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Medical and Veterinary Entomology

Gary R. Mullen
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Medical and Veterinary Entomology
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Medical and Veterinary
Entomology
Third Edition

Edited by

Gary R. Mullen
Department of Entomology and Plant Pathology, Auburn University, Auburn, AL, United States

Lance A. Durden
Department of Biology, Georgia Southern University, Statesboro, GA, United States

Jonas G. King, Graphics Assistant


Department of Biochemistry, Molecular Biology, Entomology and Plant Pathology,
Mississippi State University, Starkville, MS, United States
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This book and the individual contributions contained in it are protected under copyright by the Publisher (other than as may be
noted herein).

Notices
Knowledge and best practice in this field are constantly changing. As new research and experience broaden our understanding,
changes in research methods, professional practices, or medical treatment may become necessary.

Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using any information,
methods, compounds, or experiments described herein. In using such information or methods they should be mindful of their
own safety and the safety of others, including parties for whom they have a professional responsibility.
To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume any liability for any injury
and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of
any methods, products, instructions, or ideas contained in the material herein.
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A catalogue record for this book is available from the British Library

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Contents

Contributors xv 3. Arthropod Toxins and Venoms


Preface xvii
Acknowledgments xix Justin O. Schmidt
Nature of Toxins and Venoms and Their Effects 24
Arthropod Venoms as Antimicrobial, Analgesic,
1. Introduction and Other Therapeutic Agents 28
References and Further Reading 29
Lance A. Durden and Gary R. Mullen
General Entomology 1 4. Epidemiology of Vector-Borne
Medical-Veterinary Entomology Literature 2 Diseases
A Brief History of Medical-Veterinary
Entomology 2 Christopher M. Barker and William K. Reisen
Identification and Systematics of Arthropods of Components of Transmission Cycles 34
Medical-Veterinary Importance 4 Host Immunity 35
Types of Problems Caused by The Vertebrate Host 35
Arthropods 4 The Arthropod Vector 36
Annoyance 4 Modes of Transmission 37
Toxins and Venoms 5 Vertical Transmission 37
Allergic Reactions 5 Horizontal Transmission 38
Invasion of Host Tissues 6 Transmission Cycles 40
Arthropod-Borne Diseases 6 Interseasonal Maintenance 41
Food Contaminants 6 Vector Incrimination 42
Fear of Arthropods 7 Surveillance 43
Delusional Disorders 7 Emerging Vector-Borne Diseases 47
Formicophilia 8 New Tools 49
Host Defenses 8 References and Further Reading 49
Minor Arthropod Problems of Medical-
Veterinary Interest 9 5. Forensic Entomology
References and Further Reading 10
William L. Krinsky
2. Morphological Adaptations of History 51
Parasitic Arthropods Legal Cases Involving Liability 52
Nathan D. Burkett-Cadena Structural Entomology 52
Stored-Product Entomology 52
Body Shape and Wings 17 Occupational Hazards Associated With
Mouthparts 17 Arthropods 52
Legs 19 Veterinary and Wildlife Entomology 52
Sensory Structures 21 Legal Cases Involving Homicides, Suspicious
References and Further Reading 22 and Accidental Deaths, and Abuse 53

v
vi Contents

Sudden Death With Arthropod Association 53 Human Body Louse (Pediculus humanus
Automobile-Accident Death 53 humanus) 87
Arthropods as Signs of Neglect or Abuse or as Human Head Louse (Pediculus humanus
Agents of Murder 53 capitis) 88
Illicit Drug Transport, Use, and Overdose 53 Human Crab Louse (Pthirus pubis) 88
Suspicious Deaths 54 Lice of Veterinary Importance 89
Stages of Decomposition 54 Lice of Cattle 89
Insect Succession and Postmortem Interval 55 Lice of Other Livestock Animals 90
References and Further Reading 59 Lice of Cats and Dogs 92
Lice of Laboratory Animals 92
6. Cockroaches (Blattaria) Lice of Poultry and Other Birds 93
Public Health Importance 94
Richard J. Brenner and Richard D. Kramer
Epidemic Typhus 95
Taxonomy 61 Louse-Borne Relapsing Fever 96
Morphology 61 Trench Fever 97
Life History 63 Other Pathogens Transmitted by Human Body
Behavior and Ecology 64 Lice 98
Common Cockroach Species 65 Lice as Intermediate Hosts of Tapeworms 98
Oriental Cockroach (Blatta orientalis) 65 Veterinary Importance 98
Turkestan Cockroach (Blatta lateralis) 65 Lice of Livestock 98
American Cockroach (Periplaneta Lice of Wildlife 101
americana) 65 Lice of Cats and Dogs 101
Australian Cockroach (Periplaneta australasiae) 66 Lice of Laboratory Animals 102
Brown Cockroach (Periplaneta brunnea) 66 Lice of Poultry and Other Birds 102
Smokybrown Cockroach (Periplaneta Prevention and Control 102
fuliginosa) 67 References and Further Reading 104
Florida Woods Cockroach (Eurycotis floridana) 67
Brown-Banded Cockroach (Supella longipalpa) 68 8. True Bugs (Hemiptera)
German Cockroach (Blattella germanica) 68
William L. Krinsky
Asian Cockroach (Blattella asahinai) 69
Surinam Cockroach (Pycnoscelus Kissing Bugs (Reduviidae) 109
surinamensis) 69 Taxonomy 109
Public Health Importance 70 Morphology 109
Pathogenic Agents 70 Life History 111
Intermediate Hosts 72 Behavior and Ecology 111
Cockroach Allergies 73 Public Health Importance 112
Veterinary Importance 74 Veterinary Importance 118
Prevention and Control 75 Prevention and Control 118
Sanitation 75 Bed Bugs (Cimicidae) 119
Harborage Elimination 75 Taxonomy 119
Physical Control 75 Morphology 119
Biological Control 75 Life History 120
Insect Growth Regulators 76 Behavior and Ecology 121
References and Further Reading 76 Public Health Importance 122
Veterinary Importance 123
7. Lice (Phthiraptera) Prevention and Control 124
References and Further Reading 124
Lance A. Durden
Taxonomy 79 9. Beetles (Coleoptera)
Morphology 80
William L. Krinsky
Life History 83
Behavior and Ecology 84 Taxonomy 129
Lice of Medical Importance 86 Morphology 129
Contents vii

Life History 130 Murine Trypanosomiasis 165


Behavior and Ecology 130 Other Flea-Borne Pathogens and Parasites 165
Public Health Importance 131 Fleas as Intermediate Hosts or Vectors of
Meloidae (Blister Beetles) 133 Helminths 165
Oedemeridae (False Blister Beetles) 134 Prevention and Control 166
Staphylinidae (Rove Beetles) 134 References and Further Reading 167
Tenebrionidae (Darkling Beetles) 135
Dermestidae (Larder Beetles) 137 11. Flies (Diptera)
Scarabaeidae (Scarab Beetles) 137
Reid R. Gerhardt and Lawrence J. Hribar
Coccinellidae (Lady Beetles) 137
Veterinary Importance 138 Taxonomy 171
Ingestion of Toxic Beetles 138 Morphology 173
Transmission of Pathogens 138 Life History 177
Intermediate Hosts of Parasites 139 Behavior and Ecology 177
Nest Associates and Ectoparasites 140 Families of Minor Medical or Veterinary
Dung Beetles and Biocontrol 140 Interest 178
Prevention and Control 141 Tipulidae (Crane Flies) 178
References and Further Reading 141 Bibionidae (March Flies) 179
Sciaridae (Dark-Winged Fungus Gnats) 179
10. Fleas (Siphonaptera) Chaoboridae (Phantom Midges) 180
Corethrellidae (Frog-Biting Midges) 180
Lance A. Durden and Nancy C. Hinkle
Chironomidae (Chironomid Midges) 180
Taxonomy 145 Rhagionidae (Snipe Flies) 182
Morphology 146 Athericidae (Athericid Flies) 182
Life History 148 Stratiomyidae (Soldier Flies, Latrine Flies) 182
Behavior and Ecology 149 Phoridae (Humpbacked Flies,
Fleas of MedicaleVeterinary Importance 150 Scuttle Flies) 183
Human Flea (Pulex irritans) 150 Syrphidae (Flower Flies, Hover Flies) 184
Cat Flea (Ctenocephalides felis) 151 Piophiliae (Skipper Flies) 184
Dog Flea (Ctenocephalides canis) 151 Drosophilidae (Small Fruit Flies) 185
Oriental Rat Flea (Xenopsylla cheopis) 151 Chloropidae (Grass Flies, Eye Gnats) 185
European Rabbit Flea (Spilopsyllus cuniculi) 152 Public Health Importance 186
Sticktight Flea (Echidnophaga gallinacea) 152 Veterinary Importance 186
Chigoe (Tunga penetrans) 152 Prevention and Control 187
Northern Rat Flea (Nosopsyllus fasciatus) 153 References and Further Reading 188
European Chicken Flea (Hen Flea in Europe)
(Ceratophyllus gallinae) 153 12. Phlebotomine Sand Flies and Moth
European Mouse Flea (Leptopsylla segnis) 154 Flies (Psychodidae)
Public Health Importance 154
Flea-Associated Allergies 156 Leonard E. Munstermann
Plague 156 Taxonomy 192
Murine Typhus 159 Psychodinae 192
Other Flea-Borne Rickettsial Agents 161 Sycoracinae 192
Other Flea-Borne Pathogens 161 Phlebotominae 193
Bacteria 161 Morphology 193
Viruses 162 Psychodinae 193
Tungiasis 162 Phlebotominae 194
Fleas as Intermediate Hosts of Helminths 162 Life History 195
Veterinary Importance 163 Psychodinae 195
Flea Allergy Dermatitis 164 Phlebotominae 195
Tungiasis 164 Behavior and Ecology 196
Myxomatosis 164 Psychodinae 196
Squirrel Pox 165 Phlebotominae 196
viii Contents

Public Health Importance 198 Veterinary Importance 251


Psychodinae 198 Bovine Onchocerciasis 252
Phlebotominae 198 Leucocytozoonosis 252
Vesicular Stomatitis Virus Disease 198 Other Parasites and Pathogens of Veterinary
Chandipura Virus Disease 200 Importance 253
Sand Fly Fever 200 Simuliotoxicosis 253
Changuinola Virus Disease 201 Prevention and Control 254
Bartonellosis 201 Onchocerciasis Control 255
Leishmaniasis 202 References and Further Reading 256
Veterinary Importance 207
Leishmaniasis 208 15. Mosquitoes (Culicidae)
Vesicular Stomatitis Virus Disease 208
Woodbridge A. Foster and Edward D. Walker
Prevention and Control 208
Psychodinae 208 Taxonomy 261
Phlebotominae 209 Morphology 264
References and Further Reading 210 Life History 269
Behavior and Ecology 272
13. Biting Midges (Ceratopogonidae) Genetics 279
Public Health Importance 280
Gary R. Mullen and C. Steven Murphree
Mosquito Bites 280
Taxonomy 213 Mosquito-Borne Viruses 280
Morphology 214 Togaviridae (Alphavirus) 283
Life History 215 Flaviviridae (Flavivirus) 287
Behavior and Ecology 216 Bunyaviridae (Orthobunyavirus and
Public Health Importance 218 Phlebovirus) 297
Oropouche Fever 221 Malaria 300
Other Viral Agents 221 Filariasis 305
Mansonellosis 222 Veterinary Importance 310
Veterinary Importance 223 Mosquito-Borne Viruses of Animals 310
Bluetongue Disease 223 Nonhuman Malarias 312
Epizootic Hemorrhagic Disease 225 Dog Heartworm 313
African Horse Sickness 227 Other Filarial Nematodes of Animals 314
Schmallenberg Virus 228 Prevention and Control 314
Other Viral Agents 229 Control of Pathogen Transmission 317
Blood Protozoans 229 References and Further Reading 317
Equine Onchocerciasis 230
Other Filarial Nematodes 231 16. Horse Flies and Deer Flies
Equine Allergic Dermatitis 231 (Tabanidae)
Prevention and Control 232
References and Further Reading 232 Bradley A. Mullens
Taxonomy 327
14. Black Flies (Simuliidae) Morphology 329
Life History 330
Peter H. Adler and John W. McCreadie
Behavior and Ecology 332
Taxonomy 237 Public Health Importance 336
Morphology 238 Loiasis 336
Life History 240 Tularemia 337
Behavior and Ecology 240 Other Tabanid-Transmitted Human
Public Health Importance 243 Pathogens 338
Biting and Nuisance Problems 243 Veterinary Importance 338
Human Onchocerciasis 245 Surra and Related Trypanosomiases 338
Mansonellosis 250 Equine Infectious Anemia 339
Other Diseases Related to Black Flies 250 Anaplasmosis 339
Contents ix

Elaeophorosis 340 18. Tsetse Flies (Glossinidae)


