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Encyclopedia of Entomology

Encyclopedia of Entomology

Edited by
John L. Capinera
University of Florida

Second Edition

Volume 4
Professor John L. Capinera
Dept. Entomology and Nematology
University of Florida
Gainesville FL 326110620

Library of Congress Control Number: 2008930112

ISBN: 978-1-4020-6242-1
This publication is available also as:
Electronic publication under ISBN 978-1-4020-6359-6 and
Print and electronic bundle under ISBN 978-1-4020-6360-2

This work is subject to copyright. All rights are reserved, whether the whole or part of the material is con-
cerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, repro-
duction on microfilms or in other ways, and storage in data banks. Duplication of this publication or parts
thereof is only permitted under the provisions of the German Copyright Law of September 9, 1965, in its cur-
rent version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for
prosecution under the German Copyright Law.

2008 Springer Science+Business Media B.V.

The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of a spe-
cific statement, that such names are exempt from the relevant protective laws and regulations and therefore
free for general use.

Editor: Zuzana Bernhart, Dordrecht/ Sandra Fabiani, Heidelberg

Development Editor: Sylvia Blago, Heidelberg
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Editorial Board
Cyrus Abivardi Eugene J. Gerberg
Swiss Federal Institute of Technnoloy University of Florida
Donald R. Barnard Donald W. Hall
United States Department of Agriculture University of Florida
Jean-Luc Boev Marjorie A. Hoy
Royal Belgian Institute of Natural Sciences University of Florida
Drion Boucias John B. Heppner
University of Florida Florida State Collection of Arthropods
Paul. M. Choate Pauline O. Lawrence
University of Florida University of Florida
Whitney Cranshaw Heather J. McAuslane
Colorado State University University of Florida
Thomas C. Emmel James L. Nation
University of Florida University of Florida
J. Howard Frank Herb Oberlander
University of Florida United States Department of Agriculture
Severiano F. Gayubo Frank B. Peairs
Universidad de Salamanca Colorado State University
This project is the labor of many people, including some who labored diligently behind the scenes.
Among those to whom I am greatly indebted for their behind-the-scenes assistance are Pam Howell and
Carole Girimont (first edition) and Pam Howell (second edition) for document processing and editing
assistance; Mike Sanford, Pat Hope, and Jane Medley (first edition) and Hope Johnson (second edition)
for assistance with the images, and Marsha Capinera for compiling the list of contributors. Ron Cave,
Andrei Sourakov, and Lyle Buss helped greatly by supplying numerous photographs for the second
Howard Frank deserves special mention for his editing acumen and assistance. Drion Boucias con-
tributed the lengthy unattributed sections on insect pathology. The unattributed biographic sketches
with last names beginning with A to J were contributed by Howard Frank. All other unattributed
sections were contributed by John Capinera.
Some biologists have called this the Age of Insects. Among animals, certainly the diversity of insects is
unrivaled. Nearly one million species have been described to date, and some entomologists estimate that
as the tropics are fully explored, we will find that there are actually more than three million insect spe-
cies. The large number of insects is often attributed to the divergence of plants (angiosperms), which
provide numerous hosts and places to feed, but if plant feeders are excluded from the tabulation the
biodiversity of insects remains unrivaled. Virtually every environment has been exploited by these resil-
ient organisms. Even if one dislikes insects, they are impossible to ignore, and a little knowledge about
them could be indispensable should one have a close encounter of an unpleasant kind.
Insects are remarkable biological organisms. They are small enough to escape the detailed scrutiny
of most people, but I have yet to meet anyone whom, once provided the opportunity to examine insects
closely (through a microscope) is not completely amazed by the detail and complexity of these exqui-
sitely designed (by natural selection) beasties. They are fascinating in function as well as form. Insects are
the only invertebrates to fly, they are disproportionately strong, and their ecological adaptability defies
belief. For example, some insects produce their own version of anti-freeze, which allows them to be fro-
zen solid yet to regain normal function upon thawing. Their sensory abilities are beyond human com-
prehension; a male insect can sometimes locate a female by her perfume (pheromone) from several
kilometers distance. Although not normally considered intelligent, insects display surprisingly complex
behaviors, and altruistic social systems that could well serve as models for human societies.
Insects and their close relatives are important for many reasons besides their sheer diversity. Their
effect on humans is profound. Insects are our chief competitor for food and fiber resources throughout
the world. Annual crop losses of 10 to 15% are attributed to insects, with both pre-harvest and post-
harvest losses considerably more at times. Insects also are the principal vector of many human, animal,
and plant diseases, including viruses, mollicutes, bacteria, fungi, and nematodes. The ability to transmit
diseases magnifies their effect, and makes it more difficult to manage injury. Over the course of human
history, insect-transmitted disease has caused untold human suffering. For example, introduction of
flea-transmitted bubonic plague to Europe centuries ago killed millions of people and caused severe
disruption to western civilization. Though less dramatic, mosquito-transmitted malaria kills thousands
annually throughout the world, and unlike plague, which is now mostly a historical footnote, the toll
continues to mount.
Advances in technology, particularly the introduction of chemical insecticides, have done much to
remove the threat of insect-related damage from the consciousness of most humans. Insecticides are
applied preventatively to avoid pre- and post-harvest damage to crops, to our dwellings, and to our land-
scape. This is an oft-overlooked but remarkable achievement that has increased stability in the supply
and price of resources, and in the lives of resource producers. No longer are people faced with starvation
or economic ruin due to the ravages of insects; in almost all parts of the world, the ready availability of
insecticides can be used to prevent massive insect population outbreaks. However, we realize increas-
ingly that this approach is not without its own set of health, environmental and economic costs, and
alleviating dependency on insecticides, or making alternatives to insecticides more readily available, has
assumed greater priority.

We are faced with an interesting dichotomy. There is a wealth of information about insects, but it is
known mostly to insect scientists (entomologists). The public (non-entomologists or 99.99% of all peo-
ple) has little knowledge about insects, and poor access to vital information about these important
organisms. So this encyclopedia is presented to bridge the gap to better enable those with a need to
know to find fundamental information provided by more than 450 experts in the field of entomology.
We provide a broad overview of insects and their close relatives, including taxonomy, behavior, ecology,
physiology, history, and management. Importantly, we provide critical links to the entomological litera-
ture, much of which presently is unavailable for search electronically. The contributors are distinguished
entomologists from around the world. They hope that the availability of this encyclopedia will help oth-
ers to reap the benefits of centuries of discovery, and to discover the wonders that make the study of
insects so compelling. It was constructed with college and university students in mind, but others may
find it a handy reference.

John L. Capinera, Gainesville (Florida) April, 2008

Highlights of the Encyclopedia of
Major Taxa of Insects and Their Stoneflies (Plecoptera)
Near Relatives Stylopids (Strepsiptera)
Symphylans (Symphyla)
Alderflies and Dobsonflies (Megaloptera) Termites (Isoptera)
Angel Insects (Zoraptera) Thrips (Thysanoptera)
Bark-Lice, Book-Lice, or Psocids (Psocoptera) Ticks (Ixodida)
Beetles (Coleoptera) Wasps, Ants, Bees, and Sawflies (Hymenoptera)
Bristletails (Archeognatha) Webspinners (Embiidina)
Bugs (Hemiptera)
Butterflies and Moths (Lepidoptera)
Caddisflies (Trichoptera) Other Groups
Centipedes (Chilopoda)
Chewing and Sucking Lice (Phthiraptera) Anagrus Fairyflies (Hymenoptera: Mymaridae)
Cockroaches (Blattodea) Ants (Hymenoptera: Formicidae)
Diplurans (Diplura) Aphids (Hemiptera: Aphididae)
Dragonflies and Damselflies (Odonata) Apoid Wasps (Hymenoptera: Apoidea:
Earwigs (Dermaptera) Spheciformes)
Fleas (Siphonaptera) Argasid (Soft) Ticks (Acari: Ixodida: Argasidae)
Flies (Diptera) Assassin Bugs, Kissing Bugs and Others
Gladiators (Mantophasmatodea) (Hemiptera: Reduviidae)
Grasshoppers, Katydids, and Crickets Bark Beetles, Dendroctonus spp. (Coleoptera:
(Orthoptera) Curculionidae: Scolytinae)
Lacewings, Antlions, and Mantispids Bees (Hymenoptera: Apoidea: Apiformes)
(Neuroptera) Bess Beetles (Coleoptera: Passalidae)
Mayflies (Ephemeroptera) Biting Midges, Culicoides (Diptera:
Millipedes (Diplopoda) Ceratopogonidae)
Mites (Acari) Black Flies (Diptera: Simuliidae)
Pillbugs and Sowbugs, or Woodlice (Isopoda) Blister Beetles (Coleoptera: Meloidae)
Praying Mantids (Mantodea) Bulb Mites, Rhizoglyphus (Acari: Acaridae)
Proturans (Protura) Burnet Moth Biology (Lepidoptera:
Rock Crawlers (Grylloblattodea) Zygaenidae)
Scorpionflies (Mecoptera) Brush-Footed Butterflies (Lepidoptera:
Scorpions (Scorpiones) Nymphalidae)
Sea Spiders (Pycnogonida) Butterflies (Lepidoptera: Rhopalocera)
Silverfish (Zygentoma) Carpenter Bees (Hymenoptera: Apidae:
Snakeflies (Raphidioptera) Xylocopinae)
Spiders (Araneae) Carrion Beetles (Coleoptera: Silphidae)
Springtails (Collembola) Cicadas (Hemiptera: Cicadoidea)
Stick and Leaf Insects (Phasmida) Clearwing Moths (Lepidoptera: Sesiidae)
Highlights of the Encyclopedia of Entomology

Coreid Bugs and Relatives: Coreidae, Marine Insects and the Sea-Skater Halobates
Stenocepahidae, Alydidae, Rhopalidae, and (Hemiptera: Gerridae)
Hyocephalidae (Hemiptera: Coreoidea) Metalmark Butterflies (Lepidoptera: Riodinidae)
Crane Flies (Diptera: Tipulidae and Others) Microdon spp. (Diptera: Syrphidae)
Dance Flies, Balloon Flies, Predaceous Flies Minute Pirate Bugs (Hemiptera: Anthocoridae)
(Diptera: Empidoidea, exclusive of Mosquitoes (Diptera: Culicidae)
Dolichopodidae) Moths (Lepidoptera: Heterocera)
Darkling Beetles (Coleoptera: Tenebrionidae) Nuttalliellidae (Acari)
Earwigflies (Mecoptera: Meropeidae) Orchid Bees (Hymenoptera: Apidae)
Fairyflies (Hymenoptera: Mymaridae) Parasitic Hymenoptera (Hymenoptera: Parasitica)
Fireflies (Coleoptera: Lampyridae) Paederus Fabricius (Coleoptera: Staphylinidae:
Flea Beetles (Coleoptera: Chrysomelidae: Paederinae)
Alticinae) Pelecinid Wasps (Hymenoptera: Pelecinidae)
Four-Legged Mites (Eriophyoidea or Tetrapodili) Periodical Cicadas, Magicicada spp. (Hemiptera:
Fruit Flies (Diptera: Tephritidae) Cicadidae)
Fungus Gnats (Diptera: Mycetophilidae and Phytoseiid Mites (Acari: Phytoseiidae)
Others) Powderpost Beetles (Coleoptera: Bostrichidae:
Gall Midges (Diptera: Cecidomyiidae) Lyctinae)
Gall Wasps (Hymenoptera: Cynipidae) Pine Tip Moths, Rhyacionia spp. (Lepidoptera:
Giant Water Bugs (Hemiptera: Belostomatidae) Tortricidae)
Greater Fritillaries or Silverspots, Speyeria [= Plant Bugs (Hemiptera: Miridae)
Argynnis] (Lepidoptera: Nymphalidae) Planthoppers (Hemiptera: Fulgoroidea)
Ground Beetles (Coleoptera: Carabidae): Plume Moths (Lepidoptera: Pterophoridae)
Taxonomy Powderpost Beetles (Coleoptera: Bostrichidae:
Harvester Ants, Pogonomyrmex (Hymenoptera: Lyctinae)
Formicidae) Predatory Stink Bugs (Hemiptera: Pentatomidae:
Horse Flies and Deer Flies (Diptera: Tabanidae) Asopinae)
Jerusalem Crickets (Orthoptera: Riffle Beetles (Coleoptera: Elmidae)
Stenopelmatidae) Robber Flies (Diptera: Asilidae)
Jumping Spiders (Arachnida: Araneae: Salticidae) Rove Beetles (Coleoptera: Staphylinidae)
June Beetles, Phyllophaga spp. (Coleoptera: Sac Spiders (Arachnida: Araneae: Tengellidae,
Scarabaeidae: Melolothinae: Melolothini) Zorocratidae, Miturgidae, Anyphaenidae,
Katydids (Orthoptera: Tettigoniidae) Clubionidae, Liocranidae, and Corinnidae)
Kissing Bugs (Hemiptera: Reduviidae: Triatominae) Sap Beetles (Coleoptera: Nitidulidae)
Lace Bugs (Hemiptera: Tingidae) Sawflies (Hymenoptera: Symphyta)
Ladybird Beetles (Coccinellidae: Coleoptera) Sawflies (Hymenoptera: Tenthredinidae)
Leaf Beetles (Coleoptera: Chrysomelidae) Scale Insects and Mealybugs (Hemiptera:
Leaf-Cutting Ants (Formicidae: Myrmicinae: Coccoidea)
Attini) Scarab Beetles (Coleoptera: Scarabaeoidea)
Leaf-Miner Flies (Diptera: Agromyzidae) Skin-Piercing and Blood-Feeding Moths, Calyptra
Leafhoppers (Hemiptera: Cicadellidae) spp. (Lepidoptera: Noctuidae: Calpinae)
Longicorn, Longhorned, or Round-Headed Soil Mites (Acari: Oribatida and Others)
Beetles (Coleoptera: Cerambycidae) Spittlebugs (Hemiptera: Cercopoidea)
Long-Legged Flies (Diptera: Dolichopodidae) Spongillaflies (Neuroptera: Sisyridae)
Mantidflies (Neuroptera: Mantispidae) Stilt Bugs (Hemiptera: Berytidae)
Highlights of the Encyclopedia of Entomology

Stink Bugs (Hemiptera: Pentatomidae), Alarm Pheromones

Emphasizing Economic Importance Alary Muscles
Tachinid Flies (Diptera: Tachinidae) Alimentary Canal and Digestion
Tent Caterpillars, Malacosoma spp. (Lepidoptera: Biological Clock of the German Cockroach,
Lasiocampidae) Blattella germanica (L.)
Tiger Beetles (Coleoptera: Carabidae: Collyrinae Chordotonal Sensory Organs
and Cicindelinae) Corazonin
Termites (Isoptera) in South America Cryptobiosis
Tiphiid Wasps (Hymenoptera: Tiphiidae) Diapause
Treehoppers (Hemiptera: Membracidae) Dorsal Vessel: Heart and Aorta
Underwing Moths Catocala (Lepidoptera: Ecdysone Agonists, a Novel Group of Insect
Noctuidae) Growth Regulators
Velvet Ants (Hymenoptera: Mutillidae) Ecdysteroids
Water Penny Beetles (Coleoptera: Psephenidae) Embryogenesis
Weevils, Billbugs, Bark Beetles, and Others Endocrine Regulation of Insect Reproduction
(Coleoptera: Curculionoidea) Eyes and Vision
Weta (Orthoptera: Anostostomatidae, Fatty Acid Binding Proteins
Rhaphidophoridae) Fireflies: Control of Flashing
Whiteflies (Hemiptera: Aleyrodidae) Hemocytes in Insects: Their Morphology and
Morphology and Anatomy Immunity
Integument: Structure and Function
Abdomen of Hexapods Entomopathogenic Fungi and Their Host Cuticle
Antennae of Hexapods Juvenile Hormone
Head of Hexapods Learning in Insects: Neurochemistry and
Internal Anatomy of Insects Localization of Brain Function
Alimentary Canal and Digestion Locomotion and Muscles
Legs of Hexapods Metamorphosis
Mouthparts of Hexapods Midgut and Insect Pathogens
Thorax of Hexapods Multifunctional Semiochemicals
Wings of Insects Nervous System
Eggs of Insects Neurological Effects of Insecticides and the Insect
Nervous System Nervous System
Eyes and Vision Nutrition in Insects
Integument: Structure and Function Oogenesis
Nygma Pheromones
Wing Coupling Polyphenism in Insects and Juvenile Hormone
Prothoracicotropic Hormone
Physiology Reflex Bleeding
Regulation of Sex Pheromone Production in
Accessory Pulsatile Hearts Moths
Adipokinetic and Hypertrehalosemic Reproduction
Neurohormones Reselin
Highlights of the Encyclopedia of Entomology

Sex Attractant Pheromones Arthropod-Associated Plant Effectors (AAPEs):

Social Insect Pheromones Elicitors and Suppressors of Crop Defense
Sound Production in the Cicadoidea Attraction of Insects to Organic Sulfur
Storage Proteins Compounds in Plants
Storage Protein Receptors Blister Beetle Antennal Twisting Behavior
Stress-Induced Host Plant Free Amino Acids and Construction Behavior of Insects
Insects Crypsis
Taste and Contact Chemoreception Cycloalexy
Tracheal System and Respiratory Gas Exchange Deflection Marks
Ultrastructure of Insect Sensilla Deimatic Behavior
Venoms and Toxins in Insects Drumming Communication and Intersexual
Venoms of Ectoparasitic Wasps Searching Behavior of Stoneflies
Venoms of Endoparasitic Wasps (Plecoptera)
Vitellogenesis Environmental Influences on Behavioral
Water Balance Development in Insects
Facultative Predators
Flash Colors
Genetics Glowworms
Gregarious Behavior in Insects
Bacteriophage WO Hilltopping
Behavior of Insects: Genetic Analysis by Crossing Host Location in Parasitic Wasps
and Selection Host Plant Selection by Insects
Behavior: Molecular Genetic Analyses Inverted Copulation
Genetic Modification of Drosophila By P Learning in Insects
Elements Marking Insects for Studying Ecology and
Genetic Sexing Behavior
Genetic Transformation Mosquito Oviposition
Genomes of Insects Multifunctional Semiochemicals
Homeotic Genes in Coleoptera Myrmecomorphy
Honey Bee Sexuality: A Historical Perspective Myrmecophiles
Meiotic Drive in Insects Parental Care in Heteroptera (Hemiptera:
Molecular Diagnosis Heteroptera)
Polytene Chromosomes Puddling Behavior in Butterflies
Sex Ratio Modification by Cytoplasmic Agents Red Imported Fire Ant Territorial Behavior
Sibling Species Reflex Bleeding
Transgenic Arthropods for Pest Management Sociality of Insects
Programs Sound Production in the Cicadoidea
RNA Interference Sphragis
Spider Behavior and Value in Agricultural
Behavior Thermoregulation in Insects
Vibrational Communication
Acoustic Communication in Insects Visual Mating Signals
Acoustical Communication in Heteroptera Walking Stick Defensive Behavior and
(Hemiptera: Heteroptera) Regeneration of Appendages
Highlights of the Encyclopedia of Entomology

Ecology and Evolution Insectivorous Plants

Life Tables
Adaptation of Indigenous Insects to Introduced Mosquito Larval Feeding Ecology
Crops Mosquito Overwintering Ecology
Allelochemicals Natural Enemy Attraction to Plant
Ant-Plant Interactions Volatiles
Aposematism Night Blooming Plants and Their Insect
Arthropod-Associated Plant Effectors Pollinators
(AAPEs): Elicitors and Suppressors of Nutrient Content of Insects
Crop Defense Outbreaks of Insects
Bioclimatic Models in Entomology Overwintering in Insects
Biogeography Phase Polymorphism in Locusts
Brachelytry Phylogenetics
Bromeliad Fauna Phytotelmata
Cannibalism Plant Extrafloral Nectaries
Carnivorous Plants Plant Secondary Compounds and Phytophagous
Carnivory and Symbiosis in the Purple Insects
PitcherPlant Pollination and Flower Visitation
Castes Pollination by Osmia Bees (Hymenoptera:
Cave Adapted Insects Megachilidae)
Cave Habitat Colonization Pollination by Yucca Moths
Chemical Ecology of Insects Pollution and Terrestrial Arthropods
Cold Tolerance in Insects Polyphenism
Conservation of Insects Polyacetylenes and Their Thiophene
Cryptobiosis Derivatives
Decomposer Insects Predation: The Role of Generalist Predators
Diapause inBiodiversity and Biological
Eggs of Insects Control
Endophytic Fungi and Grass-Feeding Pyrophilous Insects
Insects Pyrrolizidine Alkaloids and Tiger Moths
Food Habits of Insects (Lepidoptera: Arctiidae)
Fossil Record of Insects Retournement and Deversement of the
Furanocoumarins Aedeagusin Coleoptera
Gall Formation Rhythms of Insects
Galpagos Islands Insects: Colonization, Seed Predation by Insects
Structure, and Evolution Sexual Selection
Geological Time Sociality of Insects
Glowworms Species Concepts
Gregarious Behavior in Insects Speciation Processes Among Insects
Ground Beetle (Coleoptera: Carabidae) Symbiosis Between Planthoppers and
FeedingEcology Microorganisms
Hypertely Trichomes and Insects
Hypericin Tritrophic Interactions
Inquilines and Cleptoparasites Water Pollution and Insects
Highlights of the Encyclopedia of Entomology

