg-
ments of leg I–II in Eupodoidea (Prostigmata).
Rosette A group of the leaves which are arising from a short stem and are, there-
fore, lying close together on or near the ground.
Rotation Practice of growing different crops on the same land in a regular and
recurring sequence. Rotations tend to be most effective against pest species that
have a narrow host range and limited rate of dispersal.
Rutellum In Sarcoptiformes, the hypertrophied setae on the hypostome which are
often toothed.
Scabies A contagious skin disease of animals caused by parasitic mite, Sarcoptes
scabiei.
Secondary Pest A pest that usually does little damage but it can become a serious
pest under certain conditions.
468 26 Glossary
Sensillum Also known as sensilla or sensillae. A sensory structure, sensilla is often

used for the bothridial seta in Acariformes.
Silk Substance produced by some species of spider mites, although not all; silk is
important in their biology providing benefits such as protecting them from rain,
predation and desiccation.
Solenidion A hollow optically inactive chemosensory setae in limbs of acariform
mites.
Spermatodactyl A structure on the chelicerae of some gamasid males that is
involved in sperm transfer (podospermy). The structure of the spermatodactyl
can be used to identify species. Typically, podospermy involves the male taking
up a bundle of sperm from the genital opening on his venter and inserting the
sperm packet into a structure in the female located between legs three and four.
Spinneret A structure that produces silken fibres. In spider mites, the spinneret is
in the form of a short seta-like structure on the palp tarsus.
Staggered Planting Planting different fields in a commodity or a farm over a
period of several weeks in contrast to simultaneous planting where planting of all
fields is done over a period of a week or less.
Stigma The opening to the tracheal system.
Stippling Localized damage to foliage characterized by numerous pale dots or
points where the chlorophyll has been removed by mites during feeding.
Stylet A narrow, pointed structure, typically referring to a mouthpart in the Acari.
Stylophore Chelicerae formed of fused cheliceral bases bearing stylet-like move-
able digits as in Raphignathidae (Prostigmata), which includes the plant parasitic
spider mites and their relatives (Tetranychoidea).
Stylostome A ramifying structure produced in the skin of a host by the feeding of
a chigger or larval water mite.
Subcapitulum Also known as infracapitulum. The venter of the capitulum; the
ventral faces of the fused palp coxae apparently formed independently in the two
superorders of mites.
Surveillance Also called scouting. Refers to the constant watch on a pest for the
detection of the species presence and determination of population density, dis-
persion and dynamics on search crop at fixed interval.
Tanglefoot A sticky substance used in the traps to immobilize mites. Tanglefoot is
used as a barrier on leaf discs for studying development of mites and for pesti-
cidal efficacy against mites.
Tectum Any shelflike projection of the cuticle. In Mesostigmata the leading dorsal
edge of the basic capitulum is referred to as tectum, gnathotectum or epistome.
26 Glossary 469
Tectum capituli The tectum or epistome of Mesostigmata.