Other Pathogens of Veterinary Importance 340
Prevention and Control 340 William L. Krinsky
References and Further Reading 341 Taxonomy 369
Morphology 370
17. Muscid Flies (Muscidae) Life History 371
Behavior and Ecology 372
Roger D. Moon
Public Health Importance 374
Taxonomy 345 African Sleeping Sickness 374
Morphology 346 Veterinary Importance 377
Life History 350 Nagana 377
Behavior and Ecology 351 Prevention and Control 379
Species of Medical-Veterinary Importance 353 References and Further Reading 381
House Fly (Musca domestica) 353
Bazaar Fly (Musca sorbens) 354 19. Myiasis (Muscoidea, Oestroidea)
Bush Fly (Musca vetustissima) 354
Face Fly (Musca autumnalis) 354 Philip J. Scholl, Douglas D. Colwell, and
Cluster Fly (Pollenia rudis) 354 Ramón Cepeda-Palacios
Stable Fly (Stomoxys calcitrans) 354 Taxonomy 384
Horn Fly (Haematobia irritans irritans) and Morphology 389
Buffalo Fly (Haematobia irritans exigua) 355 Life History 390
False Stable Fly (Muscina stabulans) and Its Ecology and Behavior 392
Relatives 355 Myths 394
Little House Fly (Fannia canicularis) and Its Flies Involved in Myiasis 394
Relatives 355 Psychodidae (Moth Flies, Drain Flies) 394
Garbage Flies (Hydrotaea spp.) 356 Stratiomyidae (Soldier Flies) 394
Sweat Flies (Hydrotaea spp.) 356 Phoridae (Humpback Flies) 395
Bird Nest Parasites (Passeromyia spp., Phil- Syrphidae (Flower Flies, Hover Flies,
ornis spp., and Protocalliphora spp.) 356 Rat-Tailed Maggots) 395
Public Health Importance 356 Piophilidae (Skipper Flies) 395
House Fly (Musca domestica) 357 Neottiophilidae (Nest Skipper Flies) 396
Bazaar Fly (Musca sorbens) 357 Drosophilidae (Pomace Flies, Vinegar Flies,
Bush Fly (Musca vetustissima) 358 Fruit Flies, and Wine Flies) 396
Face Fly (Musca autumnalis) and Cluster Fly Chloropidae (Grass Flies and Australian
(Pollenia rudis) 358 Frog Flies) 396
Stable Fly (Stomoxys calcitrans) 358 Anthomyiidae (Root Maggots) 396
False Stable Fly (Muscina stabulans) and Its Fanniidae (Faniid Flies) 396
Relatives 359 Muscidae (Dung Flies) 397
Little House Fly (Fannia canicularis) and Its Tropical Nest Flies 397
Relatives 359 Calliphoridae (Blow Flies, Carrion Flies,
Garbage Flies (Hydrotaea spp.) 359 Floor Maggots, Nest Maggots, and
Sweat Flies (Hydrotaea spp.) 359 Screwworms) 398
Veterinary Importance 359 Carrion-Associated Blow Flies 398
House Fly (Musca domestica) 360 Sarcophagidae (Flesh Flies) 401
Bush Fly (Musca vetustissima) 360 Oestridae (Bot Flies) 402
Face Fly (Musca autumnalis) 361 New World Skin Bot Flies (Cuterebrinae) 403
Stable Fly (Stomoxys calcitrans) 362 Old World Skin Bot Flies (Hypodermatinae) 406
Horn Fly (Haematobia irritans irritans) and Nose Bot Flies (Oestrinae) 407
Buffalo Fly (Haematobia irritans exigua) 363 Stomach Bot Flies (Gasterophilinae) 410
Sweat Flies (Hydrotaea spp.) 364 Other Oestroid Flies 411
Bird Nest Parasites (Philornis spp.) 364 Public Health Importance 411
Prevention and Control 364 Clinical Use of Maggots 412
References and Further Reading 366 Veterinary Importance 413
x Contents

Prevention and Control 414 Veterinary Importance 454


Screwworm Eradication Program 415 Caterpillar-Induced Equine Abortion 454
Cattle Grub Control 416 Prevention and Control 455
Myiasis and Molecular Advances 417 References and Further Reading 455
References and Further Reading 417
22. Ants, Wasps, and Bees
20. Louse Flies, Keds, and Bat Flies (Hymenoptera)
(Hippoboscoidea) Hal C. Reed and Peter J. Landolt
Will K. Reeves and John E. Lloyd
Taxonomy 459
Taxonomy 421 Morphology 460
Morphology 422 Life History 463
Hippoboscidae 423 Behavior and Ecology 464
Streblidae 424 Hymenoptera Venoms 465
Nycteribiidae 424 Ant Venoms 465
Life History 424 Vespid Venoms 466
Behavior and Ecology 425 Honey Bee Venom 466
Common Species of Hippoboscids 426 Ants 466
Sheep Ked (Melophagus ovinus) 426 Fire Ants (Solenopsis spp.) 467
Dog fly (Hippobosca longipennis) 427 Harvester Ants (Pogonomyrmex spp.) 469
Hippobosca equina 428 Pavement Ant (Tetramorium caespitum) 470
Hippobosca variegata 428 Pharaoh’s Ant (Monomorium pharaonis) 470
Deer Keds (Lipoptena and Neolipoptena Wasps 471
spp.) 428 Solitary Wasps 471
Lipoptena depressa 428 Mutillidae 471
Lipoptena cervi 428 Pompilidae 471
Lipoptena mazamae 429 Sphecidae 471
Neolipoptena ferrisi 429 Social Wasps (Vespidae) 472
Pigeon Fly (Pseudolynchia canariensis) 429 Yellowjackets (Dolichovespula and Vespula) 473
Public Health Importance 429 Hornets (Vespa spp.) 475
Veterinary Importance 430 Paper Wasps (Polistes spp.) 476
Prevention and Control 434 Bees 476
References and Further Reading 435 Solitary Bees 476
Halictidae 476
21. Moths and Butterflies (Lepidoptera) Anthophoridae 477
Social Bees 477
Gary R. Mullen and Jennifer M. Zaspel
Apidae 477
Taxonomy 439 Bumble Bees (Bombus spp.) 477
Morphology 440 Honey Bees (Apis spp.) 478
Spicule Hairs 441 Public Health Importance 479
Spine Hairs 443 Veterinary Importance 483
Life History 443 Prevention and Control 483
Behavior and Ecology 443 References and Further Reading 484
Urticating Caterpillars 444
Lachryphagous Moths 450 23. Scorpions (Scorpiones)
Erebidae 450
Gary R. Mullen and W. David Sissom
Geometridae 450
Crambidae 451 Taxonomy 489
Notodontidae 451 Buthidae 490
Nolidae 451 Pseudochactidae 491
Sphingidae 451 Chaerilidae 491
Drepanidae 451 Chactidae 491
Wound-Feeding and Skin-Piercing Moths 452 Euscorpiidae 491
Public Health Importance 452 Superstitioniidae 491
Contents xi

Typhlochactidae 491 Theridiidae 510


Troglotayosicidae 491 Thomisidae 511
Akravidae 491 Trachelidae 511
Iuridae 492 Morphology 511
Caraboctonidae 492 Life History 513
Vaejovidae 492 Behavior and Ecology 513
Bothriuridae 492 Public Health Importance 514
Hormuridae 492 Tarantism 514
Heteroscorpionidae 493 Tarantulas 515
Hemiscorpiidae 493 Australian Funnel-Web Spiders 517
Urodacidae 493 South American Wandering Spiders 518
Diplocentridae 493 Yellow Sac Spiders 518
Scorpionidae 493 Hobo Spiders 518
Morphology 494 Recluse Spiders 519
Life History 496 Widow Spiders 523
Behavior and Ecology 497 Veterinary Importance 527
Public Health Importance 498 Prevention and Control 528
Scorpions of Medical Importance 500 References and Further Reading 529
Veterinary Importance 501
Prevention and Control 501 26. Mites (Acari)
References and Further Reading 502
Gary R. Mullen and Barry M. OConnor
24. Solpugids (Solifugae) Taxonomy 533
Morphology 534
Gary R. Mullen
Life History 535
References and Further Reading 506 Behavior and Ecology 536
Public Health Importance 536
25. Spiders (Araneae) Mite-Induced Dermatitis 537
Melicharidae 537
Gary R. Mullen and Richard S. Vetter
Dermanyssidae 537
Taxonomy 507 Macronyssidae 539
Mygalomorph Spiders 507 Laelapidae 541
Actinopodidae 507 Trombiculidae 542
Barychelidae 508 Stored-Products Mites 544
Dipluridae 508 Skin-Invading Mites 547
Atracidae 508 Sarcoptidae 549
Macrothelidae 508 Human Scabies 550
Theraphosidae 508 Human Notoedric Mange 552
Araneomorph Spiders 508 Mite-Induced Allergies 552
Agelenidae 508 Internal Acariasis 555
Araneidae 508 Mite-Borne Diseases of Humans 556
Ctenidae 509 Rickettsialpox 556
Desidae 509 Tsutsugamushi Disease 557
Dysderidae 509 Intermediate Hosts of Human Parasites 559
Eutichuridae 509 Delusory Acariasis and Acarophobia 559
Gnaphosidae 509 Veterinary Importance 559
Lamponidae 509 Mite-Induced Dermatitis 560
Lycosidae 509 Dermanyssidae 560
Miturgidae 510 Macronyssidae 560
Oxyopidae 510 Laelapidae 562
Pisauridae 510 Trombiculidae, Leeuwenhoekiidae,
Salticidae 510 Walchiidae 563
Segestriidae 510 Fur Mites 564
Sicariidae (Including the Former Cheyletidae 564
Loxoscelidae) 510 Myobiidae 565
xii Contents

Listrophoridae 566 Behavior and Ecology 613


Atopomelidae 567 Tick Species of Medical-Veterinary
Chirodiscidae 567 Importance 618
Myocoptidae 567 Public Health Importance 626
Feather Mites 568 Human Babesiosis 627
Mange Mites 568 Tick-Borne Encephalitis Complex 628
Laminosioptidae 571 Colorado Tick Fever 630
Demodecidae 572 Crimean-Congo Hemorrhagic Fever 631
Psorergatidae 574 Severe Fever and Thrombocytopenia
Sarcoptidae 575 Syndrome Virus 632
Other Sarcoptid Genera 578 Heartland Virus 632
Rhyncoptidae 580 Bourbon Virus Disease 632
Psoroptidae 581 Rocky Mountain Spotted Fever 632
Psoroptic Scab Mites 581 Boutonneuse Fever 635
Chorioptic Scab Mites 583 Other Spotted Fever Group Rickettsiae 635
Caparinic Scab Mites 584 Human Ehrlichiosis 636
Harpirhynchidae 585 Human Anaplasmosis 637
Hypoderatidae 585 Human Neoehrlichiosis 637
Mite-Induced Allergies 585 Q Fever 638
Internal Acariasis 585 Lyme Disease 638
Ear Mites 586 Tick-Borne Relapsing Fever and Borrelia
Respiratory Mites 588 miyamotoi Disease 643
Entonyssidae 588 Tularemia 646
Rhinonyssinae 589 Tick Paralysis 647
Halarachnidae 589 Tick-Bite Allergies 648
Ereynetidae 590 Veterinary Importance 648
Trombiculidae, Leeuwenhoekiide, and Piroplasmoses 649
Walchiidae 591 Diseases Caused by Members of the Family
Lemurnyssidae 591 Anaplasmataceae 652
Turbinoptidae 591 Prevention and Control 659
Cytoditidae 591 Personal Protection 659
Pneumocoptidae 592 Acaricides 660
Gastronyssidae 592 Pheromone-Assisted Control 661
Mite-Borne Diseases 592 Passive Treatment 661
Mites as Intermediate Hosts of Tapeworms 593 Hormone-Assisted Control 661
References and Further Reading 596 Biological Control 661
Vaccines 661
27. Ticks (Ixodida) Management 662
Eradication 663
William L. Nicholson, Daniel E. Sonenshine,
References and Further Reading 663
Bruce H. Noden, and Richard N. Brown
Taxonomy 603 28. Molecular Tools Used in Medical
Family Ixodidae (Hard Ticks) 604 and Veterinary Entomology
Family Argasidae (Soft Ticks) 605
Dana Nayduch, Rebecca Trout Fryxell, and
Family Nuttalliellidae 606
Pia Untalan Olafson
Morphology 606
External Anatomy 606 Molecular Techniques 673
Internal Anatomy 609 Nonmolecular Techniques 677
Life History 611 Genomics: Sequencing, Cataloging, and
Ixodid Life Cycles 611 Study of an Organism’s Genes 678
Argasid Life Cycles 612 Genome Projects of Important Arthropods 678
Contents xiii

Genome Sequencing 679 Appendix: Arthropod-Related Viruses


Analyzing Gene Expression 684 of Medical and Veterinary Importance 695
RNA Analysis 684
Protein Analysis 686 Michael J. Turell
Molecular Tools to Manipulate Arthropods 687
Editing Genomes With CRISPR-Cas9 687
Glossary 705
Insect Population Suppression 687
Posttranscriptional Gene Silencing With Taxonomic Index 741
RNAi 689
Subject Index 757
Conclusions and Future Directions 689
References and Further Reading 691
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Contributors