Microbiology and Pathology Nosema disease

Microbial Control of Insects Orcytes virus and other Nudiviruses
Microbial Control of Medically Important Paecilomyces
Insects Paenibacillus
Symbionts of Insects Parvoviruses
Symbiosis Between Planthoppers and Polydnaviruses
Microorganisms Reoviruses
Symbiotic Viruses of Parasitic Wasps Rhabdoviruses
Fungal Pathogens in Insects Septobasidium
Entomopathogenic Fungi and Their Host Cuticle Serratia entomophila
Midgut and Insect Pathogens Small RNA viruses of invertebrates
Immunity Sorosporella
Pathogens of Whiteflies (Hemiptera: Trichomycetes
Diseases Of Grasshoppers and Locusts
Nematode Parasites of Insects Humans and Insects
Nematomorphs (Nematomorpha)
Entomopathogenic Nematodes and Insect Apiculture (Beekeeping)
Management Aquatic Entomology and Flyfishing
Thrips-Parasitic Nematodes Butterfly Counts
American Foulbrood Butterfly Gardening
Amoebae Careers in Entomology
Ascosphaera apis Collecting and Preserving Insects
Ascoviruses Common (Vernacular) Names of Insects
Bacillus sphaericus Conservation of Insects
Bacillus thuringiensis Costs and Benefits of Insects
Baculoviruses Cultural Entomology
Beauveria Entomodeltiology
Birnaviruses Entomophagy: Human Consumption of Insects
Cestodes Forensic Entomology
Coelomomyces History and Insects
Culicinomyces Honeybee Sexuality: An Historical Perspective
Entomophthorales Identification of Insects
Entomopoxviruses (Poxviruses) Insects as Aphrodisiacs
Granulovirus Invasive Species
Helicosporium Lacquers and Dyes From Insects
Hirsutella Literature and Insects
Iridoviruses Maggot Therapy
Laboulbeniales Marking Insects for Studying Ecology and
Lagenidium giganteum Behavior
Lecanicillium Midges as Human Food
Metarhizium Mythology and Insects
Microsporidia (Phylum Microsporidia) Native American Culture and Insects
Nosema locustae (Protozoa: Microsporidia) Popularity of Insects
Highlights of the Encyclopedia of Entomology

Pronunciation of Scientific Names and Terms Bokor, Elemr

Psychiatry And Insects: Phobias and Delusions of Bonnet, Charles
Insect Infestations in Humans Borgmeier, Thomas
Rearing of Insects Bving, Adam Giede
Scientific Names and Other Words From Latin Brauer, Friedrich Moritz
and Greek Broun, Thomas
Sericulture Bruch, Carlos
Silk Brues, II, Charles Thomas
Teaching and Training in Entomology: Institu- Brull, Gaspard Auguste
tional Models Brunner von Wattenwyl, Carl
Teaching Entomology: A Review of Techniques Buffon, Georges Louis Leclerc (Comte de)
Burgess, Albert Franklin
Burmeister, Carl Hermann Conrad
Notable and Pioneer Buxton, Patrick Alfred
Entomologists Calvert, Philip Powell
Cameron, Malcolm
Abafi-Aigner, Lajos (Ludwig Aigner) Candze, Ernest Charles Auguste
Abbott, John Carpenter, Frank Morton
Agassiz, Jean Louis Rodolphe Carter, Herbert James
Aldrovandi, Ulisse (Ulysse, Ulysses) Casey, Thomas Lincoln
Alexander, Charles Paul Caudell, Andrew Nelson
Amerasinghe, Felix P. Chagas, Carlos Justiniano Ribeiro
Andrewartha, Herbert George Chaudoir, Maximilien Stanislavovitch de
Arnett, Jr., Ross Harold Champion, George Charles
Audinet-Serville, Jean-Guillaume Chapman, Reginald Frederick
Audouin, Jean-Victor Chevrolat, Louis Alexandre Auguste
Barber, Herbert Spencer Chiang, Huai C.
Bates, Henry Walter China, William Edward
Bates, Marston Chittenden, Frank Hurlbut
Beck, Stanley Dwight Christophers, (Sir) Samuel Rickard
Beklemishev, Vladimir NikolayevicH Cockerell, Theodor Dru Alison
Bell, William J. Comstock, John Henry
Becquaert, Joseph Charles Coquillett, Daniel William
Berlese, Antonio Craw, Alexander
Bernhauer, Max Cresson, Ezra Townsend
Bertram, Douglas Somerville Criddle, Norman
Br, Lajos Crotch, George Robert
Blackburn, Thomas Crowson, Roy Albert
Blackwelder, Richard Elliot Csiki, Erno (Ernst Dietl)
Blaisdell, Frank Ellsworth Curran, Charles Howard
Blatchley, Willis Stanley Curtis, John
Bodenheimer, Friedrich (Frederick) Simon Cushman, Robert Asa
Bohart, Richard M. Cuvier, (Baron) Georges Lopold Chretien
Boheman, Carl Heinrich Frdric Dagobert
Boisduval, Jean-Baptiste Alphonse Dechauffour Dahlbom, Anders Gustav
Highlights of the Encyclopedia of Entomology

Darlington, Jr., Philip J. Frivaldszky, Jnos

Darwin, Charles Froggatt, Walter Wilson
Davis, William Thompson Gahan, Arthur Burton
Debach, Paul Ganglbauer, Ludwig
Degeer, Carl (Carolus, Karl, Charles) Germar, Ernst Friedrich
Dejean, Pierre Franois Marie Auguste Ghilarov, Mercury Sergeevich
Dethier, Vincent Gaston Girault, Alexandre Arsne
Dobzhansky, Theodosius Grigorievich Glover, Townend
Donisthorpe, Horace St. John Kelly Gmelin, Johann Friedrich
Douglas, John William Goeldi, Emil (Emilio) August
Drake, Carl John Gorgas, William Crawford
Dudich, Endre Graham, Marcus William Robert de Vere
Dufour, Lon Jean Marie Grassi, Giovanni Battista
Duponchel, Philogne August-Joseph Gravenhorst, Johan Ludwig Christian
Dyar, Harrison Grey Gressitt, Judson Linsley
Edwards, William Henry Grote, Augustus Radcliffe
Eickwort, George C. Guene, Achille
Emerson, Alfred Edwards Gurin-Mneville, Flix Edouard
Endrdi, Seb (Sebastian Endrdi) Gundlach, Johannes (Juan) Christopher
Endrdy-Younga, Sebastian (Sebestyn Gyllenhal, Leonhard
Endrdy-Younga, Sebestyn Endrdy) Haddow, Alexander John
Erichson, Wilhelm Ferdinand Hagen, Hermann August
Esaki, Teiso Hagen, Kenneth Sverre
Evans, Howard Ensign Hahn, Carl Wilhelm
Fabre, Jean-Henri Casimir Hale Carpenter, Geoffrey Douglas
Fabricius, Johann Christian Haliday, Alexander Henry
Fairchild, Alexander Graham Bell Handlirsch, Anton
Fairmaire, Lon Hansen, Viktor
Fall, Henry Clinton Harris, Thaddeus William
Fauvel, Charles Adolphe Albert Hatch, Melville Harrison
Felt, Ephraim Porter Heer, Oswald
Fenichel, Smuel Henneguy, Louis Flix
Fennah, Ronald Gordon Hennig, Willi
Fernald, Charles Henry Herman, Ott
Ferris, Gordon Floyd Herrich-Schffer, Gottlieb August
Fischer Von Waldheim, Gotthelf Hewitt, Charles Gordon
Fitch, Asa Hinton, Howard Everest
Fletcher, James Hobby, Bertram Maurice
Foote, Richard H. Hocking, Brian
Forbes, Stephen Alfred Hogstraal, Harry O.
Ford, Edmund Brisco Holland, William Jacob
Forel, Auguste Henri Hope, Frederick William
Fourcroy, Antoine-Franois (Comte de) Hopkins, Andrew Delmar
Frisch, K. von Horn, George Henry
Frivaldszky, Imre Horn, Hermann Wilhelm Walther
Highlights of the Encyclopedia of Entomology

Horsfall, William R. Maskell, William Miles

Howard, Leland Ossian Masters, George
Hbner, Jacob Matheson, Robert
Huffaker, Carl Barton Matsumura, Shonen
Hungerford, Herbert Barker Mcdunnough, James Halliday
Ihering, Hermann Von Mcglashan, Charles Fayette
Imms, Augustus Daniel Meigen, Johann Wilhelm
Jacquelin du Val, Pierre Nicolas Camille Mellanby, Kenneth
Jeannel, Ren Melander, Axel Leonard
Johannsen, Oskar Augustus Melsheimer, Frederick Valentine
Jordan, Heinrich Ernst Karl Mntris, Edouard
Kaszab, Zoltn Metcalf, Clell Lee
Kellogg, Vernon Lyman Metcalf, Zeno Payne
Kennedy, John S. Meyrick, Edward
Kershaw, John Crampton Wilkinson Miller, David
Kevan, Douglas Keith McEwan Mlachlan, Robert
Kiesenwetter, Ernst August Hellmuth von Morgan, Thomas Hunt
Kirby, William Morrison, Herbert Knowles
Klee, Waldemar G. Morse, Albert Pitts
Knipling, Edward Fred Motschulsky, Victor Ivanovich
Koebele, Albert Mller, Johann Friedrich Theodor
Kraatz, Ernst Gustav Mller, Josef
Kring, James Burton Mulsant, Etienne
Lamarck, Jean-Baptiste Needham, James George
Latreille, Pierre Andr Newell, Wilmon
Lea, Arthur Mills Newman, Edward.
Lea, H. Arnold Newsom, Leo Dale
LeConte, John Lawrence Olivier, Guillaume Antoine
Leech, Hugh Bosdin Ormerod, Eleanor Anne
Lefroy, Harold Maxwell Osborn, Herbert
Leng, Charles William Osten Sacken, C.R.
Lepeletier, Amde Louis Michel Packard, Alpheus Spring
Lindroth, Carl H. Painter, Reginald Henry
Linnaeus, Carolus (Linn, Carl von) Palm, Thure
Linsley, Earle Gorton Pass, Bobby Clifton
Loew, Hermann Patch, Edith Marion
Lorguin, Pierre Joseph Michel Paykull, Gustaf
Lugger, Otto Peairs, Leonard Marion
Macleay (Sir) William John Peck, William Dandridge
Macquart, Pierre Justin Marie Petrunkevitch, Alexander
Mallis, Arnold Pickett, Allison Deforest
Mann, William M. Pomerantsev, Boris Ivanovich
Mannerheim, Carl Gustav von Potter, Charles
Marlatt, Charles Lester Prokopy, Ronald J.
Marx, George Provancher, (labb) Lon
Highlights of the Encyclopedia of Entomology

Putzeys, Jules Antoine Adolphe Henri Stern, Vernon M.

Rabb, Robert Lamar Swammerdam, Jan
Radoshkowsky, Octavius John Swezey, Otto Herman
de Raumur, Ren Antoine Ferchault Szent-Ivny, Jzsef Gyula Hubertus
Redi, Francesco Thomas, Cyrus
Redtenbacher, Ludwig Thomson, Carl Gustav
Reed, Walter. Thunberg, Carl Peter
Rehn, James Abram Garfield Tillyard, Robin John
Reitter, Edmund Torre-Bueno, Jos Rollin De La
Remington, Charles Lee Townes, Jr., Henry K.
Rey, Claudius Treherne, John E
Richards, Owain Westmacott Uhler, Philip Reese
Riley, Charles Valentine Usinger, Robert Leslie
Robineau-Desvoidy, Jean Baptiste Uvarov, (Sir) Boris Petrovich
Rondani, Camillo Van Den Bosch, Robert
Rosen, David Van Duzee, Edward Payson
Ross, Herbert Holdsworth Varley, George C
Rothschild, Miriam Viereck, Henry Lorenz
Roubal, Jan Walker, Francis
Sabrosky, Curtis Waloff, Nadejda
Sahlberg, Carl Reinhold Walsh, Benjamin Dann
Sailer, Reece I. Walsingham, (Lord) Thomas de Gray
Sanderson, Dwight Wasmann, Erich
Saunders, William Wesmael, Constantin
Saussure, Henri Louis Frederic de Westwood, John Obadiah
Say, Thomas Wheeler, William Morton
Scheerpeltz, Otto Wiedemann, Christian Rudolph Wilhelm
Schneiderman, Howard Allen Wigglesworth, (Sir) Vincent Brian
Schwarz, Eugene Amandus Williams, Carroll Milton
Scudder, Samuel Hubbard Williston, Samuel Wendell
Selys-Longchamps, Michel Edmond de Wirth, Willis Wagner
Sharp, David Wolcott, George N.
Shelford, Victor Ernest Wollaston, Thomas Vernon
Signoret, Victor Antoine Young, Jr., David A
Silvestri, Filippo Zeller, Philipp Christoph
Sylveira Caldeira, Joo da Zetterstedt, Johann Wilhelm
Smirnoff, Vladimir A. Zachvatkin (Jasykov), Aleksei Alekseevich
Smith, John Bernhardt
Smith, Harry Scott
Smith, Ray F. Pest Management
Snodgrass, Robert Evans
Snow, Francis Huntington Integrated Pest Management (IPM)
Spielman, Andrew Economic Injury Level (EIL) and Economic
Stl, Carl Threshold (ET) Concepts in Pest
Steinhaus, Edward Arthur Management
Highlights of the Encyclopedia of Entomology

Costs and Benefits of Insects Augmentative Biological Control

Methods for Measuring Crop Losses by Insects Classical Biological Control
Sampling Arthropods Conservation Biological Control
Traps for Capturing Insects Natural Enemies Important in Biological Control
Scale and Hierarchy in Integrated Pest Mass Rearing of Natural Enemies
Management Culture of Natural Enemies on Factitious Foods
Phenology Models for Pest Management and Artificial Diets
Agroecology Classical Biological Control of Chestnut Gall
Organic Agriculture Wasp in Japan
Regulatory Entomology Conservation of Ground Beetles in
Regulations Affecting the Implementation of Annual Crops
Regulatory Entomology Practices Filth Fly Parasitoids (Hymenoptera:
Invasive Species Pteromalidae) in North America
Pest Risk Analysis Carabid Beetles (Coleoptera: Carabidae) as
Mechanical Protection of Humans from Parasitoids
Arthropod Attacks and Bites History of Biological Control of Wheat Stem
Repellents of Biting Flies Sawflies (Hymenoptera: Cephidae)
Physical Management of Insect Pests Parasitism of Lepidoptera Defoliators in
Kaolin-Based Particle Films for Arthropod Sunflower and Legume Crops, and A djacent
Control Vegetation in the Pampas of Argentina
Controlled Atmosphere Technologies for Insect Transmission of Plant Diseases by Insects
Control Plant Viruses and Insects
Cultural Control of Insects Management of Insect-Vectored Pathogens of
Plant Resistance to Insects Plants
Resistance of Solanaceous Vegetables to Insects Transmission of Xylella fastidiosa Bacteria by
Cover, Border and Trap Crops for Pest and Xylem-Feeding Insects
Disease Management Vectors of Phytoplasmas
Microbial Control of Insects
Visual Attractants and Repellents in IPM Pesticides and Pesticide
Push-Pull Strategy for Insect Pest Management Application
Area-Wide Insect Pest Management
Sterile Insect Technique Acaricides or Miticides
Filter Rearing System for Sterile Insect Boric Acid
Technology DDT
Weeds in Crop Systems for Pest Suppression Insecticides
Biological Control of Weeds Insecticide Application: The Dose Transfer
Weed Biological Control in Australia Process
Biological Control of Invasive Plants in Latin Food-Based Poisoned Baits for Insect Control
America Soil Fumigation
Host Specificity of Weed-Feeding Insects Structural Fumigation
Foreign Exploration for Insects that Feed on Insecticide Bioassay
Weeds Insecticide Formulation
Flower Strips as Conservation Areas for Pest Insecticide Resistance
Management Insecticide Toxicity
Highlights of the Encyclopedia of Entomology

Pesticide Hormoligosis Maize (Corn) Pests and Their Management

Synergism Mite Pests of Crops in Asia
Chronotoxicology Musk Thistle Suppression Using Weevils For
Detoxification Mechanisms in Insects Biological Control
Enhanced Biodegradation of Soil-Applied Palm Insects
Pesticides Pests and Their Natural Enemies (Parasitoids
Pesticide Resistance Management and/or Predators) in The Middle East
Natural Products Used for Insect Control Potato Pests and Their Management
Botanical Insecticides School IPM, or Pest Management on School
Chinaberry, Melia azedarach, a Biopesticidal Tree Grounds
Pyrethrum and Persian Insect Powder Shade Tree Arthropods and Their Management
Neem Small Fruit Pests and Their Management
Horticultural Oils Stored Grain and Flour Insects and Their
Soaps as Insecticides Management
Diatomaceous Earth Sugarcane Pests and Their Management
Photodynamic Action in Pest Control and Sweetpotato Weevils and Their Eradication
Medicine Programs in Japan
Neurological Effects of Insecticides and the Insect Tropical Fruit Pests and Their Management
Nervous System Turfgrass Insects of the United States and Their
Regulations Affecting Use of Pesticides Management
Urban Entomology
Vegetable Pests and Their Management
Pests Groups and Their Veterinary Pests and Their Management
Management Wheat Pests and Their Management
Whitefly Bioecology and Management in Latin
Agricultural Crop Pests in Southeast Asia America
Including South China Wood-Attacking Insects
Alfalfa (Lucerne) Pests and Their Management
Apple Pests and Their Management
Banana Pests and Their Management Medical and Veterinary
Bark Beetles in The Genus Dendroctonus Entomology
Cassava Pests and Their Management
Citrus Pests and Their Management African Horse Sickness Viruses
Coffee Pests and Their Management Argasid (Soft) Ticks (Acari: Ixodida: Argasidae)
Crucifer Pests and Their Management Avian (Bird) Malaria
Desert Locust, Schistocerca gregaria Forskl Bed Bugs (Hemiptera: Cimicidae: Cimex spp.)
(Orthoptera: Acrididae) Plagues Biting Midges, Culicoides spp. (Diptera:
Grasshoppers and Locusts as Agricultural Pests Ceratopogonidae)
Grasshoppers of the Argentine Pampas Black Flies Attacking Livestock: Simulium
Grasshopper and Locust Pests in Africa arcticum Malloch and Simulium luggeri
Grasshopper and Locust Pests in Australia Nicholson & Mickel (Diptera: Simuliidae)
Grasshopper Pests in North America Bluetongue Disease
Gramineous Lepidopteran Stem Borders in Bovine Hypodermosis: Phenology in Europe
Africa Brown Dog Tick, Rhipicephalus sanguineus
Mahogany Pests and Their Management (Latreille) (Acari: Ixodida: Ixodidae)
Highlights of the Encyclopedia of Entomology