Tenent Hairs Tiny hairs on the pulvilli and tarsal pads that allow some mites to
cling to smooth surfaces.
Tegula Lobe-like to acuminate tectum projecting posteriorly between coxae IV in
Tarsonemidae.
Tetrapodili An obsolete name for the Eriophyoidea.
Thumb–Claw Complex A compound structure of palp in some acarines which is
adapted to holding. It consists of a tibial seta (the claw) and the palpal tarsus (the
thumb).
Tillering Stage The development of side shoots from the base of a single-stemmed
cereal or grass plant.
Tolerance The condition when the host plant may show an ability to grow and
reproduce itself or to repair injury despite the presence of mite or insect popula-
tion adequate to damage a normally susceptible host beyond repair.
Transgenic A transgenic plant is simply a normal plant with one or more addi-
tional genes from diverse sources. Application of transgenic plants through
genetic engineering is the latest concept in management of insect and mite pests.
Trichobothria Minute sensory hairs on the tarsi in mites and spiders, often set in a
cuplike base.
Tritonymph The third and final nymphal stage or instar present in Opilioacarida,
Holothyrida, Argasidae and many Acariformes.
Tritosternum The sternum of the third body segment (between legs I) produced as
a biflagellate structure in Mesostigmata, although sometimes the flagella (lacinia)
are partial or completely fused.
Vagrant A mite which occasionally moves from one place to another but not with
any regularity, the implication is that the movement is accidental.
Varroasis Also sometimes called as ‘varroatosis’. Parasitic mite, Varroa destruc-
tor, is the major pest of honeybees; the mite reproduces mainly on the drone
pupae. Varroasis symptoms include the presence of pale or dark-brown mites on
the white pupae, and the drone or worker brood has punctured cappings and
disfigured and stunted adults.
Vector A living organism (e.g. insect, mite, nematode, etc.) able to carry and trans-
mit a pathogen (virus, bacterium, fungus), thus disseminating disease.
Web Network of threads spun by the spider mites. Through ballooning spider
mites are transported from one habitat to another.
470 26 Glossary
Weed A plant growing in the wrong place where it is not desired; an aggressive,
invasive, easily dispersed plant, one which commonly grows in cultivated ground
to the detriment of a crop.
Witches’ Broom Disease symptom with an abnormal, massed, brush-like develop-
ment of many weak shoots or roots of mainly woody plants, arising at or close to
the same point or resulting from the proliferation of buds, caused by mites,
viruses, fungi, bacteria, nematodes, etc.
Wolbachia A genus of bacteria that is a very common microbial endosymbiont of
arthropods. Some are associated with modifications of the sex ratio, with cyto-
plasmic incompatibility and with fitness costs. Usually Wolbachia is found in the
reproductive tracts (ovaries) and transovarially transmitted to progeny. However,
the effect of Wolbachia often remains unknown.
Further Reading
Van der Hammen L (1976) Glossary of acarological terminology. Boston W. Junk, The Hague
1. http://itp.lucidcentral.org/id/mites/invasive_mite/Invasive_Mite_Identification/key/0_Glossary/
Mite_Glossary.htm Glossary of acarine terms
2. http://www.ipm.ucdavis.edu/PMG/glossary.html IPM glossary
3. http://bristoltickid.blogs.ilrt.org/introduction-3/glossary-of-morphological-features/ Glossary
of morphological features of ticks

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Manjit

ISBN 978-981-10-1592-2 ISBN 978-981-10-1594-6 (eBook)

Library of Congress Control Number: 2016953350

© Springer Science+Business Media Singapore 2016

Printed on acid-free paper

This Springer imprint is published by Springer Nature

thoroughly discussed. Safe usage of pesticides based on their mode of action

Ludhiana, India M.S. Dhooria

562-I Block, B.R.S. Nagar, M.S. Dhooria

1 Acarology and Its Importance................................................................ 1

3.7 Packing and Shipping Mites......................................................... 33

5.2 Superorder Acariformes................................................................ 67

8.4 Dispersal of Populations and Species........................................... 180

11 Quarantine Acarology............................................................................. 207

13 Mite Pests of Field Crops......................................................................... 275

1 4 Mite Pests of Greenhouse Crops............................................................. 307

16.3 Plant Viral Diseases Vectored by Spider Mites............................. 336

19.1.9 Order Coleoptera.......................................................... 371

23 Medical and Veterinary Acarology......................................................... 425

25.6 Cheese........................................................................................... 455

Acarology is a sub-discipline of Zoology and is sometimes included in the field of

© Springer Science+Business Media Singapore 2016 1

cycles, a myriad of their dispersal techniques, and adaptability to diverse ecological

Predatory Mites Diverse ecological relationships among mites may be utilized in

Acariasis ‘Acariasis’ in animals is caused by a variety of mites. The zoonotic spe-

Electronic References (ER)

1. https://www.youtube.com/watch?v=_BPE9-JymTA Arachnids: the different kinds

© Springer Science+Business Media Singapore 2016 9

2.1 Developments Prior to 1850 AD

2.2 Progress During 1851–1950 AD

systems: (i) Acari, Atracheata (including Glycyphagus, Histiostoma, Phytoptus,

2.3 Developments Since 1951

The eminent scientists, EW Baker and GW Wharton, investigated thoroughly diver-

2.4 OSU Acarology Development Programme

At Duke University (USA), Dr George W. Wharton established a section ‘The

2.5 Acarological Societies

1. International Congress of Acarology (ICA)

Table 2.1 International Congress of Acarology

Table 2.1 (continued)

Table 2.2 EURAAC symposiums

9. The Acarological Society of Iran (ASI)

2.6 Journals in Acarology

1. International Journal of Acarology (IJA). [ISSN: 0164-7954 (print); and ISSN:

2.7 Important Books Published

Baker EW, Wharton GW. 1952. An introduction to Acarology, MacMillan,

Johnston DE (1968) An Atlas of Acari 1. The families of Parasitiformes and Opilioacariformes.