Peter H. Adler, Department of Plant and Environmental Jonas G. King, Department of Biochemistry, Molecular
Sciences, Clemson University, Clemson, SC, United Biology, Entomology and Plant Pathology, Mississippi
States State University, Starkville, MS, United States
Christopher M. Barker, Center for Vectorborne Diseases, Richard D. Kramer, Board Certified Entomologist (BCE),
School of Veterinary Medicine, University of California, CEO Innovative Pest Management, Inc., Brookeville,
Davis, CA, United States MD, United States
Richard J. Brenner, Director, ATIP Foundation, Arlington, William L. Krinsky, Division of Entomology, Peabody
TX, United States; Formerly Assistant Administrator of Museum of Natural History, Yale University, New
Technology Transfer, USDA, Agricultural Research Haven, CT, United States
Service, Beltsville, MD, United States Peter J. Landolt, Temperate Tree Fruit and Vegetable
Richard N. Brown, Department of Wildlife, Humboldt Research, USDA ARS, Wapato, WA, United States
State University, Arcata, CA, United States John E. Lloyd, Department of Renewable Resources,
Nathan D. Burkett-Cadena, University of Florida, Florida University of Wyoming, Laramie, WY, United States
Medical Entomology Laboratory, Vero Beach, FL, John W. McCreadie, Department of Biological Sciences,
United States University of South Alabama, Mobile, AL, United States
Ramón Cepeda-Palacios, Animal Science and Environ- Roger D. Moon, Department of Entomology, University of
mental Conservation Department, Autonomous Uni- Minnesota, St. Paul, MN, United States
versity of Baja California Sur, La Paz, Mexico
Gary R. Mullen, Department of Entomology and Plant
Douglas D. Colwell, Livestock Parasitology, Agriculture Pathology, Auburn University, Auburn, AL, United
and Agri-Food Canada, Lethbridge Research and States
Development Centre, Lethbridge, AB, Canada
Bradley A. Mullens, Department of Entomology, Uni-
Lance A. Durden, Department of Biology, Georgia versity of California, Riverside, CA, United States
Southern University, Statesboro, GA, United States
Leonard E. Munstermann, Yale University School of
Woodbridge A. Foster, Department of Evolution, Ecol- Public Health, New Haven, CT, United States
ogy, and Organismal Biology, and Department of
Entomology, The Ohio State University, Columbus, C. Steven Murphree, Biology Department, Belmont
OH, United States University, Nashville, TN, United States
Rebecca Trout Fryxell, Department of Entomology and Dana Nayduch, USDA-ARS, Arthropod-borne Animal
Plant Pathology, University of Tennessee, Knoxville, Diseases Research Unit, Manhattan, KS, United States
TN, United States William L. Nicholson, Rickettsial Zoonoses Branch,
Reid R. Gerhardt, Department of Entomology and Plant Division of Vector-Borne Diseases, National Center for
Pathology, University of Tennessee, Knoxville, TN, Emerging and Zoonotic Infectious Diseases, Centers for
United States Disease Control and Prevention, Atlanta, GA, United
States
Nancy C. Hinkle, Department of Entomology, University
of Georgia, Athens, GA, United States Bruce H. Noden, Department of Entomology and Plant
Pathology, Oklahoma State University, Stillwater, OK,
Lawrence J. Hribar, Florida Keys Mosquito Control United States
District, Marathon, FL, United States

xv
xvi Contributors

Barry M. OConnor, Museum of Zoology, University of W. David Sissom, Department of Life, Earth and Envi-
Michigan, Ann Arbor, MI, United States ronmental Sciences, West Texas A&M University,
Pia Untalan Olafson, USDA-ARS, Livestock Arthropod Canyon, TX, United States
Pests Research Unit, Kerrville, TX, United States Daniel E. Sonenshine, Department of Biological Sciences,
Hal C. Reed, Department of Biology and Chemistry, Oral Old Dominion University, Norfolk, VA, United States
Roberts University, Tulsa, OK, United States Michael J. Turell, Virology Division, United States Army
Will K. Reeves, U.S. Department of Agriculture, Animal Medical Research Institute of Infectious Diseases, Fort
and Plant Health Inspection Service, Biotechnology Detrick, Frederick, MD, United States (retired)
Resource Services, Western Compliance Assurance Richard S. Vetter, Department of Entomology, University
Branch, Fort Collins, CO, United States of California, Riverside, CA, United States
William K. Reisen, Center for Vectorborne Diseases, Edward D. Walker, Department of Microbiology and
School of Veterinary Medicine, University of California, Molecular Genetics, and Department of Entomology,
Davis, CA, United States Michigan State University, East Lansing, MI, United
Justin O. Schmidt, Southwestern Biological Institute, States
Tucson, AZ, United States; Department of Entomology, Jennifer M. Zaspel, Milwaukee Public Museum,
University of Arizona, Tucson, AZ, United States Milwaukee, WI, United States
Philip J. Scholl, United States Department of Agriculture,
Agricultural Research Service (retired), Oxford, FL,
United States
Preface

It has been 10 years since the publication of the second entomological, medical, and other terms used in the book,
edition of this book. Significant advances have been made with which she or he may not be familiar. We hope this will
in our knowledge of a plethora of arthropod-related prob- be helpful to the widest possible range of readers, spe-
lems and diseases, including changes in their geographic cialists and nonspecialists alike, in diverse disciplines
ranges and the emergence of new or previously unrecog- relating either directly or indirectly to the subject matter.
nized arthropod-borne diseases of medical and veterinary The text is illustrated with 538 figures and 22 revised, or
concern. Among notable examples of the latter are Zika and new, color maps.
Chikungunya viruses transmitted by mosquitoes and a In addition to its value to students as a textbook, this
number of tick-borne pathogens, such as Bourbon, Heart- volume is intended for a much broader audience as a
land, and severe fever with thrombocytopenia syndrome comprehensive reference source for biologists in general,
viruses, as well as newly recognized forms of ehrlichiosis, entomologists, zoologists, parasitologists, physicians, pub-
rickettsiosis, and babesiosis. A new chapter has been lic health personnel, veterinarians, wildlife biologists,
addeddChapter 3: Arthropod Toxins and Venomsdto vector biologists, military and armed forces entomologists,
provide an overview of the structural nature and diversity the general public, and others looking for a readable,
of chemical compounds that play an important medical- authoritative source of information on this important topic.
veterinary role, particularly among stinging and biting in- We welcome as new contributing authors to the book
sects and arachnids. In addition, Chapter 28: Molecular the following 14 individuals: Justin O. Schmidt (Chapter 3),
Tools Used in Medical and Veterinary Entomology has Christopher M. Barker (Chapter 4), Lawrence J. Hribar
been expanded, reflecting the many significant advances (Chapter 11), Leonard E. Munstermann (Chapter 12), C.
and applications of molecular and genetic techniques in Steven Murphree (Chapter 13), Douglas D. Colwell and
recent years, which have become such an integral part of Ramón Cepeda-Palacios (Chapter 19), Will K. Reeves
medical-veterinary entomology today. (Chapter 20), Jennifer M. Zaspel (Chapter 21), W. David
One of the primary objectives of the first and second Sissom (Chapter 23), Bruce H. Noden and Richard N.
editions has been to provide a textbook suitable for teach- Brown (Chapter 27), and Rebecca Trout Fryxell
ing courses in medical and veterinary entomology at the and Pia Untalan Olafson (Chapter 28). Together with 24
college and university levels. In keeping with that goal and continuing authors of the second edition and the Graphics
the format of the previous editions, the book is organized Editor, 40 contributors in all, they have helped significantly
from an entomological perspective, with each chapter in revising the respective chapterseproviding new
devoted to a particular taxonomic group of insects or perspectives and achieving an appropriate balance between
arachnids (including spiders, scorpions, solpugids, mites, medical and veterinary entomology as closely related
and ticks). Each chapter includes the following sub- disciplines.
headings: Taxonomy, Morphology, Life History, Behavior The success of the previous two editions is reflected by
and Ecology, Public Health Importance, Veterinary the widespread adoption of this book for teaching medical
Importance, Prevention and Control, and References and and/or veterinary-related courses at colleges and univer-
Further Reading. The separate sections on public health and sities throughout the United States and other parts of the
veterinary entomology are designed to assist instructors in world. We hope the third edition will be equally successful
using this book to teach courses in either medical or vet- in helping to educate and inspire the next generation of
erinary entomology, or courses combining these two related medical and veterinary entomologists.
disciplines. The book concludes with an Appendix titled
“Arthropod-Related Viruses of Medical and Veterinary Gary R. Mullen
Importance” and a Glossary of approximately 1,700 terms. Lance A. Durden
The latter is intended to assist the reader in understanding

xvii
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Acknowledgments

As with any undertaking of this magnitude, there are many Figs. 28.1e28.4, and Boxes 28.1e28.3); Margo A. Duncan
individuals to whom the editors are indebted. Foremost are (Gainesville, FL), for her original illustrations, particularly
the contributing authors, whose combined expertise and those in the Lepidoptera chapter (original Fig. 21.4 and
commitment to promoting medical and veterinary ento- redrawn Figs. 21.2, 21.3 and 21.5) that appear in the first,
mology have made the third edition of this book possible. second, and third editions; Rebecca L. Nims (Social Circle,
We also owe a debt of gratitude to the many other GA), for the outstanding work she did in redrawing from
individuals who contributed so generously of their time in various sources more than half the black-and-white figures
reviewing chapters or select parts of the text, offering for the first and second editions (notably for the mite
suggestions for improving and updating material for the chapter), virtually all of which have been retained in the
third edition, preparing illustrations, and providing original third edition; and Philip B. Shevlin (Auburn University,
photographs and new color images for the 538 figures. Auburn, AL) for his assistance with preparation of the
We particularly want to recognize the following people chemical structures of arthropod toxins and venoms
who have contributed significantly in preparing the figures, (Figs. 3.3e3.8).
maps, and other illustrations throughout the book: first and We also acknowledge the following individuals for
foremost, Jonas G. King (Mississippi State University, preparing original illustrations, primarily line drawings,
Starkville, MS) for his invaluable graphics skills and all- which appear within these pages: Woodbridge A. Foster
important role in ensuring the high-quality color images (The Ohio State University, Columbus, OH; Fig. 15.14);
and maps throughout the chapters; a special thanks to Jillian Susan Milna Hope (Mebane, NC; Fig. 16.1); Jonas G. King
Masters, an undergraduate student working with Jonas (Mississippi State University, Starkville, MS; Figs. 14.1
King, for her welcomed talent and generous time assisting and 18.12); Takumasa Kondo (Palmira, Valle, Colombia;
with the figures, particularly the maps; and graduate student Fig. 11.9); William L. Krinsky (Yale University, New
Anagha Gopakumar, also a member of the King lab group at Haven, CT; Figs. 5.3, 8.7 and 8.11); and Blair Sampson
Mississippi State for her part in preparing illustrations; (US Department of Agriculture, Agricultural Research
Lorenza Beati and Lance A. Durden (Georgia Southern Service, Poplarville, MS; Fig. 22.6).
University, Statesboro, GA) for preparing the stacked im- The following individuals have kindly provided original
ages of the lice (Figs. 7.5e7.7, 7.12e7.16 and 7.19) and photographs, slides, or digital images for reproduction as
fleas (Figs. 10.1, 10.8e10.12, 10.14, 10.17e10.19 and figures in this edition, or have contributed in various other
10.21, and Oriental rat flea on book cover); Nathan D. ways:
Burkett-Cadena (University of Florida, Florida Medical W. V. Adams, Jr. (Louisiana State University, Baton
Entomology Laboratory, Vero Beach, FL), for providing the Rouge, LA; Fig. 16.13); Peter H. Adler (Clemson
original color illustrations that accompany his chapter on University, Clemson, SC; Fig. 14.7); Roger D. Akre
morphological adaptations of parasitic arthropods (Washington State University, Pullman, WA; Figs. 22.14,
(Figs. 2.1e2.5), preparing original illustrations for the 22.15, 22.24 and 22.31); Karim Aoun and Aïda Bouratbine
epidemiology chapter (Figs. 4.2 and 4.3), and providing 12 (Institut Pasteur de Tunis, Tunisia; Fig. 12.14); Stephen
additional figures throughout the book (see below); E. Paul Ausmus (US Department of Agriculture; Fig. 22.25); Hans
Catts (Washington State University, Pullman, WA), for Bänziger (Chaing Mai University, Thailand; Figs. 21.19,
being able to include in this third edition his superb original 21.20 and 20.22); Martin Bennett (Creative Commons;
line drawings of myiasis-causing flies that he prepared for Fig. 19.21); Robert B. Berry (Orange Breasted Falcon
the first edition, prior to his untimely death in 1996 Project, The Peregrine Fund, Wolf, WY; Fig. 17.22);
(Figs. 19.1e19.9, 19.11, 19.13e19.15, 19.17, 19.20, 19.23, Yehuda Braverman (Kimron Veterinary Institute, Israel;
19.27, 19.32, 19.33, 19.36, 19.37 and 19.40); Victoria Fig. 12.9); Alberto B. Broce (Kansas State University,
Rhodes (Missouri Southern State University, Joplin, MO), Manhattan, KS); Nathan D. Burkett-Cadena (University of
for her excellent color illustrations, Chapter 28, Florida, Florida Medical Entomology Laboratory, Vero