Cat Flea, Ctenocephalides felis felis Bouch Rocky Mountain Wood Tick, Dermacentor
(Siphonaptera: Pulicidae) andersoni Stiles (Acari: Ixodidae)
Chagas, Carlos Justiniano Ribeiro Rodent Trypanosomiasis: A Comparison
Chagas Disease or American Trypanosomiasis Between Trypanosoma lewisi and
Chagas Disease: Biochemistry of the Vector Trypanosoma musculi
Chikungunya Simulium spp. Vectors of Onchocerca volvulus
Chironomids as a Nuisance and of Medical Skin-Piercing and Blood-Feeding Moths,
Importance Calyptra spp. (Lepidoptera: Noctuidae:
Cockroaches and Disease Calpinae)
Dermatitis Linearis Sleeping Sickness or African Trypanosomiasis
Dengue St. Louis Encephalitis
Dirofilariasis Sugar-Feeding in Blood-Feeding Flies
Eastern Equine Encephalitis Taiga Tick, Ixodes persulcatus Schulze (Acari:
Horse Flies and Deer flies Ixodida: Ixodidae)
House Fly, Musca domestica L. (Diptera: Tick Paralysis
Muscidae) Ticks (Ixodida)
Human Botfly, Dermatobia hominis (Linneaus, Ticks as Vectors of Pathogens
Jr.) (Diptera: Oestridae) Trypanosomes
Human Lice Tsetse Flies, Glossina spp. (Diptera: Glossinidae)
Human Lymphatic Filariasis (Elephantiasis) Types of Pathogen Transmission by Arthropods
Human Scabies Vector Capability of Blood-Sucking Arthropods:
Hypodermosis in Deer A Forecasting Matrix
Japanese Encephalitis Venoms And Toxins in Insects
La Crosse Encephalitis West Nile Fever
Leishmaniasis Yellow Fever
Lyme Borreliosis
Maggot Therapy
Malaria Arthropods of Economic
Mechanical Protection of Humans from Arthro- Importance
pod Attacks and Bites
Microbial Control of Medically African Armyworm, Spodoptera exempta
Important Insects (Lepidoptera: Noctuidae)
Mites (Acari) African Honey Bee, Africanized Honey Bee, or
Mosquitoes (Diptera: Culicidae) Killer Bee, Apis mellifera scutellata Lepeletier
Mosquitoes as Vectors of Viral Pathogens (Hymenoptera: Apidae)
Mosquito Oviposition African Mahogany-Feeding Caterpillar,
Myiasis Heteronygmia dissimilis Aurivillius
Onchocerciasis (Lepidoptera: Lymantriidae)
Paederina African Pine-Feeding Grasshopper, Plagiotriptus
Pederin pinivorus (Descamps) (Orthoptera:
Pathogen Transmission by Arthropods Eumastacidae)
Piroplasmosis: Babesia and Theileria Alfalfa Leafcutting Bee, Megachile rotundata
Reed, Walter Fabricius (Hymenoptera: Megachilidae)
Repellents of Biting Flies Allegheny Mound Ant, Formica exsectoides Foretl
Rocky Mountain Spotted Fever (Hymenoptera: Formicidae)
Highlights of the Encyclopedia of Entomology

American Grasshopper, Schistocerca americana Cabbage Looper, Trichoplusia ni (Hbner)

(Drury) (Orthoptera: Acrididae) (Lepidoptera Noctuidae)
Almond Seed Wasp, Eurytoma amygdali Cabbageworm, Pieris rapae (Linnaeus)
Enderlein (Hymenoptera: Eurytomidae) (Lepidoptera: Pieridae)
American Serpentine Leafminer, Liriomyza Cactus Moth, Cactoblastis cactorum (Berg)
trifolii (Burgess) (Diptera: Agromyzidae) (Lepidoptera: Pyralidae)
Argentine Ant, Linepithema humile (Mayr), Cape Honey Bees, Apis mellifera capensis Escholtz
(Hymenoptera: Formicidae: Cassava Mealybug, Phenacoccus manihoti
Dolichoderinae) Matile-Ferrero (Hemiptera:
Army Cutworm, Euxoa auxiliaris (Grote) Pseudococcidae)
(Lepidoptera: Noctuidae) Chilli Thrips, Scirtothrips dorsalis Hood
Armyworm, Pseudaletia unipuncta (Haworth) (Thysanoptera: Thripidae)
(Lepidoptera: Noctuidae) Chinch Bug, Blissus leucopterus (Say) (Hemiptera:
Asian Citrus Psyllid, Diaphorina citri Kuwayama Blissidae)
(Hemiptera: Psyllidae) Cluster Fly, Pollenia rudis (Fabricius) and
Asparagus Aphid, Brachycorynella asparagi P.pseudorudis Rognes (Diptera:
(Mordvilko) (Hemiptera: Aphididae) Calliphoridae)
Aster Leafhopper, Macrosteles quadrilineatus Coconut Mite, Aceria guerreronis (Acari:
Forbes (Hemiptera: Cicadellidae) Eriophyidae)
Australian Sheep Blowfly, Lucilia cuprina Coffee Berry Borer, Hypthenemus hampei
Wiedemann (Diptera: Calliphoridae) (Ferrari) (Coleoptera: Curculionidae:
Banana Weevil, Cosmopolites sordidus (Germar) Scolytinae)
(Coleoptera: Curculionidae) Colorado Potato Beetle, Leptinotarsa
Bed Bugs (Hemiptera: Cimicidae: Cimex spp.) decemlineata (Say) (Coleoptera:
Bee Louse, Bee Fly, or Braulid, Braula coeca Chrysomelidae)
Nitzsch (Diptera: Braulidae) Corn Delphacid, Peregrinus maidis (Ashmead)
Beet Armyworm, Spodoptera exigua (Hbner) (Hemiptera: Delphacidae)
(Lepidoptera: Noctuidae) Corn Earworm, Helicoverpa zea (Boddie)
Bertha Armyworm, Mamestra configurata Walker (Lepidoptera: Noctuidae)
(Lepidoptera: Noctuidae) Corn Leaf Aphid, Rhopalosiphum maidis (Fitch)
Black Cutworm, Agrotis ipsilon (Hufnagel) (Hemiptera: Aphididae)
(Lepidoptera: Noctuidae) Corn Leafhopper, Dalbulus maidis (Delong And
Black Fig Fly, Silba adipata McAlpine (Diptera: Wolcott) (Hemiptera: Cicadellidae)
Lonchaeidae) Cotton Leafworm, Spodoptera littoralis
Bogong moth, Agrotis infusa (Boisduval) (Boisduval)
(Lepidoptera: Noctuidae) Crapemyrtle Aphid Sarucallis kahawaluokalani
Boll Weevil, Anthonomus grandis Boheman (Kirkaldy) (Hemiptera: Aphididae)
(Coleoptera: Curculionidae) Date Palm Stem Borer, Pseudophilus testaceus
Brown Citrus Aphid, Toxoptera citricida Gah. (= Jebusea hammershmidti Reiche)
(Kirkaldy) (Hemiptera: Aphididae) (Coleoptera: Cerambycidae)
Cabbage Aphid, Brevicoryne brassicae (L.) Diamondback Moth, Plutella xylostella
(Hemiptera: Aphididae) (Linnaeus) (Lepidoptera: Plutellidae)
Cabbage Maggot or Cabbage Root Fly, Delia radi- Diaprepes Root Weevil, Diaprepes abbreviatus
cum (Linnaeus) (Diptera: Anthomyiidae) (L.) (Coleoptera: Curculionidae)
Highlights of the Encyclopedia of Entomology

Differential Grasshopper, Melanoplus differentialis Greater Date Moth, Arenipses sabella Hmps
(Thomas) (Orthoptera: Acrididae) (Lepidoptera: Pyralidae)
Douglas-Fir Beetle, Dendroctonus pseudotsugae Greenhouse Whitefly, Trialeurodes vaporariorum
pseudotsugae Hopkins (Coleoptera: (Westwood) (Hemiptera: Aleyrodidae)
Curculionidae, Scolytinae) Green Peach Aphid, Myzus persicae (Sulzer)
Driver Ants (Dorylus Subgenus Anomma) (Hemiptera: Aphididae)
(Hymenoptera: Formicidae) Gypsy Moth, Lymantria dispar Linnaeus
Dubas Bug (Old World Date Bug), Ommatissus (Lepidoptera: Lymantriidae)
binotatus (Hemiptera: Tropiduchidae) Harlequin Bug, Murgantia histrionica (Hahn)
Eastern Lubber Grasshopper, Romalea microptera (Hemiptera: Pentatomidae)
(Beauvois) (Orthoptera: Acrididae) Hazelnut and Walnut Twig Borer, Oberea
Elm Leaf Beetle, Xanthogaleruca (= Pyrrhalta) linearis Linnaeus (Coleoptera:
luteola (Mller) (Coleoptera: Cerambycidae)
Chrysomelidae) Hessian Fly, Mayetiola destructor (Say) (Diptera:
Eri Silkworm, Philosamia ricini (Lepidoptera: Cecidomyiidae)
Saturniidae) Honey Bee, Apis mellifera Linnaeus
Eurasian Spruce Bark Beetle, Ips typographus (Hymenoptera: Apidae)
Linnaeus (Coleoptera: Curculionidae, Horn Fly, Haematobia irritans (L.) (Diptera:
Scolytinae) Muscidae)
European Cherry Fruit Fly Rhagoletis cerasi House Fly, Musca domestica L. (Diptera:
(Linnaeus) (Diptera: Tephritidae) Muscidae)
European Corn Borer, Ostrinia nubilalis (Hbner) Human Botfly, Dermatobia hominis (Linneaus,
(Lepidoptera: Pyralidae) Jr.) (Diptera: Oestridae)
European Earwig, Forficula auricularia Linnaeus Japanese Beetle, Popillia japonica Newman
(Dermaptera: Forficulidae) (Coleoptera: Scarabaeidae)
Face Fly, Musca autumnalis De Geer (Diptera: Jewel Wasp, Nasonia vitripennis (Walker)
Muscidae) (Hymenoptera: Pteromalidae)
Fall Armyworm, Spodoptera frugiperda (J.E. Khapra Beetle, Trogoderma granarium Everts
Smith) (Lepidoptera: Noctuidae) (Coleoptera: Dermestidae)
Formosan Subterranean Termite, Coptotermes Large Milkweed Bug, Oncopeltus fasciatus
formosanus Shiraki (Isoptera: (Dallas) (Hemiptera: Lygaeidae)
Rhinotermitidae) Large Cabbage White Butterfly,
Fruit Stalk Borer, Oryectes elegans Prell Pieris brassicae (Linnaeus) (Lepidoptera:
(Coleoptera: Scarabaeidae) Pieridae)
Gamagrass Leafhopper Dalbulus quinquenotatus Larger Grain Borer, Prostephanus truncatus
Delong & Nault (Hemiptera: (Horn) (Coleoptera: Bostrichidae)
Cicadellidae) Leaf-Cutting Ants (Formicidae: Myrmicinae:
Glassy-Winged Sharpshooter, Homalodisca Attini)
vitripennis (Hemiptera: Cicadellidae) Lesser Date Moth, Batrachedra amydraula
Grape Phylloxera, Daktulosphaira vitifoliae Meyrick (Lepidoptera: Cosmopterygidae)
(Fitch) (Hemiptera: Aphidoidea: Lettuce Root Aphid, Pemphigus bursarius
Phylloxeridae) (Linnaeus) (Hemiptera: Aphididae)
Grapevine Leafhopper Complex (Hemiptera: Locust Borer, Megacyllene robiniae (Forster)
Cicadellidae) in Cyprus (Coleoptera: Cerambycidae)
Highlights of the Encyclopedia of Entomology

Lovebug, Plecia nearctica Hardy (Diptera: Onion Maggot, Delia antiqua (Meigen) (Diptera:
Bibionidae) Anthomyiidae)
Mango Mealybug, Rastrococcus invadens Williams Oriental Fruit Fly, Bactrocera dorsalis (Hendel)
(Hemiptera: Pseudococcidae) (Diptera: Tephritidae)
Mediterranean Fruit Fly Ceratitis capitata Pea Aphid, Acyrthosiphon pisum (Harris)
(Wiedemann) (Diptera: Tephritidae) (Hemiptera: Aphididae)
Melon Aphid, Aphis gossypii Glover (Hemiptera: Pea Leafminer, Liriomyza huidobrensis
Aphididae) (Blanchard) (Diptera: Agromyzidae)
Melon Fly, Bactrocera cucurbitae (Coquillett) Pear Psylla, Cacopsylla pyricola (Foerster)
(Diptera: Tephritidae) (Hemiptera: Psyllidae)
Melon Thrips, Thrips palmi Karny (Thysanoptera: Pepper Weevil, Anthonomus eugenii Cano
Thripidae) (Coleoptera: Curculionidae)
Melonworm, Diaphania hyalinata Linnaeus Phoracantha Longicorn Beetles (Coleoptera:
(Lepidoptera: Pyralidae) Cerambycidae)
Mexican Bean Beetle, Epilachna varivestris Pickleworm, Diaphania nitidalis (Stoll)
Mulsant (Coleoptera: Coccinellidae) (Lepidoptera: Pyralidae)
Migratory Grasshopper, Melanoplus sanguinipes Pine Weevil, Hylobius abietis (Coleoptera:
(Fabricius) (Orthoptera: Acrididae) Curculionidae)
Mole Crickets (Orthoptera: Gryllotalpidae) and Pink Hibiscus Mealybug, Maconellicoccus
Their Biological Control hirsutus Green (Hemiptera:
Monarch Butterfly, Danaus plexippus Linnaeus Pseudococcidae)
(Lepidoptera: Danaidae) Pistachio Seed Wasps, Eurytoma plotnikovi
Mormon Cricket, Anabrus simplex Haldeman Nikolskaya (Hymenoptera: Eurytomidae)
(Orthoptera: Tettigoniidae) and Megastigmas pistaciae Walker
Mountain Pine Beetle, Dendroctonus ponderosae (Hymenoptera: Torymidae)
Hopkins (Coleoptera: Curculionidae, Potato Aphid, Macrosiphum euphorbiae (Thomas)
Scolytinae) (Hemiptera: Aphididae)
Myndus crudus (Van Duzee) (Hemiptera: Potato Tuberworm, Phthorimaea operculella
Cixiidae) (Zeller) (Lepidoptera: Gelechiidae)
Neotropical Brown Stink Bug, Euschistus Plum Curculio, Conotrachelus nenuphar Herbst
heros (Fabricius) (Hemiptera: (Coleoptera: Curculionidae)
Pentatomidae) Pseudo-Curly Top Treehopper, Micrutalis
Neotropical Soybean Budborer, Crocidosema malleifera (Fowler) (Hemiptera:
aporema (Walsingham) (Lepidoptera: Membracidae)
Tortricidae) Red Imported Fire Ant, Solenopsis invicta Buren
Northern Corn Rootworm, Diabrotica barberi (Hymenoptera: Formicidae)
Smith & Lawrence (Coleoptera: Redlegged Grasshopper, Melanoplus
Chrysomelidae) femurrubrum (Degeer) (Orthoptera:
Olive Fruit Curculio, Rhynchites (= Coenorrhinus) Acrididae)
cribripennis Desbrochers (Coleoptera: Red Palm Weevil, Rhynchophorus ferrugineus
Attelabidae) (Olivier) (Coleoptera: Curculionidae)
Olive Fruit Fly, Bactrocera oleae (Rossi) (= Dacus Rhammatocerus schistocercoides Rehn
oleae) (Diptera: Tephritidae) (Orthoptera: Acrididae)
Olive Psyllids, Euplyllura spp. (Hemiptera: Rocky Mountain Wood Tick, Dermacentor
Psyllidae) andersoni Stiles (Acari: Ixodidae)
Highlights of the Encyclopedia of Entomology

Roundheaded Pine Beetle, Dendroctonus adjunc- Tarnished Plant Bug, Lygus lineolaris Palisot de
tus Blandford (Coleoptera: Curculionidae, Beauvois (Hemiptera: Miridae)
Scolytinae) Taro Caterpillar or Rice Cutworm, Spodoptera
Seedcorn Maggot or Bean Seed Fly, Delia platura litura (Fabricius)
(Meigen) (Diptera: Anthomyiidae) Tent Caterpillars, Malacosoma spp. (Lepidoptera:
Silkworm, Bombyx mori (Linnaeus) (Lepidoptera: Lasiocampidae)
Saturniidae) Termites (Isoptera) in South America
Silverleaf Whitefly, Bemisia argentifolii Bellows Timarcha Latreille (Coleoptera: Chrysomelidae,
and Perring (Hemiptera: Aleyrodidae) Chrysomelinae)
Small Green Stink Bug, Piezodorus guildinii Tomato Hornworm, Manduca quinquemaculata
(Westwood) (Hemiptera: Heteroptera: (Haworth) and Tobacco Hornworm,
Pentatomidae) Manduca sexta (Linnaeus) (Lepidoptera:
Small Hive Beetle, Aethina tumida Murray Sphingidae)
(Nitidulidae: Coleoptera) Tracheal Mite, Acarapis woodi Rennie (Acarina:
Small Honey Ant, Prenolepis imparis (Say) Tarsonemidae)
(Hymenoptera: Formicidae) Tsetse Flies, Glossina spp. (Diptera: Glossinidae)
Small Rice Stink Bug, Oebalus poecilus (Dallas) Turnip Aphid, Lipaphis erysimi (Kaltenbach)
(Hemiptera: Heteroptera: Pentatomidae) (Hemiptera: Aphididae)
Southern Green Stink Bug, Nezara viridula (L.) Turnip Root Maggot, Delia floralis (Fallen)
(Hemiptera: Heteroptera: Pentatomidae) (Diptera: Anthomyiidae)
Soybean Aphid, Aphis glycines Matsumura Twospotted Spider Mite, Tetranychus urticae
(Hemiptera: Aphididae) Koch (Acari: Tetranychidae)
Spined Soldier Bug, Podisus maculiventris (Say) Two-Spotted Stink Bug, Perillus bioculatus
(Hemiptera: Pentatomidae) (Fabricius) (Hemiptera: Pentatomidae)
Spotted Cucumber Beetle or Southern Twostriped Grasshopper, Melanoplus bivittatus
Corn Rootworm, Diabrotica undecimpunc- (Say) (Orthoptera: Acrididae)
tata Mannerheim (Coleoptera: Varroa Mite, Varroa destructor Anderson &
Chrysomelidae) Truemann (Acari: Varroidae)
Spruce Budworms, Choristoneura Lederer Variegated Cutworm, Peridroma saucia (Hbner)
(Lepidoptera: Tortricidae) (Lepidoptera: Noctuidae)
Squash Bug, Anasa tristis (DeGeer) (Hemiptera: Vegetable Leafminer, Liriomyza sativae Blanchard
Coreidae) (Diptera: Agromyzidae)
Squash Vine Borer, Melittia cucurbitae (Harris) Viburnum Leaf Beetle, Pyrrhalta viburni
(Lepidoptera: Sesiidaae) (Paykull) (Coleoptera: Chrysomelidae)
Stable Fly, Stomoxys calcitrans (Linnaeus) Vine Mealybug, Planococcus ficus Signoret
(Diptera: Muscidae) (Hemiptera: Pseudococcidae)
Sweetpotato and Silverleaf Whiteflies, Bemisia Western Balsam Bark Beetle, Dryocoetes confusus
spp. (Hemiptera: Aleyrodidae) Swain (Coleoptera: Curculionidae,
Sweetpotato Flea Beetle, Chaetocnema confinis Scolytinae)
Crotch (Coleoptera: Chrysomelidae Western Corn Rootworm, Diabrotica virgifera
Alticinae) virgifera Leconte (Coleoptera:
Sweetpotato Weevil, Cylas formicarius (Fabricius) Chrysomelidae)
(Coleoptera: Brentidae) Western Grapeleaf Skeletonizer, Harrisina
Taiga Tick, Ixodes persulcatus Schulze (Acari: brillians Barnes & McDunnough
Ixodida: Ixodidae) (Lepidoptera: Zyganeidae)
Highlights of the Encyclopedia of Entomology

Western Harvester Ant, Pogonomyrmex occidentalis Winter Moth, Operophtera brumata (L.)
(Cresson) (Hymenoptera: Formicidae) (Lepidoptera: Geometridae) and Its
Western Thatching Ant, Formica obscuripes Biological Control
(Forel) (Hymenoptera: Formicidae) Wireworms, Several Genera and Species
Wheat Stem Sawflies: Cephus cinctus Norton, (Coleoptera: Elateridae)
Cephus pygmaeus (L.) and Trachelus tabidus Yellowstriped Armyworm, Spodoptera
(F.) (Hymenoptera: Cephidae) ornithogalli (Guene) (Lepidoptera:
White Grubs, Phyllophaga, and Others Noctuidae)
(Coleoptera: Scarabaeidae)
List of Contributors
Abivardi, Cyrus Gainesville, Florida 32611-0680
Institute of Integrative Biology USA
ETH Zurich
Universtt-Str.16 (CHN) Ajjan, Iskandar
CH-8092 Zurich University of Tishreen
Switzerland P.O. Box 740
Abou-Fakhr, Efat M. Syria
Department of Plant Sciences
Faculty of Agricultural and Food Sciences Ajlan, Aziz
American University of Beirut Department of Plant Protection
P.O. Box 11-0236 College of Agricultural and Food Sciences
Bliss Street King Faisal University
Beirut P.O. Box 55009
Lebanon Hofuf, Al-Hasa 31982
Saudi Arabia
Adams, Byron J.
Department of Microbiology and Molecular
Albajes, Ramon
Universitat de Lleida
Brigham Young University
Centre UdL-IRTA
Provo, Utah 84602
Rovira Roure 177
25199 Lleida
Adler, Peter
Department of Entomology
Clemson University
Box 340365 Alborn, Hans
114 Long Hall USDA, Agricultural Research Service
Clemson, South Carolina 29634-0365 Chemistry Research Unit
USA 1600-1700 SW 23rd Drive
Gainesville, FL, 32608
Agnello, Arthur M. USA
Department of Entomology
New York State Agricultural Experiment Station Aldryhim, Yousif
Cornell University P.O. Box 2460
Geneva, New York 14456-0462 Riyadh, 1141
USA Saudi Arabia

Agrios, George Alekseev, Andrey N.