Electronic Reference (ER)

1. https://acarology.osu.edu/history. Acarology History

© Springer Science+Business Media Singapore 2016 21

3.1.1 Plant-Associated Mites

Vacuum Technique A hand-operated vacuum apparatus can be used for collecting

3.1.2 Water Mites

3.1.3 Soil Mites

Simple Floatation Technique The collected soil sample is stirred in a magnesium

3.1.4 Parasitic Mites

3.1.4.1 Bee-Associated Mites (Barlow 2009; Bush 2009)

Ludhiana, India M.S. Dhooria

1 Acarology and Its Importance................................................................ 1

3.7 Packing and Shipping Mites......................................................... 33

5.2 Superorder Acariformes................................................................ 67

8.4 Dispersal of Populations and Species........................................... 180

11 Quarantine Acarology............................................................................. 207

13 Mite Pests of Field Crops......................................................................... 275

1 4 Mite Pests of Greenhouse Crops............................................................. 307

16.3 Plant Viral Diseases Vectored by Spider Mites............................. 336

19.1.9 Order Coleoptera.......................................................... 371

23 Medical and Veterinary Acarology......................................................... 425

25.6 Cheese........................................................................................... 455

2.1 Developments Prior to 1850 AD

2.2 Progress During 1851–1950 AD

2.3 Developments Since 1951

2.4 OSU Acarology Development Programme

2.5 Acarological Societies

2.6 Journals in Acarology

2.7 Important Books Published

3.1.1 Plant-Associated Mites

3.1.2 Water Mites

3.1.3 Soil Mites

3.1.4 Parasitic Mites

3.1.4.1 Bee-Associated Mites (Barlow 2009; Bush 2009)

3.1.4.2 Mites Associated with Other Insects

3.1.4.3 Mites from Other Invertebrates

3.1.4.4 Mites from Vertebrates

3.1.4.5 Collection from Stored Products

3.1.5 Collection of Ticks

3.1.5.1 Direct Examination

3.1.5.2 Combing Technique

3.1.5.3 Dipping Technique

3.1.5.4 Dragging Technique

3.1.5.5 Carbon Dioxide (CO2) Trapping

3.1.5.6 Host Trapping

3.2 Preservation/Killing in a Medium

3.3 Preparation (Slide Mounting)

3.3.1 Preparation for Water Mites

3.3.2 Plant-Associated Mites

3.3.2.1 Temporary Preparation

3.3.2.2 Permanent Preparation

3.3.3 Preparation of Ticks for Study

3.3.4 Mounting Ticks on Glass Slides

3.3.6 Slide Preparation Technique

3.4 reparation for Scanning Electron Microscopy Studies

3.5 Preparation for Molecular Analysis Studies

3.6 Photomicrography of Mites

3.7 Packing and Shipping Mites

3.7.1 Specimens in Vials

3.7.2 Shipping Microscope Slides

3.7.3 Shipping Specimens

3.8 Working with Live Mites

3.9 Rearing Techniques

3.10 Sampling Methods

3.10.1 Direct Counting

3.10.2 Imprint Counting

3.10.3 Mite-Brushing Machine

3.10.4 Beating Foliage

3.10.5 Washing Technique

3.11 Acaricide Testing Techniques

3.11.1 Slide-Dip Method

3.11.2 Leaf-Dip Method

3.11.3 Residual Leaf Disc Assays

3.11.4 Ovolarvicide Tests

4.1 External Morphology

4.1.2 Differences Between Hard Ticks and Soft Ticks

4.1.3 Body Divisions

4.1.4 Sensory Receptors

4.2 Internal Anatomy (Fig. 4.9)

4.2.1 Digestive System

4.2.2 Central Nervous System

4.2.3 Circulatory System

4.2.5 Respiratory System