xix
xx Acknowledgments

Beach, FL; Figs. 5.1, 5.4, 6.4, 7.9, 10.4e10.7, 16.3, 20.6, Richard C. Lancaster (Agriculture and Agri-Food Canada,
25.6 and 26.11); João P. Burini (Pontifica Universidade Lethbridge, AB; Fig. 13.2); Lloyd L. Lauerman (Alabama
Católica de São Paulo, Brazil; Figs. 23.2, 23.14, 25.4, 25.5 State Veterinary Diagnostic Laboratory Auburn, AL;
and 25.10); Lyle Buss (University of Florida, Gainesville, Figs. 13.10e13.12); Matthew S. Lehnert (Kent State Uni-
FL; Fig. 21.25); Jerry F. Butler (University of Florida, versity at Stark, North Canton, OH; Figs. 21.1B,C and
Gainesville, FL; Figs. 9.3B, 20.5, 21.13, 25.25, 26.40, 21.24); John E. Lloyd, Sr. (University of Wyoming, Lar-
26.43 and 27.11); Bonnie Buxton (Philadelphia College of amie, WY; Figs. 7.19, 7.21, 20.3, 20.8 and 20.12e20.14);
Osteopathic Medicine-Georgia Campus, Suwanee, GA; Amanda D. Loftis (Ross University School of Veterinary
Fig. 15.42); Brittany E. Campbell (University of Florida, Medicine, Basseterre, Saint Kitts, West Indies; Fig. 20.1);
Gainesville, FL; Fig. 8.13); Luis Paulo Carvalho (Federal Stephen A. Marshall (University of Guelph, Guelph, ON,
University of Rond onia-Unir, Porto Velho, Brazil; Canada; Figs. 13.1, 14.2A and 14.4); Sturgis McKeever
Fig. 13.14); James Castner (University of Florida, Gain- (Georgia Southern University, Statesboro, GA; Figs. 16.5,
esville, FL; Figs. 6.3, 6.5e6.12 and 21.26); Ronald D. 19.18, 21.6e21.10, 21.14, 21.18 and 25.21, and saddleback
Cave (Indian River Research and Education Center, Uni- caterpillar on book cover); Mae Melvin (Centers for Dis-
versity of Florida, Fort Pierce, FL; Fig. 19.26); Valerie J. ease Control and Prevention, Atlanta, GA; Fig. 8.8); Hen-
Cervenka (University of Minnesota, St. Paul, MN, drick J. Meyer (North Dakota State University, Fargo, ND;
Fig. 17.23); Chan Chee Keong (Panang Island, Malaysia; Fig. 17.25); Camille Meyers (Courtesy of Peregrine
Fig. 9.6); Jillian Cowles (Vail, AZ; Fig. 3.9); Douglas A. Fund; Fig. 17.22); Steve Mihok (Russell, ON, Canada;
Craig (University of Alberta, Edmonton, AB, Canada; Fig. 16.10D); Roger D. Moon (University of Minnesota, St.
Figs. 14.2B and 14.3); Eddie W. Cupp (Owensboro, KY; Paul, MN; Figs. 17.16 and 17.19); Gary R. Mullen (Auburn
Figs. 14.12 and 14.14); Aaron T. Dossey (Gainesville, University, Auburn, AL; Figs. 8.16, 9.3A, 13.5, 13.6,
FL; Fig. 1.5); S. Allen Edgar (Auburn University, Auburn, 13.13, 13.14, 17.18, 17.20, 19.19, 21.11, 22.9, 22.19,
AL; Figs. 15.37 and 15.38); Marc E. Epstein (California 22.21, 22.22, 22.29, 22.30, 25.13 and 27.9); Bradley A.
Department of Food and Agriculture, Sacramento, CA; Mullens (University of California, Riverside, CA;
Fig. 21.12); John Flannery (Richmond County, NC; Figs. 16.8 and 16.10B); Teresa R. Mullens (Riverside, CA;
Fig. 21.1A); Woodbridge A. Foster (The Ohio State Uni- Fig. 16.10C); Leonard E. Munstermann (Yale University,
versity, Columbus, OH; Figs. 15.11, 15.17, 15.19, 15.26, New Haven, CT; Figs. 12.5e12.8 and 12.10); Amy C.
15.29 and 15.34); Wes Gaston (US Department of Agri- Murillo (University of California, Riverside, CA;
culture, Animal and Plant Health Inspection Service, Figs. 7.17AeF, 7.22, 10.13 and 26.25); Harold D. Newson
Wildlife Services; Fig. 19.41); James L. Gathany (Centers (Michigan State University, East Lansing, MI; Figs. 15.39
for Disease Control and Prevention, Atlanta, GA; Figs. 7.4, and 15.41); William L. Nicholson (Centers for Disease
8.3, 11.2, 12.2, 27.5 and 27.17); Carolyn Grissom (Shel- Control and Prevention, Atlanta, GA; Fig. 27.10; Susan M.
byville, TN; Fig. 25.17); Duane J. Gubler (Duke-National Noh (US Department of Agriculture, Pullman, WA; Fig.
University of Singapore Graduate Medical School, 27.30); Yoshiro Ohara (Kanazawa Medical University,
Singapore; Fig. 15.25); Hadel H. Go (American Museum of Uchinada, Ishikawa, Japan; Fig. 16.12); Christopher D.
Natural History, New York, NY; for a taxonomic clarifi- Paddock (Centers for Disease Control and Prevention,
cation regarding the Hymenoptera); Robert G. Hancock Atlanta, GA; Figs. 27.22 and 27.23); Jonathan D. Patterson
(University of the Cumberlands, Williamsburg, KY; (Michigan State University, East Lansing, MI; Fig. 15.40);
Fig. 15.23); Elton J. Hansens (Asheville, NC; Figs. 7.8, Laverne L. Pechuman (Cornell University, Ithaca, NY;
10.20, 16.7, 16.10A and 17.12); Mark Harvey (Western Fig. 16.6); Nicholas Perrault (Creative Commons;
Australia Museum, Welshpool, Western Australia; for Fig. 14.8); Eric Poggenphol (Eric Poggenphol Photog-
taxonomic information on the Solifugae); Nancy C. Hinkle raphy, Northampton, MA; Fig. 14.13); Lorenzo Prendini
(University of Georgia, Athens, GA; Fig. 9.11); Mac (American Museum of Natural History, New York; for his
Horton (Clemson University, Clemson, SC; Fig. 22.28); contributed expertise on scorpions); Robert J. Raven
Kevin Humphries (Huntsville, AL; Fig. 25.16); Lacy L. (Queensland Museum, South Brisbane, Australia;
Hyche (Auburn University, Auburn, AL; Fig. 21.15); Fig. 25.9); Charles Ray (Auburn University, AU Plant
Gregory D. Johnson (Montana State University, Bozeman, Diagnostic Laboratory, Auburn, AL; for providing records
MT; Fig. 7.20); Jena Johnson (Jena Johnson Photography of insects and mites biting humans); David E. Reed
and University of Georgia, Athens, GA; Figs. 14.1, 14.5, (Chanhassen, MN; Fig. 9.4); Hal C. Reed (Oral Roberts
14.6, 22.13 and 22.20; book-cover images, Asian tiger University, Tulsa, OK; Figs. 22.2, 22.5, 22.10, 22.16 and
mosquito and green-bottle fly); Florian Karolyi (University 22.18); Lawrence E. Reeves (University of Florida, Gain-
of Vienna, Austria; Fig. 16.4); Takumasa Kondo (Palmira, esville, FL; Fig. 19.24); Jon Richfield (Creative Commons;
Valle, Colombia; Fig. 22.23); William L. Krinsky (Yale Fig. 24.1); Mary Elizabeth Hayes Rogers (Waukegan, IL;
University, New Haven, CT; Figs. 5.2, 5.5 and 8.6); Fig. 13.18); Christopher Saski (Clemson University,
Acknowledgments xxi

Genomics and Computational Laboratory, Clemson, SC; Study (SCWDS), University of Georgia, Athens, GA;
for his review and editorial suggestions, Chapter 28); Philip University of Florida/IFAS Extension, Gainesville, FL; US
J. Scholl (US Department of Agriculture, Agricultural Armed Forces Institute of Pathology (AFIP), disestablished
Research Service, Oxford, FL; Fig. 19.35); Justin O. in 2011; US Department of Agriculture (USDA), including
Schmidt (US Department of Agriculture, Carl Hayden Bee the Animal and Plant Health Inspection Service (APHIS),
Research Center, Tucson, AZ; Fig. 22.27); Joseph A. Animal Research Service (ARS) at Kerrville, TX, and
Schemanchuk (Department of Agriculture and Agri-Food Laramie, WY, Wildlife Services (WS), and Foreign Animal
Canada, Lethbridge, AB, Canada; Figs. 14.15 and 14.16); Disease Diagnostic Laboratory (Plum Island, NY); US
D. Craig Sheppard (University of Georgia, Athens, GA; National Tick Collection, Georgia Southern University,
Fig. 17.21); W. David Sissom (West Texas A&M Uni- Statesboro, GA; US Forest Service; US Public Health
versity, Canyon, TX; Figs. 23.1, 23.2, 23.6, 23.9 and Service; The Wellcome Collection, London; Wikimedia
23.10e23.13, and scorpion on book cover); Rachel Stone Commons; and World Health Organization (WHO), Vector
(Wichita State University, Wichita, KS; Fig. 9.9); Daniel R. Control and Prevention Programme.
Suiter (University of Georgia, Griffin, GA; Fig. 6.2); It is our special privilege to recognize and thank the
Robert B. Tesh (University of Texas Medical Branch, following six previous chapter authors of both the first and
Galveston, TX; Fig. 8.5); Gerrit Uilenberg (Corsica, second editions of this book who are not contributors to the
France; Figs. 27.28, 27.29, 27.31 and 27.32); Richard third edition: Raj K. Gupta (Walter Reed Army Institute of
Urbanek (US Fish and Wildlife Service; Figs. 14.7 and Research, Silver Spring, MD); Robert D. Hall (University
14.8); Richard S. Vetter (University of California, River- of Missouri, Columbia, MO); Robert S. Lane (University of
side, CA; Figs. 25.12, 25.15, 25.19 and 25.20); P. Kirk California, Berkeley, CA); Louis C. Rutledge (US Army,
Visscher (University of California, Riverside, CA; retired, Mill Valley, CA); Scott A. Stockwell (US Army,
Fig. 13.7); Jan Votýpka (Czech Academy of Science, retired, Lubbock, TX); also, Gerritt Uilenberg (Corsica,
Czech Republic; Fig. 14.19); Laurel L. Walters (Lieen- France), as a chapter author of the second edition. Their
Follican Research, Bishop, CA; Fig. 12.13); Wesley D. invaluable contributions continue to be reflected in this new
Watson (University of North Carolina, Raleigh, NC; edition of Medical and Veterinary Entomology.
Fig. 9.10); Jerry Weintraub (Agriculture and Agri-Food At the same time, we are pleased to welcome as new
Canada, Lethbridge, AB; Figs. 19.28 and 19.31); Julian contributing authors the following 12 individuals: Chris-
White (Toxicology Department, Women’s & Children’s topher M. Barker, Ramón Cepeda-Palacios, Douglas D.
Hospital, North Adelaide, Australia; Figs. 25.8 and 25.23); Colwell, Lawrence J. Hribar, Leonard E. Munstermann, C.
Ralph E. Williams (Purdue University, West Lafayette, IN; Steven Murphree, Pia Untalan Olafson, Will K. Reeves,
Fig. 17.17); Germano Woehl, Jr. (Instituto Rã-bugio para Justin O. Schmidt, W. David Sissom, Rebecca Trout
Conservação da Biodiversidade, Jaraguá do Sul, Brazil; Fryxell, and Jennifer M. Zaspel.
Fig. 21.16); and Jennifer M. Zaspel (Milwaukee Public In addition, we honor the memory of the following four
Museum, Milwaukee, WI; Figs. 21.21 and 21.23). chapter authors of the first edition of this book: Roger D.
Appreciation is also extended to the following in- Akre and E. Paul Catts (Department of Entomology,
stitutions and agencies for their courtesy in granting Washington State University, Pullman, WA), Robert Traub
permission to reproduce images, providing information, (US Army, retired, Bethesda, MD); and John E. “Jack”
and otherwise affording helpful assistance to us: American Lloyd (University of Wyoming, Laramie, WY), a contrib-
Museum of Natural History, New York, NY; Auburn utor to the first and second editions, who died March 25,
University College of Veterinary Medicine, Department of 2017, while co-revising his chapter for the third edition. We
Pathobiology, Auburn, AL; Entomological Society of dedicate this book to their memory and to the many other
America, Annapolis, MD; The Carter Center, Onchocerci- medical and veterinary entomologists who have devoted
asis Elimination Program for the Americas (OEPA), their careers to protecting humans and animals from inju-
Atlanta, GA; Centers for Disease Control and Prevention rious arthropods and vector-borne diseases.
(CDC) and Public Health Image Library (PHIL); Creative And, finally, we gratefully acknowledge Kristi A.
Commons; Department of Agriculture and Agri-Food Gomez (Acquisitions Editor), Pat Gonzalez (Editorial
Canada, Lethbridge, AB; Food and Agricultural Organi- Project Manager), Punitha Radjane (Senior Project
zation of the United Nations, Rome, Italy; The Joint Manager), Swapna Praveen (Permissions Coordinator),
Pathology Center (JPC), US Federal Government, Silver Allison Esposito (Copyeditor), and the fine staff and
Spring, MD; National Geographic Society, Washington, support personnel at Elsevier, Inc., who have worked so
DC; New York Entomological Society, New York, NY; closely with us in preparing and publishing the third edition
The Natural History Museum, London; Otis Historical of this book.
Archives (OHA), National Museum of Health and Gary R. Mullen
Medicine, Silver Spring, MD; The Rockefeller Foundation, Lance A. Durden
New York, NY; Southeastern Cooperative Wildlife Disease
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Chapter 1

Introduction
Lance A. Durden1 and Gary R. Mullen2
1
Department of Biology, Georgia Southern University, Statesboro, GA, United States; 2Department of Entomology and Plant Pathology,
Auburn University, Auburn, AL, United States