Plant Pathology Department Russian Parasitological Society
University of Florida P.O. Box 738
List of Contributors

191186 Arif, Basil

St. Petersburg, D-186 Great Lakes Forestry Center
Russia 1219 Queen Street East
Sault Ste. Marie, Ontario P6A 5M7
All, John Canada
Department of Entomology
Armstrong, Earlene
University of Georgia
Department of Entomology
413 Biological Sciences Building
University of Maryland
Athens, Georgia 30602
4122 Plant Sciences Building
College Park, Maryland 20742
Alomar, scar
IRTA Arthurs, Steven
Centre de Cabrils Department of Entomology
Carretera de Cabrils s/n Biological Control Laboratory
08348 Cabrils Texas A&M University
Barcelona, Catalonia College Station, Texas 77843-2475
Spain USA

Asaro, Christopher
Alvarez, Juan M.
Department of Entomology
Aberdeen Research and Education Center
University of Georgia
University of Idaho
413 Biological Sciences Building
P.O. Box 870
Athens, Georgia 30602-2603
Aberdeen, Idaho 83210-0870
Attathom, Tipvadee
Amalin, Divina M. Department of Entomology
USDA-APHIS Kasetsart University
13601 Old Cutler Road Kamphaengsaen Campus
Miami, Florida 33158 Nakhon Pathom, 73140
USA Thailand

Amerasinghe, Felix P. Baldwin, Rebecca W.

International Management Institute Entomology and Nematology Department
127 Sunil Mawatha, Pelawatta P.O. Box 110620
Battaramulla, 10120 University of Florida
Sri Lanka Gainesville, Florida, 32611

Anderson, Robert Barbara, Kathryn A.

Canadian Museum of Nature Entomology and Nematology Department
P.O. Box 3443, Station D University of Florida
Ottawa, Ontario K1P 6P4 Gainesville, Florida 32611-0620
Canada USA
List of Contributors

Barfield, Carl Behmer, Spencer T.

Entomology and Nematology Department Department of Entomology
University of Florida Texas A&M University
Gainesville, Florida 32611-0620 College Station, Texas 77843-2475

Barrera, Juan F. Bellotti, Anthony C.

Departamento de Entomologa Tropical CIAT - Centro Internacional de Agricultura
El Colegio de la Frontera Sur Tropical
Tapachula, Chiapas, 30700 Apartado Areo 6713, Cali
Mxico Colombia
Barth, Martin
Beard, Charles E.
Friedrich-Miescher-Laboratorium der
Box 340315 - 114 Long Hall
Entomology, Soils and Plant Sciences
Spemannstr. 39
Clemson University
720726 Tbingen
Clemson, South Carolina 29634-0315
Baumgrtner, Johann
Bennett, Gary W.
International Centre of Insect Physiology and
Center for Urban and Industrial Management
Ecology (ICIPE)
Department of Entomology
P.O. Box 17319
Purdue University
Addis Ababa
1158 Smith Hall
West Lafayette, Indiana 47907-1158
Baz, Arturo USA
Universidad de Alcala
Department Biologia Animal Bentz, Barbara J.
Alcala de Henares Rocky Mountain Research Station
Madrid, E-28801 Forestry Sciences Lab
Spain 860 N 1200 E
Logan, Utah 84321
Becnel, James J. USA
Center for Medical, Agricultural and Veterinary
Entomology Berisford, C. Wayne
USDA-ARS Department of Entomology
Gainesville, Florida 32611-0970 University of Georgia
USA 413 Biological Sciences Building
Athens, Georgia 30602
Beeman, Richard W. USA
GMPRC Berlinger, Menachem J.
1515 College Avenue Entomology Laboratory
Manhattan, Kansas Agricultural Research Organization
USA Gilat Regional Experiment Station
List of Contributors

P.O. Box 7710 B-1000 Bruxelles

Beer-Sheva, 84843 Belgium
Borgemeister, Christian
Bernays, Elizabeth A. Institute of Plant Diseases and Plant Protection
Department of Entomology University of Hannover
University of Arizona Herrenheuser Str. 2
P.O. Box 2100 30419 Hannover
Tucson, Arizona 85721-0036 Germany
Bostanian, Noubar J.
Berry, Colin
Horiticultural Research and Development
Cardiff School of Biosciences
Cardiff University
Agriculture and Agri-Food Canada
P.O. Box 911
430, Boulevard Gouin
Museum Avenue
St.-Jean-sur-Richelieu, Quebec J3B 3E6
Cardiff, Wales CF10 3US

Boucher, Stphanie
Bidochka, Michael
Lyman Entomological Museum
Department of Biological Sciences
McGill University, Macdonald Campus
Brock University
Ste-Anne-de-Bellevue, Qubec H9X 3V9
St. Catharines, Ontario

Blackwell, Alison Boucias, Drion

Center for Tropical Veterinary Medicine Entomology and Nematology Department
University of Edinburgh University of Florida
Easter Bush Veterinary Centre Gainesville, Florida 32611-0620
Roslin, Midlothian EH25 9RG USA
Bowles, David E.
Blum, Murray S. Resource Protection Division
Department of Entomology Freshwater Conservation Branch/River Studies
University of Georgia Program
413 Biological Sciences Building Texas Parks and Wildlife Department
Athens, Georgia 30602 P.O. Box 1685
USA San Marcos, Texas 78666
Boev, Jean-Luc
Dpartement dEntomologie Brambila, Julieta
IRSNB-KBIN Florida Department of Agriculture and
Royal Belgian Institute of Natural Sciences Consumer Services
Rue Vautier 29 Division of Plant Industry
List of Contributors

1911 SW 34th Street Burden, Beverly

Gainesville, Florida 32614 Department of Biological Sciences
USA Louisiana State University - Shreveport
One University Place
Brandenburg, Rick Shreveport, Louisiana 71115
Department of Entomology USA
North Carolina State University
P.O. Box 7613 Buss, Eileen A.
Raleigh, North Carolina 27695-7613 Entomology and Nematology Department
USA University of Florida
Gainesville, Florida 32611-0620
Brault, Aaron C.
Centers for Disease Control and Prevention
National Center for Infectious Diseases
Division of Vector-Borne Infectious Diseases Byers, George W.
P.O. Box 2087 Snow Entomological Division
Foothills Campus Natural History Museum
Fort Collins, Colorado 80522 University of Kansas
USA 1460 Jayhawk Boulevard
Lawrence, Kansas 66045-7523
Brewer, J. Wayne USA
Department of Entomology
Auburn University Cabana, Jean
301 Funchess Hall AFA Environment Inc.
Auburn University, Alabama 36849-5413 1100, Rene-Levesque Boulevard West
USA 25th Floor
Montreal, Quebec H3B 5C9
Brown, Harley P. Canada
Department of Zoology
University of Oklahoma Caceres, Carlos
730 Van Vleet Oval, Room 314 FAO/IAEA
Norman, Oklahoma 73019-6121 Agricultural and Biotechnology Laboratory
USA A-2444 Seibersdorf
Brown, Susan J.
Division of Biology
Kansas State University Calabrese, Diane M.
9 Anderson Hall 1000 Robin Road
Manhattan, Kansas 66506 Silver Spring, Maryland 20901-1873

Broza, Meir Cane, James H.

University of Haifa USDA-ARS
Oranim, Tivon 36006 Bee Biology and Systematics Laboratory
Israel Utah State University
List of Contributors

5310 Old Main Hill Cherry, Ron

Logan, Utah 84322-5310 Everglades Research and Education Center
USA University of Florida
3200 E. Palm Beach Road
Capinera, John L. Belle Glade, Florida 33430
Entomology and Nematology Department USA
University of Florida
Gainesville, Florida 32611-0620 Chiappini, Elisabetta
USA Istituto di Entomologia e Patologia Vegetale
Facolt di Agraria
Cardwell, Kitty F. Universit Cattolica del Sacro Cuore
USDA/CSREES Via Emilia Parmense, 84
Washington, D.C. Piacenza, 29100
USA Italy

Cave, Ronald D.
Choate, Paul M.
Entomology and Nematology Department
Entomology and Nematology Department
University of Florida
University of Florida
Indian River Research and Education Center
Gainesville, Florida 32611-0620
Ft. Pierce, Florida 34945-3138
Chouinard, Grald
Cdola, Claudia V.
Institut de Recherche et de Dveloppement en
Universidad Nacional de La Plata
3300 Sicotte
Consejo Nacional de Investigaciones Cientficas y
Saint-Hyacinthe, Quebec J2S 7B8
2 No 584 (1900)
La Plata
Argentina Chow, Yien-Shing
National Museum of Natural Science
Chapman, Reg 1, Kuan Chien Road
Arizona Research Laboratories Taichung 404, Taiwan
Division of Neurobiology Republic of China
University of Arizona
P.O. Box 210077 Christian, Peter
Tuscon, Arizona 85721-0077 National Institute for Biological Standards and
USA Control
Potters Bar, Hertfordshire
Cheng, Lanna UK
Scripps Institution of Oceanography
University of California, San Diego Cilek, James E.
9500 Gilman Drive Biting Fly and Tick Control Section
LaJolla, California 92093-0202 John A. Mulrennan, Sr. Public Health Entomology
USA Research and Education Center
List of Contributors

Florida A&M University Cranshaw, Whitney

4000 Frankford Avenue Department of Bioagricultural Sciences and Pest
Panama City, Florida 32405-1933 Management
USA Colorado State University
Fort Collins, Colorado 80523-0001
Clement, Stephen L. USA
Western Regional Plant Introduction Station Cresswell, James
Washington State University School of Biological Sciences
59 Johnson Hall University of Exeter
P.O. Box 646402 Hather Laboratories
Pullman, Washington 99164-6402 Prince of Wales Road
USA Exeter, EX4 4PS
Collier, Rosemary H.
Horticulture Research International Crist, Thomas O.
Wellesbourne, Warwick CV35 9EF Department of Zoology
UK Miami University
Pearson Hall Room 212
Conlong, Des E. Oxford, Ohio 45056-1400
SASA Experiment Station USA
Private Bag X02
Mount Edgecombe Crooker, Allen
Kwa Zulu-Natal, 4300 Biology Department
South Africa Hartwick College
Oneonta, New York 13820
Constantino, Reginaldo USA
Depto de Zoologia
Universidade de Braslia Cuda, James P.
70910-900 Braslia, DF Entomology and Nematology Department
Brazil University of Florida
Gainesville, Florida 32611-0620
Coons, Lewis B. USA
Integrated Microscopy Center
University of Memphis Cumming, Jeffrey M.
201 Life Sciences Building Canadian National Collection of Insects
Rooms LS 101-113 Arachnids and Nematodes
Memphis, Tennessee 38152-6040 Agriculture and Agri-Food Canada
USA K.W. Neatby Building, C.E.F.
Ottawa, Ontario K1A 0C6
Costas, MIguel Canada
Facultad de Biologa
Universidad Complutense de Madrid Cushing, Paula E.
28040 Madrid Department of Zoology
Spain Denver Museum of Nature and Science
List of Contributors

2001 Colorado Boulevard Danoff-Burg, James A.

Denver, Colorado 80205-5798 Department of Ecology, Evolution and
USA Environmental Biology
Columbia University
Cusson, Michel 1020 Schermerhorn Extension, MC5557
Natural Resources Canada 1200 Amsterdam Avenue
Canadian Forest Service New York, New York 10027
1055 rue du P.E.P.S. USA
P.O. Box 3800
Sainte-Foy, Quebec G1V 4C7 Davidson, Diane W.
Canada Department of Biological Sciences
University of Utah
257 South, 1400 East
Czl, Gyozo
Salt Lake City, Utah 84112-0840
Hungarian Natural History Museum
Hungary DeClercq, Patrick
Laboratory of Agrozoology, Department of Crop
Daane, Kent M. Protection
Division of Insect Biology Faculty of Agricultural and Applied Biological
ESPM Sciences
University of California Ghent University
Berkeley, California 94720 Coupure Links 653
USA B-9000 Ghent
Dam, Nicole M. van
Denell, Robin E.
Netherlands Institute of Ecology
Division of Biology
Centre for Terrestrial Ecology
Kansas State University
P.O. Box 40
9 Anderson Hall
6666 ZG Heteren
Manhattan, Kansas 66506
The Netherlands

Dame, David Denmark, Harold A.

Entomology and Nematology Department Florida Department of Agriculture and
University of Florida Consumer Services
Gainesville, Florida 32611-0620 Division of Plant Industry
USA 1911 SW 34th Street
Gainesville, Florida 32614
Daniels, Jaret C. USA
Entomology and Nematology Department
University of Florida Devorshak, Christina
Gainesville, Florida 32611-0620 USDA-APHIS-PPQ
USA Center for Plant Health Science and Technology
List of Contributors

Plant Epidemiology and Risk Analysis Laboratory 1200 University Street

Raleigh, North Carolina Spearfish, South Dakota 57799-9003

Dicke, Marcel Dunford, James, C.

Laboratory of Entomology Department of Entomology and
Wageningen University Nematology
P.O. Box 8031 University of Florida
6700EH Wageningen Gainesville, Florida, 32611
The Netherlands USA

Dietrich, C.H.
Center for Biodiversity Dyby, Susanne D.
Illinois Natural History Survey Res. Le Rambouillet Bat. B
607 East Peabody Drive 1 Alle des Lucioles
Champaign, Illinois 61820 Beaulieu-sur-Mer, 06310
USA France

Dorchin, Netta Easton, Emmett

Museum Koenig Department of Plant and Environmental
Adenauerallee 160 Protection Sciences
53113 Bonn University of Hawaii at Manoa
Germany 3050 Maile Way
Gilmore Hall 310
Downer, R.A. Honolulu, Hawaii 96822-2271
Laboratory for Pest Control Application USA
Ohio Agricultural Research and Development
Ebert, Timothy A.
Laboratory for Pest Control Application
Ohio State University
1680 Madison Avenue
Ohio Agricultural Research and Development
Wooster, Ohio 44691-4096
Ohio State University
1680 Madison Avenue
Downie, D.A.
Wooster, Ohio 44691-4096
Department of Zoology and Entomology
Rhodes University
Grahamstown 6140
South Africa Edmunds, Malcolm
Department of Environmental
Downing, Holly Management
Unit 9003 University of Central Lancashire
College of Arts and Sciences Preston, PR1 2HE
Black Hills State University UK
List of Contributors

Ellis, Amanda Felicioli, Antonio

Florida Department of Agriculture and Universita degli Studi di Pisa
Consumer Services Dipartimento di Anatomia, Biochimica e
Division of Plant Industry Fisiologia veterinaria
P.O. Box 147100 Viale delle Piagge n 2, 56100 Pisa
Gainesville, Florida 32614-7100 Italy
Feng, Qili
Ellis, James D. Great Lakes Forestry Center
Entomology and Nematology Department 1219 Queen Street East
University of Florida Sault Ste. Marie, Ontario P6A 5M7
Gainesville, Florida 32611-0620 Canada

Emmel, Thomas C. Finch, Stan

McGuire Center for Lepidoptera Research and Horticulture Research International
Conservation Wellesbourne, Warwick CV35 9EF
University of Florida UK
Gainesville, Florida 32611-0620
USA Finke, Mark D.
6811 Horned Owl Trail
Epsky, Nancy D. Scottsdale, Arizona 85262-8519
Subtropical Horticulture Research Station
13601 Old Cutler Road Fishel, Fred
Miami, Florida 33158 University of Florida,
USA Pesticide Information Office
PO Box 110710
Farag, Mohamed A. Gainesville, Florida 32611-0710
Department of Chemistry and Biochemistry USA
Texas Tech University
Lubbock, Texas 79409-1061
Fitzgerald, T.D.
Department of Biology
Fasulo, Thomas R. State University of New York at Cortland
Entomology and Nematology Department Cortland, New York 13045
University of Florida USA
Gainesville, Florida 32611-0620
USA Floate, Kevin D.
Agriculture and Agri-Food Canada
Feir, Dorothy Lethbridge Research Centre
Biology Department 5403 1st Avenue
St. Louis University P.O. Box 3000
St. Louis, Missouri 63103 Lethbridge, Alberta T1J 4B1
USA Canada
List of Contributors

Flowers, R. Wills Rondebosch 7700

Center for Biological Control/Entomology Cape Town
Division of Agricultural Sciences Republic of South Africa
Florida A&M University
Tallahassee, Florida 32307 Galante, Eduardo
University of Alicante
Fornasari, Luca 03080 San Vicente del Raspeig
Clas de lErmitage Alicante
636, ave. E. Jeanbrau Spain
34090 Montpellier
France Gall, Lawrence F.
Peabody Museum of Natural History
Frank, J. Howard Yale University
Entomology and Nematology Department P.O. Box 208118
University of Florida New Haven, Connecticut 06520-8118
Gainesville, Florida 32611-0620 USA
Gangwere, Stan K.
Department of Biological Sciences
Franz, Gerald
Wayne State University
Detroit, Michigan 48202
Agricultural and Biotechnology Laboratory
A-2444 Seibersdorf
Garcia, Lloyd
North Carolina Department of Agriculture and
Froeba, Jason G. Consumer Sciences
Entomology and Nematology Department Plant Industry Division
University of Florida P.O. Box 27647
Gainesville, FL 32611-0620 Raleigh, North Carolina 27611

Gabrys, Beata Gayubo, Severiano F.

Institute of Biotechnology and Environmental Area de Zoologa
Sciences Facultad de Biologa
University of Zielona Gora Universidad de Salamanca
Monte Cassino 21b 37071 Salamanca
65-561 Zielona Gora Spain
Gerberg, Eugene J.
Gde, Gerd Entomology and Nematology Department
Zoology Department University of Florida
University of Cape Town Gainesville, Florida 32611-0620
Private Bag USA
List of Contributors

Gerlach, Gnter Goebel, Rgis

Botanischer Garten Mnchen - Nymphenburg SASEX-CIRAD
Menzinger Str. 65 Entomology Department
80638 Mnchen Private Bag X02
Germany Mount Edgecombe, 4300
South Africa
Gerling, Dan
Department of Zoology Goettel, Mark
Tel Aviv University Lethbridge Research Centre
Ramat Aviv, 69978 Agriculture and Agri-Food Canada
Israel 5403 1st Avenue South
P.O. Box 3000
Lethbridge, Alberta T1J 4B1
Ghiradella, Helen Canada
Department of Biology
State University of New York at Albany Gold, Clifford S.
Albany, New York 12222 CIAT (International Center for Tropical
USA Agriculture)
P.O. Box 6247
Giberson, Donna Kampala
Department of Biology Uganda
University of Prince Edward Island
550 University Ave. Goodman, Katie
Charlottetown, PEI C1A 4P3 Department of Biological Sciences
Canada Western Illinois University
Macomb, Illinois 61455
Giblin-Davis, Robin M. USA
Fort Lauderdale Research and Education Center
University of Florida Goula, Marta
3205 College Avenue University of Barcelona
Davie, Florida 33314-7719 Barcelona
USA Spain

Greco, Nancy M.
Gibson, G.A.P. CEPAVE
Agriculture and Agri-Food Canada Universidad Nacional de La Plata
960 Carling Avenue Consejo Nacional de Investigaciones Cientficas y
Ottawa, Ontario K1A 0C6 Tcnicas
Canada 2 No 584 (1900)
La Plata
Glenn, D. Michael Argentina
USDA-ARS, Appalachian Fruit Research Station
2217 Wiltshire Rd. Grewal, Parwinder S.
Kearneysville, WV 25430 Department of Entomology
USA The Ohio State University
List of Contributors

Columbus, Ohio 43210 Hall, Donald W.