Medical entomology is the study of insects, insect-borne arthropods, are causing newly recognized diseases
diseases, and other associated problems that affect (e.g., Lyme disease and human granulocytic anaplasmosis)
humans and public health. Veterinary entomology is the and the resurgence of diseases that had been suppressed for
study of insects and insect-related problems that affect many years (e.g., malaria, Chikungunya fever, and Zika
domestic animals, particularly livestock and companion encephalitis). Emerging and resurging arthropod-borne
animals (dogs, cats, horses, caged birds, etc.). In addition, diseases are recognized as a growing health concern by
veterinary entomology includes insect-associated problems public health and veterinary officials (Wilson and Spiel-
affecting captive animals in zoological parks and in wildlife man, 1994; Walker et al., 1996; Gubler, 1998; Winch,
in general. Medical-veterinary entomology combines 1998; and Gratz, 1999; Marcondes, 2016).
these two disciplines.
Traditionally, the fields of medical and veterinary
entomology have included health-related problems
GENERAL ENTOMOLOGY
involving arachnids (particularly mites, ticks, spiders, and Basic concepts of entomology such as morphology, tax-
scorpions). This broad approach that encompasses onomy and systematics, developmental biology, and ecol-
insects and arachnids is followed in this text. Alterna- ogy provide important background information for medical
tively, the study of health-related problems involving and veterinary entomologists. General entomology books
arachnids is called medical-veterinary arachnology or, that the reader will find helpful in this regard include those
if just mites and ticks are considered, medical-veterinary by Gillot (1995), Elzinga (2000), Chapman (1998),
acarology. Romoser and Stoffolano (1998), Gullan and Cranston
Historically, both medical and veterinary entomology (2005), Triplehorn and Johnson (2005), and Pedigo and
have played major roles in the development of human Rice (2009). References that provide a more taxonomic or
civilization and animal husbandry. Outbreaks of insect- biodiversity-oriented approach to general entomology
borne diseases of humans have profoundly influenced hu- include works by Arnett (2000), Richards and Davies
man history; such diseases include yellow fever, plague, (1994), Bosik (1997), Daly et al. (1998), and Marshall
louse-borne typhus, malaria, African trypanosomiasis, (2006). General insect morphology is detailed in Snodgrass
Chagas disease, and lymphatic filariasis. Likewise, live- (1993), whereas a useful glossary of general entomology is
stock scourges such as bovine babesiosis, bovine theiler- provided by Torre-Bueno (1962) and was updated and
iosis, scabies, pediculosis, and botfly infestations, all of revised by Nichols (1989). An encyclopedia of entomology
which are caused or transmitted by arthropods, have greatly (Resh and Cardé, 2009) and a dictionary of entomology
influenced animal production and husbandry practices. (Gordh and Headrick, 2001) are also available. Texts on
Arthropod-related disorders continue to cause significant urban entomology, the study of insect pests in houses,
health problems to humans, domestic animals, and wildlife. buildings, and urban areas, which also has relevance to
At the same time, new strains of known pathogens, as well medical-veterinary entomology, have been prepared by
as previously unrecognized disease agents transmitted by Ebeling (1975), Hickin (1985), Mallis et al. (2004), and

Medical and Veterinary Entomology. https://doi.org/10.1016/B978-0-12-814043-7.00001-7 1


Copyright © 2019 Elsevier Inc. All rights reserved.
2 Medical and Veterinary Entomology

Robinson (1996). General texts on acarology include works l Advances in Disease Vector Research, published by
by Woolley (1987), Evans (1992), and Krantz and Walter Springer-Verlag.
(2009). l Bulletin of the World Health Organization, published
by the World Health Organization.
l Journal of Wildlife Diseases, published by the Wildlife
MEDICAL-VETERINARY ENTOMOLOGY
Disease Association.
LITERATURE l Emerging Infectious Diseases, published by the Centers
Textbooks or monographs pertaining to medical ento- for Disease Control and Prevention (CDC).
mology, veterinary entomology, or the combined discipline l American Journal of Tropical Medicine and Hygiene,
of medical-veterinary entomology are listed under these published by the American Society of Tropical Medi-
headings at the end of this chapter. Most of these publi- cine and Hygiene.
cations emphasize arthropod morphology, biology, sys- l Memorias Do Instituto Oswaldo Cruz, published in
tematics, and disease relationships, whereas some texts Brazil.
emphasize molecular aspects of medical-veterinary ento-
Various Internet websites pertaining to medical-
mology, such as Crampton et al. (1997) and Marquardt
veterinary entomology also can be accessed for useful in-
et al. (2005). Other works are helpful regarding common
formation. The CDC in Atlanta, Georgia, USA, provides
names of arthropods of medical-veterinary importance
two helpful resources: (1) a pictorial key to arthropods of
(Pittaway, 1992), surveillance techniques (Bram, 1978),
public health importance, available as downloadable files
control measures (Drummond et al., 1988), repellents
(PDF), https://www.cdc.gov/nceh/ehs/Publications_Keys.
(Debboun et al., 2007; Strickman et al., 2009), or ecto-
htm; and (2) a comprehensive annual guide titled the
parasites (Andrews, 1977; Marshall, 1981; Kim, 1985; “Yellow Book” that details travel medicine updates on a
Uilenberg, 1994; Barnard and Durden 1999; Clayton et al.,
global basis (CDC, 2018). Although all aspects of travel
2015). Publications that devote substantial sections to ar-
medicine and infectious diseases are covered, much of the
thropods associated with wildlife and the pathogens they
information addresses vector-borne diseases.
transmit include those by Davidson et al. (1981), Fowler
(1986), Davidson and Nettles (1997), and Samuel et al.
(2001). A BRIEF HISTORY OF MEDICAL-
Several journals and periodicals are devoted primarily VETERINARY ENTOMOLOGY
to medical and/or veterinary entomology. These include:
Problems caused by biting and annoying arthropods and the
l Journal of Medical Entomology, published for the Ento- pathogens they transmit have been the subject of writers
mological Society of America by Oxford University since antiquity (Service, 1978). Homer (mideeighth cen-
Press. tury BC), Aristophanes (c. 448e380 BC), Aristotle
l Medical and Veterinary Entomology, published by the (384e322 BC), Plautus (c. 254e184 BC), Columella (5
Royal Entomological Society (UK). BCeAD 65), and Pliny (AD 23e79) all wrote about the
l Journal of Vector Ecology, published by the Society of nuisance caused by flies, mosquitoes, lice, and/or bedbugs.
Vector Ecologists. However, the study of modern medical-veterinary ento-
l Review of Medical and Veterinary Entomology, pub- mology is usually recognized as beginning in the late 19th
lished by CAB International. century, when blood-sucking arthropods were first proved
l Annals of Medical Entomology, published in Bhopal, to be vectors of human and animal pathogens.
India. Englishman Patrick Manson (1844e1922) (Fig. 1.1A)
l Vector-Borne and Zoonotic Diseases, published in the was the first to demonstrate pathogen transmission by a
United States by Mary Ann Liebert, Inc. blood-feeding arthropod. Working in China in 1877, he
showed that the mosquito Culex pipiens fatigans is a vector
Journals specializing in parasitology, tropical medicine,
of Wuchereria bancrofti, the causative agent of Bancroftian
or wildlife diseases that also publish articles on medical-
filariasis. After this landmark discovery, the role of various
veterinary entomology include:
blood-feeding arthropods in transmitting pathogens was
l Parasitology, published by the British Society for recognized in relatively rapid succession.
Parasitology. In 1891, Americans Theobald Smith (1859e1934)
l Journal of Parasitology, published by the American So- (Fig. 1.1B) and Frederick L. Kilbourne (1858e1936)
ciety of Parasitologists. implicated the cattle tick, Rhipicephalus (Boophilus)
l Parasite-Journal de la Société Française de Parasitolo- annulatus, as a vector of Babesia bigemina, the causative
gie, published in France. agent of Texas cattle fever (bovine babesiosis). This paved
Introduction Chapter | 1 3

(A) (B) (C)

FIGURE 1.1 Historical figures in medical-veterinary entomology. (A) Patrick Manson (1844e1922). (B) Theobald Smith (1859e1934). (C) Ronald
Ross (1857e1932). A and B, Courtesy of The Wellcome Collection, London: C, Courtesy of Wikipedia Commons.

the way for a highly successful R. annulatus eradication (1833e1915) (Fig. 1.2A) presented persuasive evidence for
program in the United States directed by the US Depart- his theory that what we know today as the yellow fever
ment of Agriculture (USDA). The eradication of this tick mosquito, Aedes aegypti, was the vector of this virus.
resulted in the projected goal d elimination of indigenous However, it was not until 1900 that American Walter Reed
cases of Texas cattle fever throughout the southern United (1851e1902) (Fig. 1.2B) led the US Yellow Fever Com-
States. mission at Havana, Cuba, which proved A. aegypti to be the
In 1898, Englishman Sir Ronald Ross (1857e1932) principal vector of yellow fever virus.
(Fig. 1.1C), working in India, demonstrated the role of In 1903, Englishman David Bruce (1855e1931)
mosquitoes as vectors of avian malarial parasites from (Fig. 1.2C) demonstrated the ability of the tsetse fly Glos-
diseased to healthy sparrows. Also in 1898, the cyclical sina palpalis to transmit, during blood feeding, the try-
development of malarial parasites in anopheline mosquitoes panosomes that cause African trypanosomiasis.
was described by Italian Giovani Battista Grassi Other important discoveries continued well into the
(1854e1925). In the same year, Frenchman Paul Louis 20th century. In 1906, American Howard Taylor Ricketts
Simond (1858e1947), working in Pakistan (then part of (1871e1910) (Fig. 1.3) proved that the Rocky Mountain
India), showed that fleas are vectors of the bacterium that wood tick, Dermacentor andersoni, is a vector of Rickettsia
causes plague. rickettsii, the causative agent of Rocky Mountain spotted
In 1848, American physician Josiah Nott (1804e1873) fever. In 1907, F. Percival Mackie (1875e1944) showed
of Mobile, Alabama, published circumstantial evidence that that human body lice are vectors of Borrelia recurrentis,
led him to believe that mosquitoes were involved in the the spirochete that causes louse-borne (epidemic) relapsing
transmission of yellow fever virus to humans. In 1881, the fever. In 1908, Brazilian Carlos Chagas (1879e1934)
Cuban-born Scottish physician Carlos Finlay (Fig. 1.4) demonstrated transmission of the agent that

(A) (B) (C)

FIGURE 1.2 Historical figures in medical-veterinary entomology. (A) Carlos Finlay (1833e1915). (B) Walter Reed (1851e1902). (C) David Bruce
(1855e1931). A and C, Courtesy of The Wellcome Collection, London: B, Courtesy of Wikipedia Commons.
4 Medical and Veterinary Entomology

discussed in more detail in the references listed at the end of


this chapter. Because of the chronology of many major
discoveries relevant to this topic in the 50-year period
starting in 1877, this time has been called the “golden age
of medical-veterinary entomology” (Philip and Rozeboom,
1973).

IDENTIFICATION AND SYSTEMATICS OF


ARTHROPODS OF MEDICAL-VETERINARY
IMPORTANCE
Table 1.1 provides a list of the eight orders of insects and
four orders of arachnids that are of particular interest to
medical-veterinary entomologists. Accurate identification
of these arthropods is an important first step in determining
the types of problems they can cause and, subsequently, in
implementing control programs.
FIGURE 1.3 Howard Taylor Ricketts (1871e1910). Courtesy of The Although taxonomy and identification are discussed in
Wellcome Collection, London. more detail with respect to arthropod groups treated in the
chapters that follow, some publications provide a broader
perspective on the classification, taxonomy, and identifi-
cation of a range of arthropods of medical-veterinary
importance. These include two works published by the
CDC (1979, 1994), as well as Service (1988), Hopla et al.
(1994), Lago and Goddard (1994), and Davis (1995). Also,
some medical-veterinary entomology books are very taxo-
nomically oriented, with emphasis on identificationdfor
example, Baker et al. (1956), Smith (1973), Lane and
Crosskey (1993), and Walker (1995).

TYPES OF PROBLEMS CAUSED BY


ARTHROPODS
Annoyance
Regardless of their role as blood-feeders (hematophages),
parasites, or vectors of pathogens, certain arthropods cause
severe annoyance to humans or other animals because of
FIGURE 1.4 Carlos Chagas (1879e1934). Courtesy of Wikipedia their biting behavior. These arthropods include lice, bed-
Commons. bugs, fleas, deer flies, horse flies, tsetse flies, stable flies,
mosquitoes, black flies, biting midges, sand flies, chiggers,
and ticks. Some arthropods, however, do not bite but instead
causes American trypanosomiasis, later named Chagas are annoying because of their abundance, small size, or habit
disease in his honor, by the conenose bug Panstrongylus of flying into or around the eyes, ears, and nose. Nonbiting
megistus. In 1909, Frenchman Charles Nicolle arthropods that cause annoyance include the house fly,
(1866e1936), working in Tunis, showed that human body chironomid midges, and eye gnats. Large populations of
lice are vectors of Rickettsia prowazekii, the agent of louse- household or filth-associated arthropods such as houseflies
borne (epidemic) typhus. and cockroaches also can be annoying. Nuisance arthropods
These important discoveries, as well as others of his- are commonly problems for humans at outdoor recreational
toric relevance to medical-veterinary entomology, are areas such as parks, lakes, and beaches.
Introduction Chapter | 1 5

TABLE 1.1 Principal Orders of Insects and Arachnids of Medical-Veterinary Interest

Order Common Names


Class INSECTA
Order Blattaria Cockroaches
Order Phthiraptera Lice
Order Hemiptera True bugs: bedbugs, kissing bugs, assassin bugs
Order Coleoptera Beetles
Order Siphonaptera Fleas
Order Diptera Flies: mosquitoes, black flies, no-see-ums, horse flies, deer flies, sand flies, tsetse
flies, house flies, stable flies, horn flies, bot flies, blow flies, flesh flies, louse flies,
keds, etc.
Order Lepidoptera Moths and butterflies
Order Hymenoptera Wasps, hornets, velvet ants, ants, bees
Class ARACHNIDA
Order Scorpionida Scorpions
Order Solpugida Solpugids, sun spiders, camel spiders, barrel spiders
Order Acari Mites, ticks
Order Araneae Spiders

Toxins and Venoms hemorrhaging, or blisters, the severity of which is largely


dependent on the particular types and amounts of toxins
Several terms are used when discussing chemical sub- involved.
stances that have adverse effects on humans and other an- For further information and details about arthropod
imals. A poison is any substance that when taken into the toxins and venoms, see Chapter 3.
body interferes with normal physiological functions. A
toxin is a poison of plant or animal origin, which can result
in a pathological condition called toxicosis. A venom is a
Allergic Reactions
poisonous mixture of compounds containing one or more A relatively wide spectrum of allergic reactions can occur
toxins, which is produced in venom glands and injected in humans or animals exposed to certain arthropods. Bites
into animal tissues via specialized morphological structures or stings by arthropods such as lice, bedbugs, fleas, bees,
(e.g., stings, modified spines, and chelicerae in arthropods). ants, wasps, mosquitoes, and chiggers can result in allergic
The act of injecting venom into animal tissues is called host reactions. Contact allergies can occur when certain
envenomation. beetles or caterpillars touch the skin. Respiratory allergies
Toxins produced by arthropods represent a wide range can result from inhaling allergenic airborne particles from
of chemical substances, from simple inorganic or organic cockroaches, fleas, dust mites, and other arthropods. The
compounds to complex alkaloids and heterocyclic com- recirculation of air by modern air-handling systems in
pounds. Venoms often contain various pharmacologically buildings tends to exacerbate inhalation of insect allergens.
active compounds that facilitate the spread and effective- For reviews of arthropod allergens, see Arlian (2002) and
ness of the toxic components. The compounds commonly Prester (2012).
include amines (e.g., histamine, catecholamines, serotonin), Humans and other animals usually react to repeated
peptides, polypeptides (e.g., kinins), specific proteins, and exposure to bites or stings from the same or antigenically
enzymes (e.g., phospholipase, hyaluronidase, esterases) related arthropods in one of two ways, depending on the
that vary significantly among different arthropod taxa. nature of the antigen or venom inoculated and the sensi-
Depending on what types of cells or tissues they affect, tivity of the host: (1) desensitization to the bites or stings
toxins and venoms can be characterized as, for example, with repeated exposure and (2) allergic reactions that, in
neurotoxins, cytotoxins, or hemotoxins. Frequently they extreme cases, can develop into life-threatening anaphy-
cause symptoms such as pain, itching, swelling, redness, lactic shock. However, a distinct five-stage sequence of
6 Medical and Veterinary Entomology

reactions typically occurs in most cases in which one is more detail in Chapter 4 on the epidemiology of vector-
repeatedly bitten or stung by the same, or related, species of borne diseases and in the Appendix devoted to
arthropod over time. Stage 1 involves no skin reaction but arboviruses.
leads to the development of hypersensitivity. Stage 2 is a Pathogens are transmitted by arthropods in two basic
delayed-hypersensitivity reaction. Stage 3 is an ways: either biologically or mechanically. In biological
immediate-sensitivity reaction followed by a delayed- transmission, pathogens undergo development or repro-
hypersensitivity reaction. Stage 4 is an immediate reac- duction in the arthropod host. Examples of diseases that
tion only, whereas stage 5 again involves no reaction (i.e., involve biological transmission are malaria, African
the victim becomes desensitized). These changes reflect the trypanosomiasis, Chagas disease, leishmaniasis, and
changing host immune response to prolonged and frequent lymphatic filariasis. In mechanical transmission, patho-
exposure to the same arthropod or to cross-reactive aller- gens are transmitted by arthropods via contaminated ap-
gens or venoms. pendages (usually mouthparts) or regurgitation of an
infectious blood meal. Examples of diseases that involve
mechanical transmission are equine infectious anemia and
Invasion of Host Tissues
myxomatosis. Biological transmission is by far the more
Some arthropods invade the body tissues of their host. common and efficient mechanism for pathogen mainte-
Varying degrees of invasion occur, ranging from subcu- nance and transmission.
taneous infestations to invasion of organs such as the lungs A wide range of life-cycle patterns and degrees of host
and intestines. Invasion of tissues allows arthropods to associations are characterized by arthropod vectors. Some
exploit different host niches and usually involves the ectoparasites, such as sucking lice, remain on their host for
immature stages of parasitic arthropods. life. Others, such as mosquitoes and most biting flies, have
The invasion of host tissues by fly larvae, called a more fleeting association with the host, with some being
myiasis, is the most widespread form of host invasion by associated with the host only during the brief acts of host
arthropods. Larvae of many myiasis-causing flies move location and blood-feeding. Between these two extremes is
extensively through the host tissues. As the larvae mature, a wide range of host associations exhibited by different
they select characteristic host sites (e.g., stomach, throat, arthropod groups.
nasal passages, or various subdermal sites) in which to Literature references on vector-borne diseases, together
complete the parasitic phase of their development. with their epidemiology and ecology, are provided in the
Certain mites also invade the skin or associated hair section “Arthropod-borne Diseases” at the end of this
follicles and dermal glands. Others infest nasal passages, chapter.
lungs and air sacs, cloaca, stomach, intestines, and other
parts of the alimentary tract of their hosts. Examples of
these mites include scabies mites, follicle mites, nasal
FOOD CONTAMINANTS
mites, lung mites, and a variety of other mites that infest Many arthropods can contaminate or spoil food materials.
both domestic and wild birds and mammals. In addition, to causing direct damage to food resources,
arthropods or their parts (e.g., setae, scales, shed cuticles, or
body fragments) may be accidentally ingested. This can
ARTHROPOD-BORNE DISEASES
lead to toxic or allergic reactions, gastrointestinal myiasis,
Table 1.2 lists the principal groups of insects and arachnids and other disorders. At least one case of millipedes
involved in arthropod-borne diseases and the associated (Nopoiulus kochii) infesting human intestines, for several
types of pathogens. Among the wide variety of arthropods years, has been documented (Ertek et al., 2004).
that transmit pathogens to humans and other animals, Insects such as the house fly may alight on food and
mosquitoes are the most important, followed by ticks. Vi- regurgitate pathogen-contaminated fluids before or during
ruses and bacteria (including rickettsiae) are the most feeding. While feeding they also may defecate, contami-
diverse groups of pathogens transmitted by arthropods, nating the food with potential pathogens. Because the
followed by protozoa and filarial nematodes. A standard- alimentary tract of arthropods may harbor pathogenic mi-
ized nomenclature has been proposed for parasitic diseases croorganisms, subsequent consumption of the contaminated
of animals including those with arthropod vectors (Kassai food can lead to the transmission of these pathogens to
et al., 1988). humans or other animals. Similarly, the integument of
All of the viruses listed in Table 1.2 are arthropod- household pests such as flies and cockroaches (particularly
borne viruses, usually referred to as arboviruses, which their legs and tarsi) can serve as a contact source of path-
indicates that they are typically transmitted by insects or ogens that may be readily transferred to food items. Some
other arthropod hosts. The study of arboviruses is termed of these arthropods previously may have visited fecal
arbovirology. These and related terms are discussed in matter, garbage and other decomposing matter, animal
Another random document with
no related content on Scribd:
attack on all the contemporary elements of Russian society
that were antagonistic to the Western reform.
Specimens from several of Kantemír’s satires are given in
C. E. Turner’s Studies in Russian Literature, London, 1882,
and the same article, in Fraser’s Magazine, 1877.
Parts of the First Satire, in article on Russian Literature, in
Foreign Quarterly Review, vol. i.

TO MY MIND

Immature Mind, fruit of recent study! Be quiet, urge not the pen
into my hands: even without writing one may pass the fleeting days
of life and gain honours, though one be not a poet. Many easy paths
lead in our days to honours, and bold feet need not stumble upon
them: the least acceptable is the one the nine barefooted sisters
have laid out. Many a man has lost his strength thereon, without
reaching a goal. You have to toil and moil there, and while you
labour, people avoid you like the plague, rail at you, loathe you. He
who bends over the table, fixing his eyes upon books, will gain no
magnificent palaces, nor gardens adorned with marbles; will add no
sheep to his paternal flock.
’Tis true, in our young monarch[127] a mighty hope has risen for
the Muses, and the ignorant flee in shame from him. Apollo has
found in him a strong defender of his glory, and has seen him
honouring his suite and steadily intent upon increasing the dwellers
on Parnassus.[128] The trouble is, many loudly praise in the Tsar
what in the subject they haughtily condemn.
“Schisms and heresies are begot by science.[129] He lies most
who knows most; who pores over books becomes an atheist.” Thus
Crito grumbles, his rosary in his hands, and sighs, and with bitter
tears the saintly soul bids us see how dangerous is the seed of
learning that is cast among us: our children, who heretofore gently
and meekly walked in the path of their forefathers, eagerly attending
divine service and listening in fear to what they did not understand,
now, to the horror of the Church, have begun to read the Bible; they
discuss all, want to know the cause of all, and put little faith in the
clerical profession; they have lost their good habits, have forgotten
how to drink kvas, and will not be driven with a stick to partake of salt
meat. They place no candles before the images, observe no feasts.
They regard the worldly power misplaced in clerical hands, and
whisper that worldly possessions ill become those who have
renounced a worldly life.
Sylvan finds another fault with science: “Education,” he says,
“brings famine in its track. We managed to get along before this
without knowing Latin much better than we live now. We used to
harvest more grain in our ignorance, but now that we have learned a
foreign language, we lose our corn. What of it if my argument be
weak and without sense and connection,—what matters that to a
nobleman? Proof, order of words, is the affair of low-born men; for
aristocrats it suffices boldly to assent, or contradict. Insane is he who
examines the force and limitations of his soul; who toils whole days
in his sweat, in order to learn the structure of the world and the
change or cause of things: ’tis like making pease to stick to the wall.
Will all that add one day to my life, or one penny to my coffers? Can I
by means of it find out how much my clerk and superintendent steal
a year or how to add water to my pond, or to increase the number of
barrels in my still?
“Nor is he wise who, full of unrest, dims his eyes over a smoking
fire, in order to learn the properties of ores. We have passed our A B
C, and we can tell without all that the difference between gold, silver
and copper. The science of herbs and diseases is idle talk. You have
a headache, and the physician looks for signs of it in your hand! The
blood is the cause of all, if we are to put faith in them. When we feel
weak, it is because our blood flows too slowly; if it moves fast, there
is a fever, he says boldly, though no one has ever seen the inside of
a living body. And while he passes his time in such fables, the
contents of our money-bags go into his. Of what use is it to calculate
the course of the stars, and without rhyme or reason pass sleepless
nights, gazing at one spot: for mere curiosity’s sake to lose your rest,
trying to ascertain whether the sun moves, or we with the earth? We
can read in the almanac, for every day in the year, the date of the
month and the hour of sunrise. We can manage to divide the land in
quarters without Euclid, and we know without algebra how many
kopeks there are in a rouble.” Sylvan praises but one science to the
skies,—the one that teaches how to increase his income and to save
expenses. To labour in that from which your pocket does not swell at
once, he deems a very dangerous occupation for a citizen.
Red-faced Lucas, belching thrice, speaks in a chanting voice:
“Study kills the companionship of men. We have been created by
God as social beings, and we have been given intelligence not for
our own sakes alone. What good does it do anybody, if I shut myself
up in my cabinet, and for my dead friends lose the living—when all
my comradeship, all my good fellows, will be ink, pen, sand and
paper? In merriment, in banquets we must pass our lives. Life is
short, why should we curtail it further, worry over books, and harm
our eyes? Is it not better to pass your days and nights over the
winecup? Wine is a divine gift, there is much good in it: it befriends
people, gives cause for conversation, makes glad, dispels heavy
thoughts, eases misery, gives courage to the weak, mollifies the
cruel, checks sullenness, and leads the lover more readily to his
goal. When they will begin to make furrows in the sky, and the stars
will shine through the surface of the earth; when swift rivers will run
to their sources, and past ages will return; when at Lent the monk
will eat nothing but dried sturgeon, then will I abandon my cup and
take to books.”
Medor is worried because too much paper is used for letters and
for printed books, and because he will soon be left without paper to
curl his locks with. He would not change for Seneca a pound of good
face-powder; in comparison with Egór,[130] Vergil is not worth two
farthings to him, and he showers his praises on Rex,[131] not Cicero.
This is a part of the speeches that daily ring in my ears, and for
this, O Mind, I advise you to be dumber than a dumpling. Where
there is no profit, praise encourages to work, and without it the heart
grows faint. But it is much worse, when instead of praises you earn
insults! It is harder than for a tippler not to get his wine, or for a priest
not to celebrate on Holy Week, or for a merchant to forego heady
liquor.
I know, O Mind, that you will boldly answer me that it is not easy
for an evil-minded man to praise virtue; that the dandy, miser,
hypocrite, and the like, must perforce scorn science, and that their
malevolent discourse concerns no men of culture.
Your judgment is excellent, correct; and thus it ought to be, but in
our days the words of the ill-disposed control the wise. Besides, the
sciences have other ill-wishers than those whom, for shortness’
sake, I merely mentioned or, to tell the truth, dared to mention. There
are many more. The holy keepers of the keys of heaven and those to
whom Themis has entrusted the golden scales little love, nearly all of
them, the true adornment of the mind.
You want to be an archbishop? Don a surplice, above it let a
gorgeous chasuble adorn your body, put a golden chain[132] around
your neck, cover your head with a high hat, your belly with a beard,
order the crosier to be carried in pomp before you; place yourself
comfortably in your carriage and, as your heart bursts with anger,
cast your benedictions to the right and left. By these signs you will
easily be recognised as the archpriest, and they will reverently call
you “Father.” But science? What has the Church to gain from it?
Some priest might forget a part, if he wrote out his sermon, and thus
there would be a loss of the Church’s revenues, and these are the
Church’s main privileges and greatest glory.
Do you wish to become a judge? Don a wig full of locks, scold him
who comes with a complaint but with empty hands, let your heart
firmly ignore the tears of the poor, and sleep in your arm-chair when
the clerk reads the brief. When someone mentions to you the civil
code, or the law of nature, or the people’s rights, spit in his face; say
that he lies at random and tries to impose an intolerable burden on
the judges; that it is the clerk’s business to rummage through
mountains of documents, but that it suffices for a judge to announce
his sentence.
The time has not come down to us when Wisdom presided over
everything and distributed wreaths, and was the only means for
advancement. The golden age has not come down to our
generation. Pride, indolence, wealth, have vanquished wisdom;
ignorance has taken the place of wisdom: it glorifies itself under the
mitre, walks in embroidered gowns, sits in judgment behind the red
cloth, boldly leads armies. Science trudges along in rags and
patches, and is driven from nearly all houses with contumely; they do
not want to know her and evade her friendship, just as those who
have suffered upon the sea avoid service on a ship. All cry: “We see
no good in science; the heads of learned men are full, but their
hands are empty.”
If one knows how to shuffle cards, to tell the flavours of various
wines, can dance, plays three pieces on the flute, cleverly matches
the colours in his apparel, for him, even in his tender years, all high
honours are but a small reward, and he regards himself to be the
equal of the Seven Sages.
“There is no justice in the world!” cries the brainless subdeacon.
“They have not yet made me a bishop, though I read fluently the
Book of the Hours,[133] the Psalter and the Epistles, and even
Chrysostom without stumbling, although I do not understand him.”
The warrior grumbles because he has not yet charge of his
regiment, though he knows how to sign his name. The scribe is
angry because he is not yet seated behind the red cloth, though he
is able to make a copy in a clear hand. He thinks it an insult to grow
old in obscurity, though he counts seven boyárs in his family and is
possessed of two thousand village houses, even though he can
neither read nor write.
Hearing such words, and seeing such examples, be silent, Mind,
complain not of your obscurity. His life has no terrors, though he may
deem it hard, who silently retires to his quiet nook. If gracious
Wisdom has taught you anything, rejoice in secret, meditating by
yourself over the advantages of learning. Explain it not to others,
lest, instead of praises which you expect, you be roundly scolded.