USA Entomology and Nematology Department
University of Florida
Gruner, Susan V. Gainesville, Florida 32611-0620
Entomology and Nematology Department USA
University of Florida
Gainesville, Florida 32611-0620
Hall, Jason P.W.
National Museum of Natural History
Guertin, Claude Smithsonian Institution
INRS Washington, D.C. 20560-0127
Institut Armand-Frappier USA
531 Boulevard des Prairies
Laval, Quebec H7V 1B7 Hall, Robert D.
Canada Office of Research
University of Missouri
Gupta, Virendra K. 205 Jesse Hall
Entomology and Nematology Department Columbia, Missouri 65211
University of Florida USA
Gainesville, Florida 32611-0620
Handler, Alfred M.
Gwynne, Darryl T. Center for Medical, Agricultural and Veterinary
University of Toronto at Mississauga Entomology
Mississauga, Ontario L5L 1C6 USDA-ARS
Canada P.O. Box 14565
1700 SW 23rd Drive
Habeck, Dale Gainesville, Florida 32608
Entomology and Nematology Department USA
University of Florida
Gainesville, Florida 32611-0620 Hangay, George
USA 80 Gondola Road
Narrabeen, New South Wales 2101
Hahn, Daniel A.
Entomology and Nematology Department
University of Florida
Gainesville, Florida 32611-0620 Hansen, Laurel D.
USA Department of Biology
Spokane Falls Community College
Halbert, Susan 3410 W. Fort George Wright Dr.
Florida Department of Agriculture and Spokane, WA 99224-5288
Consumer Services USA
Division of Plant Industry
1911 SW 34th Street Harman, Dan M.
Gainesville, Florida 32614 University of Maryland
USA Center for Environmental Science
List of Contributors

Appalachian Laboratory Hernandez, Santiago

Frostburg, Maryland 21532-2307 Universities of Cordoba and Extremadura
USA Cceres, 10071
Haunerland, Norbert H.
Department of Biological Sciences Hilje, Luko
Simon Fraser University Plant Protection Unit
Burnaby, British Columbia V5A 1S6 7170 CATIE
Canada Turrialba, Cost Rica

Headings, Mark Hinkle, Nancy C.

Agricultural Technical Institute Department of Entomology
Ohio State University University of Georgia
1328 Dover Road 413 Biological Sciences Building
Wooster, Ohio 44691-8905 Athens, Georgia 30602

Heath, Allen
Hirsch, Helmut V.B.
AgResearch Wallaceville
Department of Biology
Wallaceville Animal Research Centre
State University of New York at Albany
PO Box 40063, Upper Hutt, 5140
1400 Washington Avenue
New Zealand
Albany, New York 12222
Held, David W.
Entomology Department
University of Kentucky Ho, Chyi-Chen
S-225 Agricultural Science Center Taiwan Agricultural Research Institute
BuildingNorth Department of Applied Zoology
Lexington, Kentucky 40546-0091 Wufeng
USA Taichung, Taiwan
Republic of China
Henneman, M.L.
Department of Biological Sciences Hodges, Greg S.
University of Bristol Florida Department of Agriculture and
Bristol, BS8 1TH Consumer Services
UK Division of Plant Industry
1911 SW 34th Street
Heppner, John B. Gainesville, Florida 32614
Florida State Collection of Arthropods USA
Florida Department of Agriculture and
Consumer Services Horton, David R.
Division of Plant Industry USDA/ARS
P.O. Box 147100 5230 Konnowac Pass Road
Gainesville, Florida 32614-7100 Wapato, Washington 98951
List of Contributors

Hou, Roger F. Hadassah Medical School

Department of Entomology Jerusalem
National Chung Hsing University Israel
Taichung 402, Taiwan
Republic of China Isman, Murray B.
Faculty of Agricultural Sciences
Howard, Forrest W. University of British Columbia
Fort Lauderdale Research and Education Center 248-2357 Main Mall
University of Florida Vancouver, British Columbia V6T 1Z4
Ft. Lauderdale, Florida 33314-7719 Canada

Hoy, Marjorie A. James, Rosalind R.

Entomology and Nematology Department USDA/ARS, Bee Biology and Systematics
University of Florida Laboratory
Gainesville, Florida 32611-0620 Department of Biology, UMC 5310
USA Utah State University
5310 Old Main Hill
Huber, John Logan, Utah 84321-5310
Canadian Forestry Service USA
960 Carling Avenue
Ottawa, Ontario K1A 0C6 Jaffe, Klaus
Canada Universidad Simn Bolvar
Apartado 89000, Caracas 1080
Hunter, Fiona F. Venezuela
Department of Biological Sciences
Brock University Jeremas, Xavier
St. Catharines, Ontario L2S 3A1 Museum of Natural Sciences
Canada C./ Catalunya 28
08758 Cervell (prov. Barcelona)
Hurd, Hilary Spain
Keele University
School of Life Sciences
Keele, Staffordshire ST55BG Jolivet, Pierre
UK 67, Bd Soult
75012 Paris
Hurd, Lawrence E. France
Department of Biology
Washington & Lee University Jurez, M. Patricia
Lexington, Virginia 24450 Facultad de Ciencias Mdicas
USA Universidad Nacional de la Plata
Calle 60 y 120
Ioffe-Uspensky, Inna C.C. 455
Department of Parasitology La Plata, 1900
The Hebrew University of Jerusalem Argentina
List of Contributors

Kabissa, Joe C.B. Private Bag X134

Tanzania Cotton Lint and Seed Board Pretoria, 0001
Pamba House South Africa
Garden Avenue
P.O. Box 9161 Khan, Zeyour R.
Dar Es Salaam ICIPE
Tanzania P.O.B. 30772
Kalkar, . Kenya
Department of Entomology
Clemson University Kima, Peter E.
Clemson, South Carolina Microbiology and Cell Science Department
USA University of Florida
Building 981, Box 110700
Katsoyannos, Byron Gainesville, Florida 32611-0700
Aristotle University of Thessaloniki USA
Department of Agriculture
Laboratory of Applied Zoology and Parasitology Klassen, Waldemar
Thessaloniki, 541 24 Tropical Research and Education Center
Greece University of Florida
18905 SW 280th Street
Kathirithamby, Jeyaraney Homestead, Florida 33031-3314
Department of Zoology USA
Oxford University
South Parks Road Klotz, John H.
Oxford OX1 3PS, Department of Entomology
UK University of California
Riverside, California
Keeley, Larry L. USA
Department of Entomology
Texas A&M University Klowden, Marc J.
2475 TAMU Division of Entomology
College Station, Texas 77843-2475 University of Idaho
USA Moscow, Idaho 83844-2339
Kerr, Peter H.
California Department of Food and Agriculture Koehler, Philip G.
Plant Pest Diagnostics Branch Entomology and Nematology Department
3294 Meadowview Rd. University of Florida
Sacramento, California 95832 Gainesville, Florida 32611-0602

Kfir, Rami Kok, L.T.

ARC-Plant Protection Research Institute Department of Entomology
Rietondale Research Station College of Agriculture and Life Sciences
600 Soutpansberg Road Virginia Polytechnic Institute and State University
List of Contributors

216 Price Hall Michigan State University

Blacksburg, Virginia 24061-0319 204 Center for Integrated Plant Systems
USA East Lansing, Michigan 48824-1311
Kondratieff, Boris C.
Department of Bioagricultural Sciences and Pest Lawrence, Pauline O.
Management Entomology and Nematology Department
Colorado State University University of Florida
Fort Collins, Colorado 80523 Gainesville, Florida 32611-0620

Kouloussis, Nikos A. Leather, Simon R.

Aristotle University of Thessaloniki Division of Biology
Laboratory of Applied Zoology and Parasitology Imperial College London
Thessaloniki, 541 24 Silwood Park Campus
Greece Ascot, SL5 7PY
Krafsur, Elliot S.
206 Hidden Valley Circle Lebiush-Mordechi, Sarah
Shepherdstown, West Virginia 25443 Entomology Laboratory
USA Agricultural Research Organization
Gilat Regional Experiment Station
Lacey, Lawrence A. P.O. Box 7710
USDA-ARS Beer-Sheva, 84843
Yakima Agricultural Research Lab Israel
5230 Konnowac Pass Road
Wapato, Washington 98951 Lecoq, Michel
USA Centre de Coopration Internationale en Recher-
che Agronomique pour le Dveloppement
Lambdin, Paris L. Prifas
Department of Entomology and Plant Pathology TA 40/D
University of Tennessee 34398 Montpellier, Cedex 5
2431 Center Drive France
205 Ellington Plant Sciences Building
Knoxville, Tennessee 37996-4560 Lee, Clarence M.
USA Department of Biology
Howard University
Lapointe, Stephen L. 415 College Street N.W.
USDA-ARS, U.S. Horticultural Research Lab Washington, D.C. 20059
2001 South Rock Road USA
Fort Pierce, Florida 34945
USA Lee, How-Jing
Department of Entomology
Landis, Douglas A. National Taiwan University
Department of Entomology Taipei 106, Taiwan
Center for Integrated Plant Systems Republic of China
List of Contributors

Legaspi, Benjamin C., Jr. Lloyd, James E.

USDA-ARS, FAMU-Center for Biological Entomology and Nematology Department
Control University of Florida
6383 Mahan Drive Gainesville, Florida 32611-0620
Tallahassee, Florida 32308 USA
Long, Lewis S.
Legaspi, Jesusa C. Entomology and Nematology Department
USDA-ARS, FAMU-Center for Biological University of Florida
Control Gainesville, Florida 32611-0620
6383 Mahan Drive USA
Tallahassee Florida 32308
USA Lord, Cynthia C.
Florida Medical Entomology Laboratory
Legg, David University of Florida
Department of Renewable Resources 200 9th Street S.E.
College of Agriculture Vero Beach, Florida 32962-4699
University of Wyoming USA
P.O. Box 3354
Laramie, Wyoming 82071 Lord, Jeffrey
Northern Plains Area Grain Marketing and
Leppla, Norman C.
Production Research Center
Entomology and Nematology Department
Biological Research Unit
University of Florida
1515 College Avenue
Gainesville, Florida 32611-0620
Manhattan, Kansas 66502-2796
Leskey, Tracy
USDA-ARS Lounibos, L. Philip
Appalachian Fruit Research Station Florida Medical Entomology Laboratory
45 Wiltshire Road University of Florida
Kearneysville, West Virginia 25430 200 9th Street S.E.
USA Vero Beach, Florida 32962-4699
Liburd, Oscar
Entomology and Nematology Department Luna, Mara G.
University of Florida Department of Ecology and Evolutionary Biology
Gainesville, Florida 32611-0620 University of California at Irvine
USA 321 Steinhaus Hall
Irvine, California 92697-2525
Liebman, Matt USA
Department of Agronomy
Iowa State University Lysyk, Tim
Ames, Iowa 50011-1010 Laboratory of Vector Ecology
USA Agriculture and Agri-Food Canada
List of Contributors

P.O. Box 3000 1600/1700 SW 23rd Drive

5403 1st Avenue South P.O. Box 14565
Lethbridge, Alberta T1J 4B1 Gainesville, Florida 32604
Canada USA

MacVean, Charles Manley, Donald G.

Instituto de Investigaciones PeeDee Research and Education Center
Universidad del Valle de Guatemala Clemson University
Aptdo. Postal 82 2200 Pocket Road
Guatemala City Florence, South Carolina 29506-9706
Guatemala USA

Magalhes, Bonifcio Mapes, Carol C.

EMBRAPA Recursos Genticos e Biotecnologia Biology Department
Parque Estao Biolgica Kutztown University
Final WS Norte Kutztown, Pennsylvania 19530
CEP 70770-900 USA
Marcos-Garcia, Maria Angeles
Maimala, S.
University of Alicante
Department of Agriculture
03080 San Vicente del Raspeig
Ministry of Agriculture and Cooperation
Chatuchak, Bangkok

Marshall, David Martnez, Javier

Ecology and Evolutionary Biology Universities of Cordoba and Extremadura
University of Connecticut Cceres, 10071
75 N. Eagleville Rd., U-3043 Spain
Storrs, Connecticut 06269
USA Martnez, Maricela
Instituto Mexicano de Tecnologa del Agua
Malakar-Kuenen, Raksha (IMTA)
Division of Insect Biology Paseo Cuaunhahuac 8532 Progreso
ESPM Jiutepec, Morelos
University of California Mexico
Berkeley, California 94720
USA Mason, Peter G.
Eastern Cereal and Oilseed Research Centre
Mankin, Richard Agriculture and Agri-Food Canada
USDA/ARS 960 Carling Avenue
Center for Medical, Agricultural and Veterinary Ottawa, Ontario K1A 0C6
Entomology Canada
List of Contributors

Matthews, Deborah L. McSorley, Robert

Department of Entomology and Entomology and Nematology Department
Nematology University of Florida
University of Florida Gainesville, Florida 32611-0620
Gainesville, Florida 32611 USA
Meagher, Robert
Matsumoto, Yoshiharu USDA-ARS
University of Tokyo Center for Medical and Veterinary Entomology
Tokyo 1700 SW 23rd Drive
Japan Gainesville, Florida 32608-1069
Matthiessen, John N.
CSIRO Entomology Medal, Julio
Underwood Avenue Entomology and Nematology Department
Floreat, Western Australia 6014 University of Florida
Australia Gainesville, Florida 32611-0620
McAuslane, Heather J.
Entomology and Nematology Department Meinke, Lance J.
University of Florida Department of Entomology
Gainesville, Florida 32611-0620 University of Nebraska
USA Lincoln, Nebraska 68583-0816
McCravy, Kenneth W.
Department of Biological Sciences Meinking, Terri L.
Western Illinois University Department of Dermatology
1 University Circle University of Miami School of Medicine
Macomb, Illinois 61455 1600 NW 10 Avenue
USA P.O. Box 016960 (R-117)
Miami, Florida 33101
McFadyen, Rachel USA
CRC Australian Weed Management
Natural Resource Sciences Menalled, Fabin D.
Meiers Road Department of Agronomy
Indooroopilly, Queensland Iowa State University
Australia Agronomy Hall
Ames, Iowa 50011-1010
McIver, James USA
Forestry and Range Sciences Laboratory
Pacific Northwest Research Station Merkl, Ott
USDA Forest Service Hungarian Natural History Museum
1404 Gekeler lane H-1088 Budapest
LaGrande, Oregon 97850 Baross utca, 13
USA Hungary
List of Contributors

Meyer, Jason Sakuragaoka 1-1-1

Entomology and Nematology Department Setagaya-ku
University of Florida Tokyo, 156-8502
Gainesville, Florida 32611-0620 Japan
Mizell, Patricia A.
Meyerdirk, Dale E. North Florida Research and Education Center
USDA APHIS, PPQ University of Florida
National Biological Control Institute 155 Research Road
4700 River Road Quincy, Florida 32351
Unit 135 USA
Riverdale, Maryland 20737-1228
Mizell, Russell F., III
Miller, Barry R. North Florida Research and Education Center
Centers for Disease Control and Prevention University of Florida
National Center for Infectious Diseases 155 Research Road
Division of Vector-Borne Infectious Diseases Quincy, Florida 32351
P.O. Box 2087 USA
Foothills Campus
Fort Collins, Colorado 80522 Moriya, Seiichi
USA Insect Ecology Laboratory
National Agricultural Research Center
Miller, Dini M. 3-1-1, Kannondai
Department of Entomology Tsukuba, Ibaraki 305-8666
Virginia Tech University, 216A Price Hall Japan
Blacksburg, Virginia 24061
USA Morrill, Wendell L.
Department of Entomology
Miller, Laura T. Montana State University
West Virginia Department of Agriculture Bozeman, Montana 59717-0001
Plant Industries Division USA
1900 Kanawha Boulevard, East
Charleston, West Virginia 25305-0191
Moya-Raygoza, Gustavo
Universidad de Guadalajara
Mitcham, Elizabeth Departamento de Botnica y Zoologa
Department of Plant Sciences, Mail Stop 2 Apartado Postal 139
University of California Zapopan, Jalisco 45101
One Shields Avenue Mexico
Davis, California 95616
USA Mutun, Serap
Abant Izzet Baysal University
Mitsuhashi, Jun Department of Biology
Department of Bioscience Bolu, 14280
Tokyo University of Agriculture Turkey
List of Contributors

Nadel, Hannah Nayar, Jai K.

USDA-ARS SJVASC Florida Medical Entomology Laboratory
San Joaquin Valley Science Center University of Florida
9611 Riverbend Ave. 200 9th Street SE
Parlier, California 93648 Vero Beach, Florida 32962

Nagoshi, Rod Nealis, Vince

USDA-ARS Natural Resources Canada-Canadian Forest
Center for Medical, Agricultural and Veterinary Service
Entomology Pacific Forestry Centre
P.O. Box 14565 506 W. Burnside Road
Gainesville, Florida 32608 Victoria, British Columbia V8Z 1M5
USA Canada

Napper, Emma Negron, Jose F.

Rothamsted USDA Forest Service
Harpenden, Hertfordshire AL5 2JQ Rocky Mountain Research Station
UK 240 West Prospect
Fort Collins, Colorado 80526
Naranjo, Steven E. USA
Neuenschwander, Peter
Western Cotton Research Laboratory
International Institute of Tropical Agriculture
4135 E. Broadway Road
08 B.P. 0932
Phoenix, Arizona 85040
Nation, James L. Neven, Lisa, G.
Entomology and Nematology Department USDA-ARS, Yakima Agricultural Research
University of Florida Laboratory
Gainesville, Florida 32611-0620 5230 Konnowac Pass Road
USA Wapato, WA 98951 USA

Navajas, Maria Nguyen, Khuong

Institut National de la Recherche Agronomique Entomology and Nematology Department
Campus International de Baillarguet University of Florida
CS 30016 Gainesville, Florida 32611-0620
Montferrier sur Lez cedex, 34988 USA
Nicolson, Sue
Navarette, Ignacio Department of Zoology & Entomology
Universities of Cordoba and Extremadura University of Pretoria
Cceres, 10071 Pretoria 0002
Spain South Africa
List of Contributors

Noling, Joseph W. Okuda, Takashi

University of Florida National Institute of Agrobiological
Citrus Research & Education Center Sciences
700 Experiment Station Rd. Anhydrobiosis Research Unit, 1-2 Ohwashi
Lake Alfred, Florida 33850 Tsukuba, 305-8634
USA Japan

Norris, Douglas E. ONeill, Kevin M.

Johns Hopkins University Department of Land Resources and
Bloomberg School of Public Health Environmental Sciences
W. Harry Feinstone Department of Molecular Montana State University
Microbiology and Immunology Bozeman, MT 59717
615 North Wolfe Street Rm E5008 USA
Baltimore, Maryland 21205
USA Orphanides, George M.
Agricultural Research Institute
Northfield, Tobin P.O. Box 22016
University of Florida Nicosia, 1516
North Florida Research and Education Center Cyprus
155 Research Rd.
Otis, Gard W.
Quincy, Florida 32351
Department of Environmental Biology
University of Guelph
Guelph, Ontario N1G 2W1
OBrien, Lois B.
College of Engineering Sciences, Technology and
Overholt, William Allan
Florida A&M University
Indian River Research and Education Center
Tallahassee, Florida 32307-4100
University of Florida
Fort Pierce, Florida
OHara, James, E.
Agriculture and Agri-Food Canada Panizzi, Antnio R.
960 Carling Avenue Embrapa Soja
Ottawa, Ontario K1A 0C6 Caixa Postal 231
Canada Londrina, PR 86001-970
Oi, David H.
USDA-ARS Papadopoulos, Nikos T.
Center for Medical, Agricultural and Veterinary Aristotle University of Thessaloniki
Entomology Laboratory of Applied Zoology and
1600 SW 23rd Drive Parasitology
Gainesville, Florida 32608-1067 Thessaloniki, 541 24
USA Greece
List of Contributors

Parra, Jos Roberto Postali P.O. Box 443051

Departamento de Entomologia Moscow, Idaho 83844-3051
Fitopatologia e Zoologia Agrcola. USA
Esalq/USP, C.P. 9 - 13418-900
Piracicaba, SP Pea, Jorge E.
Brazil Tropical Research and Education Center
University of Florida
Par, Paul W. 18905 SW 280th Street
Department of Chemistry and Biochemistry Homestead, Florida 33031-3314
Texas Tech University USA
Lubbock, Texas 79409-1061
USA Pendland, J.C.
Entomology and Nematology Department
Paulson, Gregory S. University of Florida
Shippensburg University Gainesville, Florida 32611-0620
Department of Biology USA
Shippensburg, Pennsylvania 17257
USA Pereyra, Patricia C.
Peairs, Frank B. Universidad Nacional de La Plata
Department of Bioagricultural Sciences and Pest Consejo Nacional de Investigaciones Cientficas y
Management Tcnicas
Colorado State University 2 No 584 (1900)
Fort Collins, Colorado 80523-1177 La Plata
USA Argentina

Peck, Daniel C. Philogne, Bernard J.R.