FOOTNOTES:
[127] Peter II., born 1715; ascended the throne in 1729, the
year the satire was written in.
[128] Immediately upon arriving in Moscow, Peter II. confirmed
the privileges of the Academy of Sciences.
[129] Compare Feofán Prokopóvich’s Spiritual Reglement, p.
212.
[130] A famous shoemaker in Moscow; died in 1729.
[131] A German tailor of Moscow.
[132] With the image of the Holy Virgin or the Saviour,—the so-
called panagia.
[133] Prayer-book containing the prayers for every hour; it was
commonly used as a text-book for reading.
Vasíli Kiríllovich Tredyakóvski. (1703-1769.)
Like Lomonósov, Tredyakóvski was of humble origin, his
father having been a priest in the city of Astrakhán; also, like
his more illustrious colleague a few years later, he walked to
Moscow and there entered the School of the Redeemer. He
later passed a few years abroad, where he became
acquainted with French literature. Upon his return to St.
Petersburg in 1730, he translated a French book; in this
translation the spoken Russian is for the first time used, free
from Slavic influence. Even before this, Tredyakóvski had
written verses in the syllabic versification, but in 1735 he
discovered that the tonic versification was the only one
adapted to the Russian language, and at once set out to write
in that measure. His chief deserts do not lie in poetry, for his
verses show an absolute absence of talent, and he later
became a byword for insipidity. He was the first man to point
out the necessity of using the Russian language for literary
purposes, and to indicate the line in which Russian poetry
must develop. By his enormous industry in translating from
foreign languages he became an important factor in the
dissemination of learning. The following ode is really an
imitation of Boileau’s Sur la prise de Namur.

ODE ON THE SURRENDER OF DANTZIG

What strange intoxication emboldens my voice to singing? Muses,


dwellers of Parnassus, does not my mind perceive you? I hear your
sweet-sounding strings, your beautiful measure and moods, and a
fire arises in my thoughts. O nations, listen all! Stormy winds, do not
blow: my verse sings of Anna.
Pindar, and after him Flaccus, have in high-flowing diction risen
from the mist to the bright stars, like swift eagles. But if my song to-
day were to equal my sincere and eternal zeal for Anna, Orpheus of
Thrace and Amphion of Thebes would be in ecstasy from it.
Now I strike the dulcet lyre to celebrate the magnificent victory to
the greater downfall of the enemy. Oh, what victorious might has
adorned our joy, for the might of the adversary was equal to ours.
There is no limit to our pure joy that surpasses all example, that has
given balm to our hearts.
Has Neptune himself built the walls, those that stand by the sea?
Do they not resemble the Trojan walls, for they would not let in the
innumerable Russian army, mightily opposing it? Do not all call the
Vistula Skamander? Do they not all regard Stoltzenberg as Mount
Ida?
That is not Troy, the mother of fables: there is not one Achilles
here; everyone of the rank and file is in bravery a Hercules. What
might is that that hurls lightning? Is it not Minerva gleaming in her
helmet? ’Tis evident from her looks, from her whole appearance, that
she is a goddess: without her ægis she is terrible,—’tis Anna, chief of
all empresses.
That also is a Russian army that has closely invested Dantzig, the
city of the foe. Each warrior, hastening to the battle, it behooves to
call a Mars. Each is ready boldly to shed his blood, or to crown the
undertaking for Anna’s sake. Each one is strong with Anna’s fortune:
Anna is their strong hope; and, knowing that Anna is gracious to
them, they are faithful and not undecided.
Golden beam of the European and Asian Sun! O Russian
monarch, the key to your happiness is the kindness to your subjects
and your benign rule! The whole world honours your name, and the
universe will not hold your glory seeing that, O beautiful flower of
virtues!
What do I see? Does not my eye deceive me? A youth has
opposed himself to Hercules, lifting high his brows behind ramparts,
beyond the river! ’Tis Dantzig, having taken foolish counsel, as if
drunk with heady wine, that dares to oppose the great autocrat! In its
blindness it does not see the abysses, nor all death-bearing valleys.
It receives Stanislaus in its midst, who seeks twice a crown, and
hopes to be defended to the end through nearby Neptune: fearing
the Russian thunder it invokes the aid of a distant people from the
banks of the Seine: but they beat the drums at the waters of
Wechselmünde for a retreat.
Dantzig is proud of its fire and steel, and its regiments of soldiers,
and directs its engines of war against the Russians on the hills.
Being rich in stores it calls to Stanislaus; it in vain implores its
soldiers that have no brave hearts, but think only of this, how to save
their lives, and run.
O Dantzig, oh! What are you daring! Come to your senses, collect
yourself, for you are hurling yourself to destruction. Why have you
stopped? You are hesitating! Surrender! Wherefore have you such
boldness and do not tremble before Anna? Many tribes of their own
free will and without strife submit to her: China bows down before
her twice, in order not to pay her tribute.
Nowhere has there been the like of Anna in kindness, nor is there
anywhere in the world one so able to wage war with the unyielding.
Her sword wound with the olive branch is only ominous in war.
Abandon, Dantzig, your evil purpose: you see, the Alcidæ are ready
with cruel miseries for your inhabitants. You hear Anna’s angry voice:
save yourself!
You are closely pressed by thousands of athletes; you are mightily
struck by the flash of angry lightnings. You cannot withstand: the
thunder is ready not in jest. Your ramparts are without defence; the
earth opens up abysses; roofs fly into the air; your walls are emptied
of men.
If all the powers combined were to aid you, O Dantzig; if the
elements defended you; if from all the ends of the world soldiers
came to spill their blood for you,—yet nothing would be able to save
you from suffering and to stop your misery, and wring you out of
Anna’s hands.
Your adversaries see to-day the bravery of Russian soldiers:
neither fire nor water harms them, and they advance with open
breasts. How readily they advance! How forgetful they are of their
lives! The cannon’s thunder frightens them not! They make the
assault, as if going to a wedding feast! Only through smoky darkness
one may see that their brows are facing the forts.
Within the walls of the wretched city all are struck down with fear:
everything falls and flies to dust,—the besiegers are on the walls!
The last magistrates, seeing from their tower their vain hope in the
distant armies and Stanislaus who had taken refuge within their
walls, besides themselves, exclaim: “We are fated to fall!”
What I have prophesied is about to happen,—Dantzig begins to
tremble: all think of surrendering, as before they all decided to fight,
and of saving themselves from the engines of war, from flying bombs
and from all the pests the city is oppressed by. All cry, for the burden
was too heavy to carry, “It is time now to open the gates to Anna’s
army.”
So it is done: the sign for surrender is given, and Dantzig is at our
feet! Our soldiers are happy in their success; the fires have gone out;
there is an end to misery. Immediately Glory took its flight and
announced with its thundering trumpet: “Anna is fortunate! Anna is
unconquerable; Anna, exalted by all, is their common glory and
honour.”
Lyre! abate your song: it is not possible for me properly to praise
diadem-bearing Anna and her great goodness, any more than I can
fly. It is Anna’s good fortune that she is loved by God. He always
watches over her, and through Him she is victorious. Who would
dare to oppose her? May Anna live many years!
Princess Natálya Borísovna Dolgorúki. (1714-
1771.)
The Princess Dolgorúki was the daughter of Count
Sheremétev, who was intimately connected with the reforms
of Peter the Great. In 1729 she was betrothed to Prince Iván
Aleksyéevich Dolgorúki, the favourite of Peter II.; Feofán
Prokopóvich performed the ceremony of the betrothal, and
the whole Imperial family and the most distinguished people
of the capital were present. A few days later Peter II. died,
and Anna Ioánnovna ascended the throne. Dolgorúki was
banished to Siberia, and she married him in order to follow
him into exile. They passed eight years in the Government of
Tobólsk, when her husband was taken to Nóvgorod and
executed. For three years she remained in ignorance of his
fate, when the Empress Elizabeth permitted her to return to
St. Petersburg. In 1758 Princess Dolgorúki entered a
monastery at Kíev, and ten years later she wrote her
Memoirs, at the request of her son Michael. In 1810 her
grandson, the poet Dolgorúki (see p. 422), had these
Memoirs printed. The Princess Dolgorúki has become a
synonym for a devoted Russian woman, and she has
frequently been celebrated in poetry, especially by Rylyéev,
Kozlóv and Nekrásov. There is also an English book treating
of her life: The Life and Times of Nathalia Borissovna,
Princess Dolgorookov, by J. A. Heard, London, 1857.