Department of Entomology, Barton Laboratory Department of Biology
New York State Agricultural Experiment Station University of Ottawa
Cornell University Poste 4166
630 West North Street 30 Marie Curie Street
Geneva, New York 14456 Ottawa, Ontario K1N 6N5
USA Canada

Peck, Stewart B. Pickett, John A.

Department of Biology Rothamsted
4640 CTTC Building Harpenden, Hertfordshire AL5 2JQ
Carleton University UK
Ottawa, Ontario K1S 5B6
Canada Pinzauti, Mauro
Universita degli Studi di Pisa
Pellmyr, Olle Dipartimento di Coltivazione e Difesa delle
Department of Biological Sciences Specie Legnose
University of Idaho Via S. Michele degli Scalzi, n2, 56100 Pisa
Room 252, Life Sciences Building Italy
List of Contributors

Pollet, Marc Pszczolkowski, Maciej

Research Group Terrestrial Ecology Missouri State University and State Fruit
University of Ghent, Ghent, Belgium Experiment Station
and 9740 Red Spring Road
Royal Belgian Institute of Natural Sciences Mountain Grove, Missouri 65711
Brussels USA
Punzo, Fred
Potter, Daniel A.
Department of Biology
Department of Entomology
University of Tampa
University of Kentucky
Box 5F
S-225 Agricultural Science Center Building North
401 West Kennedy Boulevard
Lexington, Kentucky 40546-0091
Tampa, Florida 33606
Powers, Ann M.
Centers for Disease Control and Prevention Purcell, Alexander H.
National Center for Infectious Diseases Division of Insect Biology
Division of Vector-Borne Infectious Diseases University of California
P.O. Box 2087 201 Wellman
Foothills Campus Berkeley, California 94720-3112
Fort Collins, Colorado 80522 USA
Puterka, Gary
Preston, Catherine A. USDA-ARS, PSRL
USDA - ARS 1301 N. Western
Center for Medical, Agricultural and Veterinary Stillwater, Oklahoma 74074
Entomology USA
P.O. Box 14565
1700 SW 23rd Drive
Ragsdale, David W.
Gainesville, Florida 32608
Department of Entomology
University of Minnesota
1980 Folwell Avenue
Prischmann, Deirdre
St. Paul, Minnesota 55108
2923 Medary Ave.
Brookings, South Dakota 57006-9401
USA Raina, Ashok K.
Prokopy, Ronald J. Southern Regional Research Center
Department of Entomology Formosa Subterranean Termite Research Unit
University of Massachusetts 1100 Robert E. Lee Boulevard
Fernald Hall P.O. Box 19687
Amherst, Massachusetts 01003 New Orleans, Louisiana 70179-0687
List of Contributors

Randolph, Sarah Rey, Jorge

Department of Zoology Florida Medical Entomology Laboratory
University of Oxford University of Florida
South Parks Road 200 9th Street S.E.
Oxford, 0X1 3PS Vero Beach, Florida 32962

Rashidan, Kia Ribes, Eva

University of Saint Boniface University of Barcelona
200 Avenue de la Cathedral Barcelona
Winnipeg, Manitoba R2H 0H7 Spain
Richman, David B.
The Arthropod Museum
Ratcliffe, Brett C.
Entomology, Plant Pathology and Weed Science
Systematics Research Collections
University of Nebraska
New Mexico State University
Lincoln, Nebraska 68588-0514
MSC 3BE, Box 30003
Las Cruces, New Mexico 88003
Redborg, Kurt E.
Department of Biology Riddick, Eric W.
Coe College USDA-ARS
1220 First Avenue Biological Control and Mass Rearing Research
Cedar Rapids, Iowa 52402 Unit
USA 810 Highway 12 East
P.O. Box 5367
Reina, David Mississippi State, Mississippi 39762-5367
Faculty of Veterinary Sciences USA
Universities of Cordoba and Extremadura
Cceres, 10071 Riley, David G.
Spain Coastal Plain Region
University of Georgia
Resh, Vincent H. P.O. Box 748
Environmental Science, Policy and Management Tifton, Georgia 31793-0748
University of California USA
201 Wellman Hall
Rinkevich, Frank D.L.
Berkeley, California 94720
Department of Biology
Millersville University
Millersville, Pennsylvania 17551
Retnakaran, Arthur USA
Great Lakes Forestry Center
1219 Queen Street East Rivers, David B.
Sault Ste. Marie, Ontario P6A 5M7 Department of Biological Sciences
Canada Butler University
List of Contributors

4600 Sunset Avenue Jacksonville, Florida 32224

Indianapolis, Indiana 42608 USA
Rothschild, Marjorie
Roderick, George Integrated Microscopy Center
Environmental Science (ESPM) University of Memphis
University of California 201 Life Sciences Building
Berkeley, California 94720-3114 Rooms LS 101-113
USA Memphis, Tennessee 38152-6040
Rogers, Michael E.
Department of Entomology Rust, Michael K.
University of Kentucky Department of Entomology
S-225 Agricultural Science Center North University of California, Riverside
Lexington, Kentucky 40546-0091 Riverside, California
Roltsch, William J.
Rutledge, C. Roxanne
California Department of Food and Agriculture
Florida Medical Entomology Laboratory
Biological Control Program
University of Florida
3288 Meadowview Road
200 9th Street S.E.
Sacramento, California 95832
Vero Beach, Florida 32962-4699
Roe, Kelly
Sanborn, Allen F.
Department of Biological Sciences
Barry University
Western Illinois University
School of Natural and Health Sciences
Macomb, Illinois 61455
11300 NE Second Avenue
Miami Shores, Florida 33161-6695
Rosenberg, David M. USA
Freshwater Institute
501 University Crescent Snchez, Norma E.
Winnipeg, Manitoba R3T 2N6 CEPAVE
Canada Universidad Nacional de La Plata
Consejo Nacional de Investigaciones Cientficas y
Ross, Edward S. Tcnicas
California Academy of Sciences 2 No 584 (1900)
Department of Entomology La Plata
Golden Gate Park Argentina
San Francisco, California 94118
USA Sanford, Malcolm T.
Entomology and Nematology Department
Rossi, Anthony University of Florida
Department of Natural Sciences Gainesville, Florida 32611-0620
University of North Florida USA
List of Contributors

Sarzynski, Erin Schneider, David C.

Entomology and Nematology Department Ocean Sciences
University of Florida Memorial University of Newfoundland
Gainesville, Florida 32611-0620 St. Johns, Newfoundland A1B 3X7
USA Canada

Schning, Caspar
Sastry, Shivashankar Department of Population Biology
National Institute of Mental Health University of Copenhagen
Saint Elizabeth Hospital Universitetsparken 15
U.S. Department of Health and Human Services 2100 Copenhagen
Washington, D.C. Denmark
Schulthess, Fritz
Schabel, Hans G. Contonou
College of Natural Resources Republic of Benin
University of Wisconsin
Stevens Point, Wisconsin 54481 Schuster, Jack
USA Universidad del Valle
Aptd 82
Guatemala City
Scheffrahn, Rudolph H.
Fort Lauderdale Research and Education Center
University of Florida Scotti, Paul D.
3205 College Avenue The Horticulture and Food Research Institute of
Fort Lauderdale, Florida 33314 New Zealand
USA Mt. Albert Research Centre
Scherer, Clay W. New Zealand
DuPont Professional Products
Seal, Dakshina R.
Wilmington, Delaware
Tropical Research and Education Center
University of Florida
18905 SW 280th Street
Schmelz, Eric Homestead, Florida 33031-3314
USDA, Agricultural Research Service USA
Chemistry Research Unit
1600-1700 SW 23rd Drive Serrano, David
Gainesville, Florida 32608 Department of Entomology and Nematology
USA University of Florida
Gainesville, Florida32611
Schmidt, Justin O.
Southwestern Biological Institute Setamau, Mamoudou
1961 W. Brichta Dr. Texas A&M Agricultural Experiment Station
Tucson, Arizona 85745 Weslaco, Texas 78596-8399
List of Contributors

Shaaya, Eli Shirk, Paul D.

Department of Food Science USDA-ARS
Agricultural Research Organization Center for Medical, Agricultural and Veterinary
The Volcani Center Entomology
P.O. Box 6 Box 110970
Bet Dagan, 50250 Gainesville, Florida 32611-0970
Israel USA

Shanower, Thomas G. Showler, Allan T.

Northern Plains Agricultural Research 2413 East Highway 83
Laboratory Building 201
Pest Management Research Unit Weslaco, Texas 78596
1500 N. Central Avenue USA
Sidney, Montana 59270
USA Shukle, Richard H.
Department of Entomology
Shapiro-Ilan, David I.
Purdue University
West Lafayette, Indiana 47907
Southeastern Fruit and Tree Nut Research
21 Dunbar Road
Sikes, Derek S.
Byron, Georgia 31008
University of Alaska Museum
907 Yukon Drive
Fairbanks, AK 99775-6960
Shepard, B. Merle USA
Coastal Research and Education Center
Clemson University Sims, Kelly R.
2865 Savannah Highway Department of Entomology and Nematology
Charleston, South Carolina 29414-5333 University of Florida
USA Gainesville, Florida 32611
Shields, Vonnie D.C.
Biological Sciences Department Sinclair, Bradley J.
Towson University Entomology, Ontario Plant Laboratories,
800 York Road Canadian Food Inspection Agency
Towson, Maryland 21252-3042 K.W. Neatby Building, C.E.F.
USA Ottawa, Ontario, K1A 0C6
Shippy, Teresa D.
Division of Biology Sivinski, John
Kansas State University USDA-ARS
Manhattan, Kansas Center for Medical, Agricultural and Veterinary
USA Entomology
List of Contributors

1700 SW 23rd Drive Soler Cruz, M.D. Amparo

Gainesville, Florida 32604-2565 Department of Parasitology
USA University of Granada
18071 Granada
Skevington, Jeff Spain
Agriculture and Agri-Food Canada
K.W. Neatby Building, C.E.F. Solter, Leellen F.
960 Carling Ave. Illinois Natural History Survey
Ottawa, Ontario K1A 0C6 607 East Peabody Drive
Canada Champaign, Illinois 61820
Slaney, David
Somma, Louis A.
Ecology and Health Research Center
Department of Entomology and Nematology
Departmentof Public Health
University of Florida
Wellington School of Medicine and Health
Gainesville, Florida 32611
University of Otago
P.O. Box 7343
Soroka, Juliana J.
Wellington South
Agriculture and Agri-Food Canada
New Zealand
Saskatoon Research Centre
107 Science Place
Smagghe, Guy Saskatoon, Saskatchewan S7N 0X2
Laboratory of Agrozoology Canada
Faculty of Agricultural and Applied Biological
Sciences Sourakov, Andrei
Ghent University Entomology and Nematology Department
Coupure Links 653 University of Florida
B-9000 Ghent Gainesville, Florida 32611-0620
Belgium USA

Smith, Hugh Spafford, Helen

University of California Cooperative Extension School of Animal Biology (M0895)
624-A West Foster Road University of Western Australia
Santa Maria, California 93455 35 Stirling Highway
USA Crawley, W.A. 6009
Smith, John P.
John A. Mulrennan, Sr., Public Health Stange, Lionel A.
Entomology Research & Education Center Florida Department of Agriculture
Florida A&M University Division of Plant Industry
4000 Frankford Avenue P.O. Box 110980
Panama City, Florida 32405-1933 Gainesville, Florida 32611-0980
List of Contributors

Steck, Gary J. Sullivan, Daniel J.

Florida State Collection of Arthropods Department of Biological Sciences
Division of Plant Industry Fordham University
1911 SW 34th Street Bronx, New York 10458
Gainesville, Florida 32608-1201 USA
Swisher, Marilyn E.
Stocks, Ian Department of Family, Youth and Community
308 Long Hall Sciences
Department of Entomology, Soils, and Plant University of Florida
Sciences Gainesville, Florida 32611-0310
Clemson University USA
Clemson, South Carolina 29634-0315
USA Sword, Gregory
School of Biological Sciences
Steinkraus, Donald C. University of Sydney
Department of Entomology The Macleay Building A12
University of Arkansas Sydney, NSW 2006
Virology Laboratory Australia
319 Agriculture Building
Fayetteville, Arkansas 72701
Tabachnick, Walter J.
Florida Medical Entomology Laboratory
University of Florida
Stewart, Kenneth W.
200 9th Street SE
Department of Biological Science
Vero Beach, Florida 32962-4699
University of North Texas
P.O. Box 305220
Denton, Texas 76203-5220
Tanaka, Seiji
Laboratory of Insect Life Cycles and Physiology
Division of Insect and Animal Sciences
Stonedahl, Gary
Independent Administrative Institution
106 Briza Court
Ohwashi 1-2, Tsukuba, Ibaraki 305-8634
Bellingham, Washington 98226

Striganova, Bella R. Tartar, Aurlian

Laboratory of Soil Zoology and General Entomology and Nematology Department
Entomology University of Florida
A. N. Severtsov Institute of Ecology and Gainesville, Florida 32611-0620
Evolution USA
Russian Academy of Sciences
Leninsky Prospect, 33 Taylor, Steven J.
Moscow 119071 Center for Biodiversity
Russia Illinois Natural History Survey
List of Contributors

607 East Peabody Drive (MC-652) Tinsaara, William

Champaign, Illinois 61820-6970 Bioversity International
USA P.O. Box 24384, Kampala
Teal, Peter E.A.
USDA-ARS Tipping, Christopher
Center for Medical, Agricultural and Veterinary Delaware Valley College
Entomology 700 East Butler Ave.
1700 SW 23rd Drive Doylestown, Pennsylvania 18901
P.O. Box 14565 USA
Gainesville, Florida 32604
Triplehorn, Charles
Tew, James E. Department of Entomology
Ohio State University Museum of Biological Diversity
1608 Madison Avenue Ohio State University
Wooster, Ohio 44691-1030 1315 Kinnear Road
USA Columbus, Ohio 43212-1192
Thomas, Michael C.
Florida State Collection of Arthropods Trumble, John T.
Florida Department of Agriculture and Department of Entomology
Consumer Services University of California
P.O. Box 147100 Riverside, California 92521-0001
Gainesville, Florida 32614-7100 USA
Tsai, James H.
Thompson, Sarah Fort Lauderdale Research and Education Center
Department of Entomology University of Florida
North Carolina State University 3205 College Avenue
P.O. Box 7613 Fort Lauderdale, Florida 33314-7799
Raleigh, North Carolina 27965-7613 USA

Thompson, Vinton Tzanakakis, Minos E.

Roosevelt University Aristotle University of Thessaloniki
Chicago, Illinois 60605 Department of Applied Zoology and
USA Parasitology
Thessaloniki, 541 24
Tilgner, Erich Greece
Department of Entomology
University of Georgia Ueshima, Norihiro
413 Biological Sciences Building Matsusaka University
Athens, Georgia 30602 Matsusaka, Mie
USA Japan
List of Contributors

Uspensky, Igor Verma, K.K.

Department of Chemistry HIG1/327, Housing Board Colony
A. Silberman Institute of Life Sciences Borsi, DURG - 491001
Hebrew University of Life Sciences India
Jerusalem 91904
Israel Vickerman, Danel B.
Department of Entomology
Valles, Steven M. University of California
USDA-ARS Riverside, California 92521-0001
Center for Medical, Agricultural, and Veterinary USA
1600 SW 23rd Drive Villegas, Baldomero
Gainesville, Florida 32608 California Department of Food and Agriculture
USA Biological Control Program
3288 Meadowview Road
Vandergast, Amy G. Sacramento, California 95832
U.S. Geological Survey USA
San Diego, California
USA Vincent, Charles
Horticultural Research and Development Center
Agriculture and Agri-Food Canada
VanderMeer, Robert K.
430 Boulevard Gouin
Saint-Jean-sur-Richelieu, Quebec J3B 3E6
Center for Medical, Agricultural and Veterinary
P.O. Box 14565
Wallace, John R.
1700 SW 23rd Drive
Department of Biology
Gainesville, Florida 32608
Millersville University
Millersville, Pennsylvania 17551
Vzquez, M. ngeles
Departamento de Zoologa y Antropologa Fsica Wang, Ping
C/ Jos Antonio Novais 2, pl.X Lab. 9 Entomology Department
Universidad Complutense de Madrid Cornell University
28040 Madrid New York State Agricultural Experiment Station
Spain Geneva, New York 14456
Vega, Fernando E.
USDA-ARS Wang, Qiao
Insect Biocontrol Laboratory Institute of Natural Resources
Beltsville Agricultural Research Center Massey University
Building 011A, Room 214 PB 11122
Beltsville, Maryland 20705 Palmerston North
USA New Zealand
List of Contributors

Weber, Donald C. Comstock Hall

USDA, ARS, PSI, Insect Biocontrol Laboratory Ithaca, New York 14853-0901
Bldg. 011A, Rm. 107, BARC-West USA
Beltsville, MD 20705
USA Wheeler, Alfred G.
Department of Entomology
Weber, Richard G. Clemson University
Department of Entomology and Applied Ecology 114 Long Hall
University of Delaware Box 340365
Newark, Delaware 19716 Clemson, South Carolina 29634-0365

Webster, Thomas C. Wiener, Linda

Atwood Research Facillity St. Johns College
Kentucky State University Santa Fe, New Mexico 87501
Frankfort, Kentucky 40601 USA
Wild, Alex
Weinstein, Philip Department of Entomology
Ecology and Health Research Center, Department University of Arizona
of Public Health PO Box 2100
Wellington School of Medicine and Health Tucson, Arizona 85721-0036
Sciences USA
University of Otago
P.O. Box 7343 Williamson, R. Chris
Wellington South Department of Entomology
New Zealand University of Wisconsin
237 Russell Labs
Weintraub, Phyllis G. Madison, Wisconsin 53706-1520
Agricultural Research Organization USA
Gilat Research Station
D.N. Negev, 85280 Willis, John S.
Israel Department of Cellular Biology
University of Georgia
Weissman, David B. 724 Biological Sciences Building
Department of Entomology Athens, Georgia 30602-2607
California Academy of Sciences USA
Golden Gate Park
San Francisco, CA 94118 Willis, Judith H.
USA Department of Cellular Biology
University of Georgia
Weston, Paul A. 724 Biological Sciences Building
Department of Entomology Athens, Georgia 30602-2607
Cornell University USA
List of Contributors

Willmott, Keith R. Blacksburg, Virginia 24061-0406

McGuire Center for Lepidoptera Research and USA
University of Florida Yu, Simon J.
Gainesville, Florida 32611 Entomology and Nematology Department
USA University of Florida
Gainesville, Florida 32611-0620
Wing, Steven R. USA
University of Florida
P.O. Box 115001 Zachvatkin, Yuri A.
Gainesville, Florida 32611 Department of Entomology
USA K. A. Timiryazev Agricultural Academy
Timiryazeva St., 49, Build. 12
Moscow 127550
Worner, Sue
National Centre for Advanced Bio-Protection
Zhu, Kun Yan
Bio-Protection and Ecology Division
Department of Entomology
Lincoln University
Kansas State University
P.O. Box 84, Canterbury
123 Waters Hall
New Zealand
Manhattan, Kansas 66506-4004
Wysiecki, Mara L.
CEPAVE Zaspel, Jennifer
Universidad Nacional de La Plata Entomology and Nematology Department
Consejo Nacional de Investigaciones Cientficas y University of Florida
Tcnicas Gainesville, Florida 32611-0620
2 No 584 (1900) USA
La Plata
Argentina Zuparko, Robert L.
Essig Museum of Entomology
Yousten, Allan A. University of California
Department of Biology 201 Wellman Hall
Virginia Tech Berkeley, California 94720-3112
2119 Derring Hall USA

Abafi-Aigner, Lajos (Ludwig original family name: Abafi-Aigner and changed

Aigner) his German Christian name Ludwig to the
Hungarian equivalent Lajos. However, despite
george hangay his successes in publication and writing his busi-
Narrabeen, New South Wales, Australia ness begun to decline in the 1880s and within a
few years he faced financial difficulties, which
Ludwig Aigner was born on the 11 February 1840 ultimately led to the closure of his famous book-
at Nagyjcsa, Torontl Shire, Transylvania, Hungary, shop. Disillusioned, he discontinued most of his
now Romania. His family moved to Temesvr, a business activities, and from 1890 he devoted all
large town in Transylvania, where he received a his time and energy to lepidopterology. In 1895,
formal education in commerce and begun his he published the results of his studies in the
career as a book merchant. His family was of ethnic Termszetrajzi Fzetek (Notebooks of Natural
German stock and young Ludwig only learned History), the journal of the National Museums
Hungarian when, in 1858, they moved to Pozsony Natural History Department and he was one of
(now Bratislava and in the Slovak Republic), a the authors of Fauna Regni Hungariae (Catalogue
large town with predominantly Hungarian inhab- of Hungarys Fauna). He resurrected Rovartani
itants. From here he soon moved on to Pest (now Lapok (Entomological Papers), which was estab-
Budapest) and in 1863, as it was the custom in lished in 1884 but ceased to exist in 1886. His
those years, he wandered all over Austria and treatment of the butterfly fauna of Hungary won
Germany. He completed his studies in Kln and the coveted Bugt Prize. Based on this work he
Stuttgart before returning to Pest. He always had published Butterflies of Hungary in 1907. The
an interest in entomology and he became a keen book was (and probably still is) one of the most
amateur lepidopterologist. However, besides ento- popular entomological publications in Hungary.
mology, he had a great variety of other interests It has inspired countless young entomologists and
too, especially in the field of publishing, writing, made the name of Abafi-Aigner well known to
historical research as well as aspirations in busi- every naturalist in the country. He passed away
ness. He found success in publishing and in estab- on the 19 June 1909.
lishing a popular bookshop. In 1870 he was
initiated as a Freemason and eventually he rose to
the highest positions in the Order. For 12 years he Reference
has worked on an extensive monograph of the
history of Freemasonry. He used his Hungarian Horvth C (1990) A rovartan tudsa. E let s Tudomny
pen name Abafi in a hyphenated form with his 45:290, 312
A Abaxial Surface

Abaxial Surface drawings of a quality that was very high for that
time. Some of the insect illustrations included
The lower surface of a leaf (contrast with adaxial not only adults, but also larvae and the plants on
surface). which they fed, and even observational notes. He
died about 1840.