FROM HER “MEMOIRS”

My mind totters when I recall all that has befallen me after my


happiness which at that time appeared to me to be eternal. I did not
have a friend to teach me that I ought to walk more warily on the
slippery road of pleasure. My Lord! What a threatening storm arose
against me, and what calamities from the whole world befell me!
Lord! Give me strength to tell of my sufferings, that I may describe
them for the information of the curious and the consolation of the
afflicted who, thinking of me, might be consoled. I have passed all
the days of my life in misery, and have experienced all: persecution,
exile, want, separation from my beloved one,—everything that one
can think of. I do not boast of my endurance, but will boast of the
mercy of the Lord who has given me so much strength to bear all
that I have borne up to now. It would be impossible for a man to
endure such strokes, if the power of the Lord did not strengthen him
from on high. Consider my bringing up, and my present state!
Here is the beginning of my misery that I had never expected. Our
Emperor had departed from this life, and before I had expected it,
there was a change of the crown. It evidently had pleased God to
chastise the people for their sins: a merciful Tsar was taken away
from them, and great was the weeping in the nation. All my relatives
came together, were sorrowing and weeping, and wondering how to
announce to me the calamity. I generally slept late, until nine o’clock;
as soon as I awoke, I noticed that the eyes of all were in tears;
though they were careful to hide it, yet it was quite obvious they had
been weeping. I knew that the Tsar was sick, and even very sick, but
I had great hope the Lord would not abandon His orphans. They
were of necessity compelled to tell me the truth. As soon as this
news reached my ears, I lost my consciousness; when I regained it, I
kept on repeating: “I am lost, lost!” No other words left my lips but
“lost.” However they tried to console me, they could not stop my
weeping, nor keep me quiet. I knew too well the custom of my
country, that all the favourites perish with the death of their
Emperors: what could I, then, expect? Yet, I did not think that the
end would be as bad as it actually was, for though my fiancé was
beloved by the Tsar, and had many distinctions, and all kinds of
affairs of State had been entrusted to him, yet I placed some hope in
his honest acts. Knowing his innocence, and that he had not been
tainted by any improper conduct, it appeared to me that a man would
not be accused without a proper judicial trial, or be subject to
disfavour, and be deprived of his honours and possessions; I learned
only later that truth is not helpful in misfortune.
So I wept unconsolably. My relatives, in their search for means of
consoling me, pointed out to me that I was yet a young person, and
had no reason to grieve so senselessly; that I could reject my fiancé
if things went badly with him, and that there were other suitors who
were not of less worth than he, even if they had not his high honours.
And indeed there was a suitor who was very anxious to have me, but
I did not like him, though all my relatives wanted me to marry him.
That proposition weighed so heavily upon me, that I was not able to
answer them. Consider yourself, what kind of a consolation that
could be to me, and how dishonourable such an act would have
been,—to be ready to marry him when he was great, but to refuse
him the moment he was cast into misfortune. I could not agree to
any such unscrupulous advice; I resolved at once to live and die
together with him to whom I had given my heart, and not to allow
anyone else to share my love. It was not my habit to love one to-day
and another to-morrow; such is the fashion in the world, but I proved
to the world that I was faithful in love. I have been a companion to
my husband in all his troubles, and I am telling the truth when I
assert that in all my misery I never repented having married him, and
did not murmur against the Lord for it. He is my witness: I bore
everything while loving him, and as much as was in my power, I kept
up his courage. My relatives were evidently of a different opinion,
and therefore advised me otherwise, or maybe they simply pitied me.
Towards evening my fiancé came to my house, and complained to
me of his misfortune. He told me of the pitiable death of the Emperor,
who did not lose consciousness to the last, and bid him good-bye.
While he told me all this, we both wept, and swore to each other that
nothing should separate us but death; I was ready to go with him
through all the terrestrial misfortunes. Thus it grew worse from hour
to hour. Where were those who formerly had sought our protection
and friendship? They had all hid themselves, and my relatives stood
aloof from me; they all left me for the new favourites, and all were
afraid to meet me, lest they should suffer through the suspicion
under which I was. It were better for a person not to be born in this
world, if he is to be great for a while, and then will fall into disgrace:
all will soon despise him, and no one will speak to him.
Here we remained about a week, while a vessel was being fitted
out to take us down the river. All that was terrible to me, and I ought
to pass it in silence. My governess, to whose care I had been
entrusted by my mother, did not wish to leave me, and had come
with me to the village. She thought that we would pass all the days of
our misfortune there; but things turned out differently, and she was
compelled to leave me. She was a foreigner, and could not endure
all the hardships; yet, as much as she could she did for me in those
days: went on the ill-starred vessel that was to take us away, fixed
everything there, hung the walls with tapestry to keep out the
dampness, that I might not catch a cold; she placed a pavilion on
board, partitioned off a room, in which we were to live, and wept for
me all the time.
At last there arrived the bitter day when we must depart. We were
given ten people to attend on us, and a woman for each person, in
all, five. I had intended to take my maid with me, but my sisters-in-
law dissuaded me: they gave me theirs to take her place, and gave
me another maid for an assistant to the laundresses, who could do
nothing else but wash clothes; I was compelled to agree to their
arrangement.
My maid wept, and did not want to part from me. I asked her not to
importune me with her tears, and to take things as fate had decreed.
Such was my equipment: I had not even my own serf, and not a
penny of money. My governess gave me every kopek she had; it
was not a great sum, only sixty roubles, and with that I departed. I do
not remember whether we went on foot to the vessel, or whether we
drove to it in a carriage. The river was not far from our house; there I
bid good-bye to my family, for they had been permitted to see us off.
I stepped into the cabin, and saw how it was fixed up: my
governess had done all she could to help me in my evil plight. I had
to thank her here for the love she had shown to me, and for the
education she had given me; I also bid her farewell, not expecting
ever to see her again: we grasped each other’s necks, and my
hands grew stiff with cold, and I do not remember how we were torn
from each other. I regained consciousness in the place that served
as a cabin. I was lying in the bed, and my husband was standing
over me, holding me by my hand, and making me smell some salts. I
jumped down from my bed, ran upstairs, thinking that I would still
catch a glimpse of it all, but those were all unfamiliar scenes,—we
had sailed away a long distance. Then I noticed that I had lost a
pearl that I wore on my finger; I evidently dropped it in the water as I
bade my family farewell; I was not even sorry for it,—other thoughts
were occupying me: life was lost, and I was left alone, had lost all for
the sake of one man. And thus we sailed all night long.
The next day there was a stiff breeze; there was a storm on the
river, and the thunder sounded more terrible on the water than on
land, and I am naturally very much afraid of thunder. The vessel
rolled from side to side, and every time it thundered people fell down.
My younger sister-in-law was very much frightened, and wept and
cried aloud. I thought the world had come to an end; we were
compelled to make for the shore, where we passed a sleepless night
in terror. As soon as it dawned, the storm subsided; we continued
our voyage, which lasted three weeks. Whenever the weather was
quiet, I sat near the window in the cabin; I wept or washed my
kerchiefs, while the water was nearby. At times I bought a sturgeon,
and, tying him to a rope, let him swim by my side, so that I was not
the only captive, but the sturgeon with me. Whenever the wind
began to rock the boat, my head began to ache, and I felt
nauseated; then they took me out on deck, where I lay unconscious
until the wind subsided, being covered with a fur coat: on the water
the winds are piercing. Often he sat by my side, to keep me
company. When the storm was over, I rested; but I could not eat
much from nausea.
Here is what once happened to us: There was a frightful storm,
and there was not a person on board who knew where there were
the deep places and the shallows, or where we could land. The
sailors were merely peasants that had been taken from the plough,
and who were sailing where the wind bore them. It was getting dark,
the night was near, and the wind did not permit us to make a landing.
They threw out an anchor in the middle of the stream, where it was
deepest, and the anchor was carried away. The companion of my
misfortunes would not let me go on deck, for he was afraid that I
would be crushed in the turmoil. The people were running all about
the boat: some were pumping out the water, others were tying up the
anchor; all were at work. While nothing was being done successfully,
the boat was suddenly drawn into an eddy. I heard a terrible noise,
and did not know what had happened. I arose to look out: our boat
was standing as if in a box, between two shores. I asked where we
were, but nobody could tell me, for they did not know themselves.
On one shore there was nothing but a birch wood, but it was not a
very thick forest. The earth on that shore began to settle, and the
forest slid several fathoms into the river, or eddy, where we were
standing. The forest rustled terribly under our very boat, and then we
were lifted up, and again drawn into the eddy. Thus it lasted for a
long time. All thought that we would perish, and the sailors were
ready to save their lives in boats, and to leave us to death. Finally, so
much of the land was torn loose that only a small strip was left, and
beyond it we could see some water, supposedly a lake. If that strip
were carried away, we would be in that lake. The wind was awful,
and our end would certainly have come, if God’s mercy had not
saved us. The wind calmed down, and no more land was being
carried away, and we were saved; at daylight we rode out of the
eddy into the river, and continued our voyage. That eddy had carried
part of my life away; yet I endured it all, all the terrors, for the end of
my sufferings was not yet to be: I was preparing myself for greater
woes, and God gave me strength for them.

We reached the provincial town of the island where we were to


reside. We were told that the way to that island was by water, and
that a change would be made here: the officer of the guard was to
return, and we were to be turned over to an officer of the local
garrison, with a detachment of twenty-four soldiers. We stayed here
a week, while they were fixing the boat that was to take us there, and
we were transferred from hand to hand, like prisoners. It was such a
pitiable sight that even a heart of stone would be softened. At this
departure, the officer wept, and said: “Now you will suffer all kinds of
insult. These are not ordinary men: they will treat you like common
people, and will show you no indulgence.” We all wept, as if we were
parting from a relative. We had at least gotten used to him. However
badly we were off, yet he had known us in our fortune, and he felt
ashamed to treat us harshly.
When they had fixed the boat, a new commander took us to it. It
was quite a procession. A crowd of soldiers followed us, as if we
were robbers. I walked with downcast eyes, and did not look around:
there was a great number of curious people along the road on which
they led us. We arrived at the boat. I was frightened when I saw it,
for it was quite different from the former one: out of disrespect to us,
they gave us a worthless one. The boat was in accordance with the
designation which we bore, and they did not care, if we were to
perish the next day: we were simply prisoners,—there was no other
name for us. Oh, what can there be worse than that appellation? The
honour we received was in conformity with it! The boards on the boat
were all warped, and you could see daylight through them; the
moment a breeze began to blow, it creaked. It was black with age
and soot: labourers had been making fires in it, and no one would
have thought to travel in it. It had been abandoned, and was
intended for kindling wood. As they were in a hurry with us, they did
not dare keep us back long, and gave us the first boat they could
find. But maybe they had express orders to drown us. God having
willed otherwise, we arrived safely at the appointed place.
We were compelled to obey a new commander. We tried all
means to gain his favour, but in vain. How could we have found any
means? God grant us to suffer with a clever man! But he was a
stupid officer. He had risen from a common peasant to be a captain.
He thought he was a great man, and that we must be kept as
severely as possible, since we were criminals. He regarded it below
his dignity to speak to us; yet in spite of all his arrogance, he came to
dine with us. Consider for yourself whether the man had any sense
from the way he was dressed: he wore his uniform right over his
shirt, and slippers on his bare feet; and thus he sat down to dinner
with us! I was younger than the rest, and uncontrollable: I could not
help laughing as I looked at his ridiculous get-up. He noticed that I
was laughing at him, and said, himself smiling: “Lucky for you that
my books have burnt, or I should have a talk with you!” However
bitter I felt, I tried to get him to talk more; but he never uttered
another word. Just think what a commander we were given to watch
us in all we did! What were they afraid of? That we would run away?
Not their watch kept us back, but our innocence: we were sure that
in time they would see their error, and would return us to our former
possessions. Besides, we were restrained by the fact that we had a
large family. And thus we sailed with the stupid commander a whole
month until we arrived at the town where we were to reside.
Mikhaíl Vasílevich Lomonósov. (1711-1765.)
Lomonósov was born in the village of Denísovka, in the
Government of Arkhángelsk, not far from the spot where, one
hundred and fifty years before, the English had rediscovered
Russia. In his letters to Shuválov, Lomonósov tells us of the
difficulties with which he had to contend at home and at the
School of the Redeemer at Moscow. His brilliant progress
caused him to be chosen among the first men to be sent
abroad at Government expense to study mining, and to get
acquainted with mining methods in Holland, England and
France. In spite of insufficient support from the Government
and a roving life at German universities, Lomonósov made
excellent progress in philosophy, under Christian Wolff at
Marburg, and in the sciences at Freiburg. After marrying a
German woman, wandering about and starving, Lomonósov
returned to St. Petersburg. Before reaching home, he had
sent to St. Petersburg his Ode on the Occasion of the
Capture of Khotín. It was the first time the tonic versification
was successfully applied to the language, and though the
diction of the ode is turgid and the enthusiasm forced, yet it
became the model for a vast family of odes and eulogies,
generally written to order, until Derzhávin introduced a new
style with his Felítsa.
Upon his return, Lomonósov became attached to the
University, which was mainly filled with German professors.
His own unamiable temper, combined with the not more
amiable characters of German colleagues, was the cause of
endless quarrels and exasperations. Under the most
depressing difficulties, Lomonósov, the first learned Russian,
developed a prodigious activity. He perfected the Russian
literary language, lectured on rhetoric and the sciences and
wrote text-books, odes and dramas. For a century he passed
in Russia as a great poet, and his deserts in other directions
were disregarded. But a more sober criticism sees now in
Lomonósov a great scientist who has increased knowledge by
several discoveries, and only a second-rate poet. Only where
he described phenomena of nature or scientific facts, did he
become really inspired, and write poems that have survived
him. His services to the Russian language and literature are
many. He did for them what Peter the Great did for the State:
by his own mighty personality and example be put them on
the road which they have never abandoned, and though
lacking originality, the school of Lomonósov itself survived in
Russian literature to the end of the eighteenth century.
But few of Lomonósov’s poems have been translated into
English. Ode from Job, Morning Meditations, Evening
Meditations, are given in Sir John Bowring’s Specimens of the
Russian Poets, Part II.; the Evening Meditations, in another
version, is also given by him in Part I.
Ode in Honour of the Empress Anna, in F. R. Grahame’s
The Progress of Science, Art and Literature in Russia.
Morning Meditation, and part of the Ode on the Accession
of Catherine II., in C. E. Turner’s Studies in Russian
Literature, and, the same article, in Fraser’s Magazine, 1877.
A Chronological Abridgement of Russian History; translated
from the original Russian ... and continued to the present by
the translator (J. G. A. F.), London, 1767.

LETTERS TO I. I. SHUVÁLOV[134]

Dear Sir, Iván Ivánovich:—Your Excellency’s kind consideration in


honouring me with a letter assures me, to my great joy, of your
unchanged feelings to me, and this I have for many years regarded
as one of my great fortunes. How could the august generosity of our
incomparable Empress, which I enjoy through your fatherly
intercession, divert me from my love and zeal to the sciences, when
extreme poverty, which I have endured voluntarily for the sake of

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