Abbott, John
John Abbott was born in London in 1751. In Eng-
land, he was given drawing lessons and, through Mallis A (1971) American entomologists. Rutgers University
his drawing instructor, was introduced to Dru Press, New Brunswick, NJ, 549 pp
Drury, a collector of insects who had been presi-
dent of the Linnean Society. These two encoun-
ters encouraged him to collect insects and draw Abbott s Formula
them, but his father was training him to be an
attorney. Finding legal paperwork not to his lik- A mathematical technique commonly used to
ing, he emigrated to Virginia in 1773. After 2 assess mortality in insecticide trials when there is
years in Virginia, he relocated to Georgia, where need to correct for a change (decrease) in the
he served as a private in the Third Georgia Con- background population density (i.e., in the check
tinental Battalion during the Revolutionary War. or control plots). The formula is:
For his military service he received several hun- % corrected control = 100 (% alive in the check %
dred acres of land, and worked as a planter and alive in the treatment)/(% alive in the treatment)
schoolmaster. In Virginia he had collected Amer-
ican insects and bird skins, and drew and painted
insects and birds. Some of the specimens and
paintings were shipped to England for sale. Some Abdomen
of the paintings, after sale, adorned books on
birds, insects, and spiders written by various The posterior of the three main body divisions of
authors, not necessarily with acknowledgment an insect (Fig. 1).
to Abbott. In all, Abbott produced over 3,000 Abdomen of Hexapods

Abdomen, Figure 1 Cross section of an insect abdomen, showing components of the insect circulatory
system and direction of hemolymph flow (adapted from Evans, Insect biology).
Abdomen of Hexapods
A 3

Abdomen of Hexapods primitive, the number of abdominal segments is

usually greater, as occurs in the Protura with 11
severiano f. gayubo segments (Figs. 25). An exception is the Collem-
Universidad de Salmanca, Salamanca, Spain bola, which only possess six. In addition, it is nec-
essary to keep in mind that, in certain cases, the
The abdomen constitutes the caudal tagma in the total number of visible segments does not coin-
hexapods and is usually larger than the other two, cide with what a particular individual actually
the head and the thorax. This region is also referred possesses, since some segments remain invisible
to as a visceral area because it houses the visceral upon being telescoped, particularly those of the
organs. Its form can vary depending on the group, posterior region of the abdomen.
and even on the species. The maximum number of According to Bitsch, a generalized abdominal
observed segments is 11, although certain authori- segment would be limited anteriorly by a preseg-
ties consider a twelfth segment that in fact corre- mentary domain, separated from the segmentary
sponds to a telsonic caudal region. In general, the domain proper (of greater size) by a suture that
number of segments decreases from the preimagi- begins an internal crest named the costa or antecosta.
nal phases to the adult stage, especially in those This crest anteriorly delimits an acrotergite or pre-
holometabolous insects in which the last segments costa in the tergal part and a presternite in the sternal
of the adults are formed from imaginal discs part. In this idealized model, the muscles would be
during pupation. In the groups considered most inserted in successive antecostas. No known struc-
ture is homologous to the thoracic furca.
The presence of the gonopore (double in
Ephemeroptera) in segments VIII and IX (in VII
in the case of Ephemeroptera), and fundamentally
of the external structures related to reproduction
(the genitalia), produce important modifications
in those segments. Considering the presence of
these genitalia, three regions of the abdomen are
recognized: an anterior (pregenital or visceral
region that includes the first eight segments),
median (genital region, eighth and ninth segments),
and caudal regions (postgenital region, tenth and
eleventh segments plus the telsonic region).

The Pregenital Region

In the most generalized condition, the first abdom-

inal segments conserve their basic structure, being
easily distinguished from the thoracic segments.
Nevertheless, the most frequent condition is that
which produces morphological modifications that
Abdomen of Hexapods, Figure 2 Diagram of a affect the thoracic-abdominal union. These modi-
proturan (Protura) showing abdominal segments fications usually consist of reductions that affect
and appendages: dorsal view (left), ventral view the sternal region and involve a greater or lesser
(right). desclerotization of different structures and their
A Abdomen of Hexapods

Abdomen of Hexapods, Figure 3 Diagram of chewing louse (Mallophaga) showing abdominal segments,
including numbering of segments: dorsal view (left), ventral view (right).

incorporation to the metathorax. In this sense, the modified appendages exist in Collembola. In
case of the Hymenoptera, Apocrita stands out, in Archaeognatha, very developed coxites are differ-
which a narrowing is produced between the sec- entiated, above which are inserted styli in a median
ond and the third abdominal segments, which position and the exsertile vesicles in the most
incorporate the thorax and is named the propo- internal position.
deum. The rest of the abdominal segments are The styli are elongated pieces, articulated in
called the gaster or metasoma. The region formed their base above the external face of the coxite.
by the propodeum and the thorax constitutes the They are unisegmentary and lack muscles
mesosoma. The narrowing allows a great ampli- inserted in their base, often presenting an apical
tude of movements of the metasoma, which per- spine. Taking into account their position and
mits stinging in the capture of prey in aculeates. In their embryonic development, the styli are con-
some groups, like Formicidae and Sphecidae (Acu- sidered by the majority of authorities as vestigial
leata), one or two segments of the metasoma form appendages, and more concretely as reduced
a narrower zone called the petiole. telepodites. The exsertile vesicles are considered
In the pregenital region, several appendicular internal coxal formations (internal coxalia of
structures can be found. Thus, three pairs of highly some authorities).
Abdomen of Hexapods
A 5

least in part, it is of appendicular origin. In this

sense, it is clear that in the Archaeognatha the
tibiotarsus eye eighth and ninth segments are basically similar in
femur pronotum males and females and their structures are homol-
mesonotum ogous to those already indicated for the pregenital
precoxae metanotum segments. Taking into account this relationship,
collophore the genitalia of Archaeognatha are considered
tenaculum (catch) primitive, and therefore fundamental to interpret
the genitalia of Pterygota.
In the eighth segment of the Archaeognatha,
the basal part of the appendicular structure is named
first gonocoxa or gonocoxite and bears the first
gonostylus; in the ninth is found the second gono-
manubrium coxa with its corresponding stylus. In both segments
(at times in the eighth, always in the ninth) forma-
mucro (spring)
tions homologous to the exsertile vesicles appear,
which are called gonapophyses (parameters in the
males and gonapophysis proper in the females). The
fundamental difference between both sexes lies in
Abdomen of Hexapods, Figure 4 Diagram
the presence in the males of a phallic structure.
of springtail (Collembola) showing furcular
The female genitalia in the Pterygota constitute
appendage at tip of abdomen.
the ovipositor. The gonocoxites are incorporated
into the lateral wall of the genital segments in a
In Pterygota the abdominal appendages complete manner in the eighth segment, forming
remain restricted to the larval forms (Lepidoptera the first valvifer. In the ninth segment the basal part
and Hymenoptera, Tenthredinoidea), although is incorporated into the lateral wall, originating the
rough appendicular pairs already exist in the second valvifer, while the rest is extended, forming
polypodous type of embryos. These abdominal the third pair of valves (dorsal or lateral valves of
appendages are named false legs or prolegs and some authorities), which are not homologous in
are retractile, conical and membranous projec- Archaeognatha. The other two pairs of valves are
tions, with a circular planta that bears a crown, the ventral valves, corresponding to the eighth seg-
usually with hooks, to adhere to the substrate. ment, and the internal valves, corresponding to the
ninth segment. These two pairs of valves are homol-
ogous to the gonapophysis of Archaeognatha. In
The Genital Region the case of the generalized type of ovipositor like
that of Orthoptera, these three pairs of valves are
The transformations that affect the eighth and linked through the length of their course, forming
ninth abdominal segments are a consequence of in their interior a canal for oviposition.
the development of special external structures that Among the sclerites that are situated in the
in the case of the male serve in the transfer of base of the valves (in addition to the valvifers
sperm, and in the case of the female allow for ovi- already mentioned) are found the intervalves (inter-
position. These structures together are known by valvulae of the authorities) by way of elongated
the name genitalia. transverse formations, one in the base of the valves
The origin of the genitalia is controversial, of the eighth segment and another in the base of the
although the majority of authorities accept that, at valves of the ninth segment. The typical ovipositor
A Abdomen of Hexapods

Abdomen of Hexapods, Figure 5 Comparative development of cerci on earwig (Dermaptera, top left);
grasshopper (Orthoptera, top right); scorpion fly (Mecoptera, lower left); silverfish (Zygentoma).

that was just described can experience modifica- developed, in such a way that they can become
tions according to the functions that it carries out; telescoped, forming oviposition tubes; this type of
one of the most drastic is found in Hymenoptera, ovipositor is named the ovicauda. Not being
Aculeata, where it is transformed into a sting that homologous to the genitalia, many authorities call it
serves the females as an attacking organ, either to terminalia.
capture prey or as a defense. On the other hand, the The masculine genitalia present great mor-
process of oviposition can be carried out through phological variability, which together with their
other, different structures, as occurs in the females taxonomic importance, have been the object of an
of certain Diptera. In that case, the last segments are infinity of descriptions, many of them without
retractile and the intersegmental zones are highly truly anatomical criteria. This has originated the
Abdomen of Hexapods
A 7

use of very varied terminologies that have done or on the contrary, increase in complexity, develop-
nothing but complicate its study and impede the ing spines and other types of processes named fla-
establishment of homologies even in the same group, gellum, virga or pseudovirga over the internal walls of
creating in this way a great nomenclatorial chaos. the endophallus. When the endotheca and the
In the males, in addition to the genitalia proper, endophallus are evaginated, the genitalia are converted
other structures (processes, lobes, etc). exist that into authentic intromittent organs.
intervene in functions other than those strictly The primitive position of the male genitalia
related to the transfer of sperm; among the most can be displaced through different types of turns;
common is the grasping of the female during mat- one of the most showy cases is that which occurs
ing. It has already been mentioned that the major- in some Hymenoptera, Symphyta that present the
ity of authorities consider that the interpretation of condition called strophandric, which is character-
the genitalia of Pterygota should be made by ized by a 180 rotation of the genitalia. Rotations
homology with the basic condition that is found in have also been observed in males of Diptera.
Archaeognatha. In this group, the phallic complex The postgenital region, as was mentioned in
is formed by a median organ, the phallus or penis, the beginning of this section, comprises the tenth
and a pair of segmented pieces named parameres, and eleventh segments plus the telsonic region. The
that in the case of maximum development can exist tenth segment has been detected in Protura,
in the eighth and ninth segments. The parameres Diplura, Archaoegnatha, Thysanura (Zygentoma),
correspond to the gonapophysis of the females Ephemeroptera, Plecoptera, and some Orthoptera.
(although the term gonapophysis is utilized indis- The morphology of this segment is basically simi-
tinctly for both sexes by some authorities). lar to the pregenital segments, although with cer-
Many morphological models have been pro- tain frequency it can form a ring when the tergum
posed to describe the male genitalia of Pterygota. and sternum unite, or the sternal region can be
The most complete, since it gathers and discusses membranous. In embryonic forms, a pair of appen-
early data, is that proposed by Bitsch. According dicular outlines is seen above this segment. In cer-
to this author, what together forms the copulatory tain holometabolous insects, structures of uncertain
organ (phallus or penis) and the structures associ- meaning appear, such as the socii of some
ated with the parameres (considered in the sense Hymenoptera.
expressed by the Archaeognatha) is named the The eleventh segment is recognized in the
phallic complex. majority of embryonic phases of hexapods. In
The aedeagus is a sclerotized tube, situated Archaeognatha and Thysanura it forms an annular
above a largely membranous phallobase, although structure from whose dorsal part is differentiated a
in more complex cases the phallobase presents an long and narrow process called filum terminale,
internal fold that remains membranous (endoth- while from the lateroventral position are differenti-
eca) while the external part is sclerotized (phalloth- ated the cerci that in the adults possess numerous
eca or theca). The aedeagus presents an invagination divisions. In the Pterygotes, the eleventh segment is
that forms a more or less developed internal cham- formed by the epiproct (tergal region) and the
ber (the endophallus), which communicates with paraprocts (in the lateroventral position); in the
the gonopore at its base and in the other extreme more primitive groups exist cerci (whose length
communicates with the exterior through the phal- and number of divisions are variable) situated in
lotreme. In counter-proposition to the endophallus, the membranous zones that exist between the
the part formed by the external walls of the phal- epiproct and the paraprocts. The telsonic, asegmen-
lobase and the aedeagus forms the ectophallus. The tary region constitutes the perianal membrane or
phallic complex can present variable development, periproct.
even being able to cause the aedeagus to disappear, Alimentary Canal and Digestion
A Abdominal Pumping

References Abnormality

Bitsch J (1979) Morphologie abdominal des insects. In: In insect pathology, deviation from the normal; a
Grass, P-P (ed) Trait de Zologie, VIII (II): 291600 malformation or teratology; a state of disease.
Bitsch J (1994) The morphological groundplan of Hexapoda:
critical review of recent concepts. Annales de la Socit
Entomologique de France 30:103129
Deuve T (2001) The epipleural field in hexapods. Annales de
la Socit Entomologique de France 37:195231
Matsuda R (1970) Morphology and evolution of the insect
abdomen. Pergamon Press, New York, NY A family of chewing lice (order Phthiraptera).
Snodgrass RE (1935) Principles of insect morphology. Chewing and Sucking Lice
MacGraw Hill, New York, NY

Abdominal Pumping Absolute Methods of Sampling

Contraction of the muscles associated with the Techniques used to sample insect populations that
abdomen can result in collapse and expansion of provide an estimate per unit of area (e.g., per
the air sacs. This forces relatively large volumes of square meter, per leaf or per plant). Types of abso-
air in and out of the insect through the spiracles, lute methods include unit of habitat, recapture,
promoting ventilation. This is called active ventila- and removal trapping. (contrast with relative
tion, in contrast with the more normal gas exchange methods of sampling).
mechanism of insects, diffusion or passive ventilation. Sampling Arthropods
To a small degree, abdominal pumping also pro-
motes gas exchange through the trachea, but the
trachea is quite resistant to change in shape. Acanaloniidae
Abdominal pumping is more important for larger
insects such as locusts, which display abdominal A family of insects in the superfamily Fulgoroidae
pumping almost continuously, but especially when (order Hemiptera). They sometimes are called
active. In these insects air is sucked in through planthoppers.
some spiracles and pumped out through others. Bugs
Active Ventilation

Abiotic Disease Acanthmetropodidae

A family of mayflies (order Ephemeroptera).
A disease caused by factors other than pathogens
(e.g., weather or nutrition).

Abiotic Factors Acanthopteroctetidae

Factors, usually expressed as factors affecting mor- A family of moths (order Lepidoptera). They com-
tality, characterized by the absence of life. Abiotic monly are known as archaic sun moths.
factors include temperature, humidity, pH, and Archaic Sun Moths
other physical and chemical influences. Butterflies and Moths
Acaricides or Miticides
A 9

Acanthosomatidae applying a measured amount to the integument.

The lower the LD50 or LC50, the more toxic the
A family of bugs (order Hemiptera). poison.
Bugs An LC50 is obtained when a mite is exposed to
a particular concentration of toxicant but the actual
amount of toxicant the individual experiences is
Acaricide not determined. For example, if the pesticide is
applied to foliage and the mite walks about on the
A pesticide applied to manage mite populations. foliage, the actual amount of toxicant the mite is
An acaricide is also called a miticide. exposed to depends on the activity of the mite, the
Acaricides or Miticides amount taken up through the integument or by
Figure 6 shows a concentration-response
Acaricides or Miticides curve in parts per million (ppm) for the acaricide
Omite (propargite) exhibited by adult females
marjorie a. hoy from colonies of the Pacific spider mite Tetrany-
University of Florida, Gainesville, FL, USA chus pacificus. The concentration required to kill
50% of the individuals is the LC50. The two types
An acaricide or miticide is a pesticide that pro- of F1 females (produced by crossing Chapla
vides economic control of pest mites and ticks. males and Bidart females, and vise versa) respond
Mites and ticks are collectively called either acari similarly and their concentration-response
or acarina. Some products can act as insecticides curves are about midway between those of the
or fungicides as well as acaricides. resistant (Bidart) and susceptible (Chapla) colo-
An acaricide is a pesticide used to kill mites nies, which indicates that resistance may involve
and ticks (Table 1). Always check with state and a semidominant mode of inheritance. The term
federal authorities to be sure products containing mode of inheritance describes how the trait is
these active ingredients are registered for use. inherited; for example, the resistance can be
Always read labels carefully and follow the direc- determined by a single major dominant (only
tions completely. one copy of the gene is required for the mite to
The toxicity of an acaricide is determined by a express the resistance) or recessive (two copies of
dose-response curve or a concentration-response the gene are required) gene. Or, the resistance
curve. Such curves are obtained by exposing test can be a quantitative trait determined by multiple
mites or insects to increasing concentrations or genes of equal and additive effect. In this exam-
doses of the pesticide and recording the resulting ple, the propargite resistance may be determined
mortality after a given time interval. One estimate a single semidominant gene with modifying
of toxicity used is the term LD50 (which is the dose genes, but additional tests are required to resolve
required to kill 50% of the test population). The whether more than one gene actually contributes
LC50 is the concentration required to kill 50% of to this resistance.
the test population. If the dose is introduced
through the insects mouth it is an oral LD50, if it is
introduced through the skin or integument it is a Acaricide Classification
dermal LD50, and if it is introduced through the
respiratory system it is the inhalation LD50. A mea- Pesticides are classified in several ways, including:
sured dose is applied to an arthropod by inserting (i) their mode of entry into the target pest, (ii)
a measured amount of toxicant into the gut or by chemical structure, or (iii) source.
A Acaricides or Miticides

Acaricides or Miticides, Table 1 Acaricides (miticides) currently or recently available for general and
restricted use to control mites and ticks*
Name** (chemical type) General Use (GU)*** Potential use
Some trade names Restricted Use (RU)
Abamectin (avermectin B1a; GU, Class IV (practically nontoxic) Also an insecticide; affects
produced from the bacterium nervous system and paralyzes
Streptomyces avermitilis) Affirm, insects or mites; used in citrus,
Agri-Mek, Avid, vertimec, Zephyr pears, nut tree crops
Amitraz (triazapentadiene) GU, Class III (slightly toxic) Used in pears, cotton, and on
Acarac, Mitac, cattle, and hogs to control
Ovidrex,Triatox,Topline insects, ticks and mites
Azadirachtin GU, Class IV Azadirachtin is similar to insect
(tetranortriterpenoid extracted hormones called ecdysones,
from the Neem tree) Align, Azatin, which control metamorphosis;
Turplex also may serve as a feeding
deterrent; used to control insects
and mites on food, greenhouse
crops, ornamentals and turf
Bifenazate (carbazate) Floramite Class IV Mites on greenhouse,
shadehouse, nursery, field, field,
landscape and interiorscape
ornamentals, not registered in
USA for use on food
Bifenthrin (pyrethroid) Talstar, RU, Class II (moderately toxic) Insecticide and acaricide that
Brigade, Capture affects the nervous system and
causes paralysis; used on
greenhouse ornamentals and
Carbaryl (carbamate) Adios, GU, Class I, II or III, depending General use pesticide to control
Bugmaser, Crunch, Dicarbam on insects on citrus, fruits, cotton,
formulation Hexavin, Karbaspray, forests, lawns, nuts, ornamentals,
Septene Sevin, Tornadao, Thinsec shade trees, poultry, livestock and
pets. Also works as a mollusccide
and acaricide
Chlorobenzilate (chlorinated RU, Class III, may cause tumors in Used for mite control on citrus
hydrocarbon) Acaraben, Akar, mice and in beehives; also kills ticks;
Benzilan, Folbex use cancelled in USA
Chlorfenapyr (pyrrole) Pylon, Class I Used to control spider mites,
Pyramite, Pirate broad ites, budmites, cyclamen
mite, rust mites and some insects.
Cinnamon oil (cinnamaldehyde) Exempt from registration under Broad spectrum miticide/
Cinnamite FIFRA insecticide/fungicide controls or
repels pests; could be phytotoxic
in some cases; used in ornamentals,
shade or nursery trees,
vegetables, herbs and spices
Citronella oil Exempt from FIFRA Repels insects and ticks
Demeton-S-Methyl No longer registered for use in Systemic and contact insecticide
(organophosphate) Meta-Systox, USA; Class I, highly toxic and acaricide, widely used
Azotox, Duratox, Mifatox against diverse pests
Acaricides or Miticides
A 11

Acaricides or Miticides, Table 1 (Continued)

Name** (chemical type) General Use (GU)*** Potential use
Some trade names Restricted Use (RU)
Dicofol (organochlorine) Acarin, GU, Class II or III, depending on Miticide used on fruits,
Difol, Kelthane, Mitigan formulation vegetables, ornamentals and field
Dicrotophos (organophosphate) RU Contact systemic pesticide and
Bidrin, Carbicron, Dicron, Ektafos acaricide used to control
sucking,boring and chewing
pests on coffee, cotton, rice,
pecans; used to control ticks on
Dienochlor (organochlorine) GU, Class III Contact material used for
Pentac, often formulated with plant-feeding mites on
other pesticides ornamental shrubs and trees
outdoors and in greenhouses;
disrupts egg laying of female
mites; use cancelled in USA
Dinocap (dinitrophenyl) GU, Class III Used as a fungicide and as an
Arathane, Caprane, Dicap, Dikar acaricide for ticks and mites; use
Karathane, Mildane cancelled in USA
Disulfoton (organophosphate) RU, Class I, highly toxic Systemic insecticide and acaricide
Disyston, Disystox, used to control sucking insects/
Dithiodemeton, Dithiosystox, mites on cotton, tobacco, sugar
Solvigram, Solvirex beets, cole crops, corn, peanuts,
wheat, grains, ornamentals,
Endosulfan (chlorinated RU, Class I Contact insecticide and
hydrocarbon) Afidan, Cyclodan, acaricideused to control many
Endocide, Hexasulfan, Phaser, pests on tea, coffee, fruits,
Thiodan, Thionex vegetables, grains
Ethion (organophosphate) GU, Class II Insecticide and acaricide used on
Acithion, Ethanox, Ethiol, Nialate, wide variety of food, fiber and
Tafethion, Vegfru Foxmite ornamentals, including
greenhouse crops, citrus, lawns
and turf
Eucalyptus oil Exempt from FIFRA Repels mites; repels fleas and
Fenamiphos (organophosphate) RU, Class I A nematicide that has some
Nemacur, Phenamiphos, Bay activity against sucking insects
68138 and spider mites
Fenbutatin oxide (organotin) RU Miticide used on perennial
Vendex fruits,eggplant and ornamentals
Fenitrothion (organophosphate) GU Acaricide and insecticide
Accothion, Cyfen, Dicofen, effective gainst a wide array of
Fenstan, Folithion, Mep, pests
Metathion, Micromite Pestroy,
Sumithion, Verthion
A Acaricides or Miticides

Acaricides or Miticides, Table 1 (Continued)

Name** (chemical type) General Use (GU)*** Potential use
Some trade names Restricted Use (RU)
Formothion (organophosphate) RU, Class II Systemic and contact insecticide
Aflix, Anthio, Sandoz S-6900 and acaricide, used against spider
mites on tree fruits, vines, olives,
hops, cereals, sugar cane, rice
Hexythiazox (ovicide, growth Class III Ovicide/miticide effective against
regulator) Savey spider mites on tree fruits,
christmas trees, strawberries,
hops, peppermint, caneberries
Lambda cyhalothrin (pyrethroid) RU, Class II Insecticide and acaricide used to
Charge, Excaliber, Granade, control a variety of pests in
Hallmark, Icon, Karate, Matador, cotton, cereals, hops,
Saber, Sentinel ornamentals, potatoes,
vegetables; controls ticks
Lindane (organochlorine) RU, Class II Most uses cancelled in Insecticide and fumigant; used in
Agrocide, Benesan, Benexane, USA because of potential to cause lotions, creams and shampoos for
BHC, Gammex, Gexane, HCH, Iso- cancer control of lice and mites (scabies)
tox, Kwell, Lindafor, Lintox, Lorex- in humans
ane, Steward
Methamidophos RU, Class I Systemic, residual insecticide/
(organophosphate) Monitor, acaricide/avicide with contact and
Nitofol, Tamaron, Swipe Patrole, stomach action, used to control
Tamanox chewing and sucking insects and
mites in many crops outside the
Methidathion RU, Class I Insecticide and acaricide with
(organosphosphate) Somonic, stomach and contact action used
Supracide, Suprathion to control a variety of insects and
mites in many crops
Methomyl (carbamate) Acinate, RU, Class I Broad spectrum insecticide and
Agrinate, Lannate, Lanox, Nudrin, an acaricide to control ticks, acts
NuBait as a contact and systemic
Mevinphos (organophosphate) RU, Class I Insecticide and acaricide effective
Fosdrin, Gesfid, Meniphos, Menite, against a broad spectrum of pests,
Mevinox, Mevinphos, Phosdrin, including mites and ticks; use
Phosfene cancelled in greenhouses
Monocrotophos RU, registration in USA withdrawn Systemic and contact insecticide
(organophosphate) Azodrin, in 1988 and acaricide
Bilobran, Monocil 40, Monocron,
Nuvacron, Plantdrin
Naled (organophosphate) GU, Class I Contact and somach insecticide
Bromex, Dibrom, Lucanal and acaricide, used against mites
in greenhouses
Oxamyl (carbamate) RU, Class I granular form is banned Insecticide/acaricide/nematacide
in USA that controls a broad spectrum of
mites, ticks and roundworms on
field crops, vegetables, fruits,
Acaricides or Miticides
A 13

Acaricides or Miticides, Table 1 (Continued)

Name** (chemical type) General Use (GU)*** Potential use
Some trade names Restricted Use (RU)
Neem oil Trilogy Broad spectrum fungicide and
acaricide in citrus, deciduous
fruits and nuts, vegetables, grains
Permethrin (pyrethroid) Ambush, Class II or III, depending on Broad spectrum used on nut, fruit,
Cellutec, Dragnet, Ectiban, formulation RU in agriculture vegetable, cotton, ornamentals,
Indothrin, Kafil, Kestrel, Pounce, because of adverse effects on mushrooms, potatoes, cereals, in
Pramex, Zamlin, Torpedo aquatic organisms greenhouses, home gardens, on
domestic animals
Petroleum oils (refined petro- Class IV Kills by contact a wide range of
leum distillate) Sunspray and mite and insects; complete
others coverage is essential; may act as a
feeding or oviposition deterrent.
Phytotoxicity can occur if plants
are stressed, especially by lack of
water; some plant cultivars are
more susceptible than others.
Used as dormant and as foliar
Phorate (organophosphate) RU, Class I Insecticide and acaricide used on
Agrimet, Geomet, Granutox, pests, including mites, in forests,
Phorate Rampart, Thimenox, root and field crops, ornamentals
Thimet, Vegfru and bulbs
Phosalone (organophosphate) GU, No longer for sale in USA due Broad spectrum insecticide/
to carcinogenic effects acaricide used on deciduous
trees, vegetables, cotton.
Phosmet (organophosphate) GU, Class II, some tolerances in Broad spectrum insecticide, used
foods changed in 1994 by EPA to control insect and mites on
apples, ornamentals, vines; is
used in some dog collars.
Propargite (organosulfide) GU Acaricide used in many crops but
Comite, Omite not USA
Rosemary oil (rosemary essential Meets requirements of USDA Broad spectrum contact
oil) Hexacide National Organic Program Exempt insecticide/miticide used in fruits,
from FIFRA nuts, vegetables. Could be
phytotoxic on some cultivars.
Soybean oil (essential oil) Low acute toxicity to humans,
generally recognized as safe
Spinosad (macrocyclic lactone) Broad spectrum insecticide and
Conserve miticide used on ornamentals and
in greenhouses.
Sulfur (sulfur) Cosan, Hexasul, GU, Check label for restrictions Fungicide and acaricide; used to
Sulflox, Thiolux control plant diseases, gall mites,
spider mites, used widely in food
and feed crops, ornamentals, turf
and residential sites; a fertilizer or
soil amendment, mixing with oil
can cause phytotoxicity
A Acaricides or Miticides

Acaricides or Miticides, Table 1 (Continued)

Name** (chemical type) General Use (GU)*** Potential use
Some trade names Restricted Use (RU)
Triforine (piperazine derivative) RU, Class I Fungicide used on almonds,
apples, asparagus, berries,
cheeries, hops, ornamentals,
peaches, rose; also controls spider
Wintergreen oil (contains methyl Exempt from FIFRA Used to control mites (Varroa) in
salicylate) honey bees; causes contact
mortality and reduced fecundity
when mites feed on syrup

* The list is based on chemicals currently registered in the USA, which can change as new information regarding
environmental impact and human health effects become available. Inclusion in this list does not necessarily indicate that
the products are effective acaricides; application methods and resistance levels in individual mite populations can affect
**Most have a variety of trade or other names, as well as different formulations, which can affect their toxicity.
***Restricted Use (RU) means that pesticides may be purchased and used only by certified applicators. Check with
specific state regulations for local restrictions.

Mode of Entry
90 Chapla A pesticide can enter and kill mites as stomach
reciprocal F1 females
80 poisons, contact poisons, and or as fumigants.
A systemic acaricide is absorbed into a plant or
animal and protects that plant or animal from
pests after the pesticide is translocated through-
20 out the plant or animal.
10 Bidart
Chemical Structure
101 102 103
ppm propargite Pesticides are classified as organic or inorganic.
Inorganic pesticides do not contain the element
Acaricides or Miticides, Figure 6 This is a
carbon (but include arsenic, mercury, zinc, sulfur,
concentration-response curve showing the
boron, or fluorine). Most inorganic pesticides have
responses of a colony of Tetranychus pacificus
been replaced by organic pesticides.
resistant (Bidart) and susceptible (Chapla)
to propargite (Omite). The mortality of adult
females at different concentrations has Source
been transformed into a straight line. The
concentration-responses of the reciprocal F1 Organic pesticides include botanicals (natural
females in crosses between the susceptible organic pesticides) produced by plants (such as
and resistant populations are intermediate and natural pyrethrums, nicotine, rotenone, essential
similar. oils such as those from the neem tree, soybean
Acaricides or Miticides
A 15

oil). Essential oils are any volatile oil that gives formulations also can influence toxicity to dif-
distinctive odor or flavor to a plant, flower or ferent species of both insects and mites.
fruit, such as lavender oil, rosemary oil, or citrus Many insecticides are effective acaricides (or
oil. Essential oils have been registered as pesti- at least they were before resistance to them devel-
cides since 1947 and at least 24 different ones are oped). For example, many OPs (such as azinphos-
available in registered products. These are used as methyl, parathion, ethion, dimethoate) were toxic
repellants, feeding depressants, insecticides, and to spider mites until resistance to these products
miticides. Botanicals have relatively high LD50 developed. Likewise, carbamates, formamides, and
values to mammals, so usually are considered many pyrethroids have both insecticidal and aca-
safe to humans. Some newer pesticides are ricidal properties. Other products have both fun-
derived from microbes, such as avermectin or gicidal and acaricidal properties. The reasons as to
spinosad. why these products are effective on particular tax-
Synthetic organic pesticides are commonly onomic groups are generally unknown.
used in pest management programs and can be
separated into groups based on their chemistry.
Themain groups are: chlorinated hydrocarbons Acaricide Types
(such as DDT and chlordane, which are banned
from use in most parts of the world), organo- Pesticide registrations change frequently so some
phosphates (such as malathion, parathion, azin- of the materials listed here may be obsolete. Always
phosmethyl), carbamates (carbaryl, propoxur), check with state and federal authorities to be sure
pyrethroids (permethrin, fenvalerate),and a products containing these active ingredients are
variety of newer products with very different registered for use. Always read labels carefully and
chemistries including nicitinoids, pyrroles, car- follow the directions completely.
bazates, and pyridazinones.

Chlorinated Hydrocarbons
Insecticides as Acaricides
Dienochlor (Trade name=Pentac) is a chlorinated
Many insecticides have acaricidal properties. hydrocarbon acaricide with long residual activity.
Sometimes an insecticide is more effective as an It has been used in greenhouses and on outdoor
insecticide than as an acaricide (lower concen- ornamentals. Pentac cannot be used on food crops
trations are required to kill the insect than are and has short residual activity when used outdoors.
required to kill the mite species). Some products It has a rapid effect on mites, stopping their feeding
are more toxic (often for unknown reasons) to within hours. Endosulfan and DDT have also been
mites than to insects. We think that mites have used as acaricides (as well as insecticides).
the same fundamental physiological responses
to toxic chemicals as insects, although mite
physiology and responses to pesticides have been Essential Oils
studied less often. Different mitespecies appear
to respond differently to different products, Soybean oil was first registered in 1959 for use as
which could be due to behavioral differences an insecticide and miticide. Three products cur-
(feeding behavior, location on plant, activity rently are registered to control mites on fruit trees,
levels), differences in cuticle thickness, differences vegetables and a variety of ornamentals. Soybean
in detoxification rates, or other biochemical, oil is not phytotoxic under most conditions. Many
morphological or behavioral factors. Different of these oils are approved for organic farming.
A Acaricides or Miticides

Inorganics Different types of petroleum oils are used with

different molecular weights. Most oils used are dis-
Sulfur is a good acaricide and fungicide, although it tillations of petroleum, although some oils derived
can be phytotoxic (cause plant injury), especially if from plants (sesame, almond, citrus) are used.
plants are not well watered during hot weather. Sul- Crude petroleum oil is a complex mixture of
fur is probably the oldest known acaricide. Sulfur hydrocarbons with both straight chain and ring
(dusts, wettable p owders and flowable formulations) molecules. Crude oil is separated into a range of
are usually highly effective acaricides for spider mites products by distillation and refining. The lightest
and rust mites, with two known exceptions. fractions include gasoline, kerosene, diesel and jet
Spider mites in California vineyards (Tetrany- fuel. As these lighter fractions distill or boil, they
chus pacificus and Eotetranychus willamettei) are separated into different fractions. Spray oils are
developed resistance to sulfur, probably because derived from the lighter lubricating oil fraction
sulfur was applied up to 20 times a season over and distill at a temperature range of 600 to900C.
many years to control powdery mildew. After a Currently used petroleum oils in the USA are
number of years, these spider mites became pests narrow-range oils and have had the waxes, sulfur,
because they were no longer controlled by the sul- and nitrogen compounds removed. Labels on
fur which had been applied to control powdery sprays usually describe the degree to which the
mildew. A number of years later, a predatory mite sulfur compounds have been removed and the
called Metaseiulus occidentalis was demonstrated percentage of active oil. The sulfur compounds
to have developed a resistance to sulfur. The resis- arelikely to cause phytotoxic effects, so the degree
tance to sulfur in this natural enemy of spider of removal of these compounds (called the UR
mites is based on a single major dominant gene; rating) is an important piece of information on
once the predator became resistant to sulfur it the label and commonly is greater than 92%. The
became an effective predator of spider mites in composition of oil should be greater than 60%.
San Joaquin Valley vineyards in California. Since the mid-1960s, narrow-range horticul-
The resistance to sulfur in M. occidentalis is tural oils have been used both as dormant or sum-
unusual; even very high rates of sulfur are non- mer oil sprays. These highly refined and narrow
toxic to the resistant populations. Interestingly, range petroleum oils rarely cause phytotoxicity and
populations of this predator collected from nearby increasingly are used for controlling both insect
almond orchards in California are susceptible to and mite pests on deciduous trees, citrus, and orna-
sulfur, indicating that populations are subjected to mental trees and shrubs. Oils have a wide range of
local selection and evolution. No genetic analyses activity against scales, mites, psyllids, mealybugs,
have been conducted on the resistance to sulfur in whiteflies, leafhoppers, and eggs of mites, aphids
the spider mites, so their mode of inheritance to and some Lepidoptera. Heavier dormant sprays are
sulfur resistance remains unknown. The biochem- used to control overwintering pests in deciduous
ical mechanism of resistance is unknown for both trees and vines. Summer oils are used to control
spider mites and their predators. pests during the growing season.
Oil kills mites and their eggs by contact. The
toxicity appears to be due to suffocation of the pest,
Petroleum Oils although it may also be due to chemical effects. Oils
block spiracles, reducing the availability of oxygen
Petroleum oils are excellent insecticides/acari- and suffocation occurs within 24 h. Penetration and
cides/fungicides for integrated mite management corrosion of tracheae, damage to muscles and
programs and have been used in pest management nerves may also contribute to the toxicity of oils.
programs for over 100 years. Oils are sometimes a repellent to pests. Once the oil
Acaricides or Miticides
A 17

dries it is no longer toxic to most natural enemies; Organotins

thus the very short residual activity of oil makes it a
useful material for integrated mite management Cyhexatin (Plictran) and fenbutatin-oxide (Ven-
programs, although it also means that there is no dex) are examples of tin compounds that are pri-
residual toxicity to the pests. marily acaricides and fungicides. Plictran (cyhexatin)
No resistance to oils has been reported in pest was introduced in 1967 and was widely used for
arthropods, including mites, perhaps because oils many years before resistance developed in spider
have a relatively short residual activity. Oils are easy mites. Some people had assumed that the organotins
to apply, relatively inexpensive, and safe to handle. were immune to resistance problems. The organo-
They are relatively harmless to vertebrates, dissipate tins were useful products because they were more
quickly after spraying, and leave little or no residue toxic to spider mites than to phytoseiids and thus
on crops. Oils man be used by organic farmers. were very useful in integrated mite management
A disadvantage to petroleum oils is that they programs. Fenbutatin-oxide (Vendex) is another
have little residual activity and kill only upon organotin. These products were taken off the market
contact, so thorough and precise coverage is nec- in the USA due to concerns about safety.
essary to achieve effective control. Phytotoxicity
can occur even with these narrow-range oils, espe-
cially if plants are weakened or under moisture Insecticides with Acaricidal
stress. Thus, applications should not be made dur- Activity
ing droughts, or periods of very high temperatures.
Some varieties of plants are more susceptible to Organophosphorus Pesticides
phytotoxicity than others, so caution should be
taken when using oils for the first time on a par- The organophosphates (pesticides that include
ticular crop or cultivar. Oils are not compatible phosphorus) are derived from phosphoric acid
with sulfur or some other pesticides, causing seri- and are the most toxic of all pesticides to verte-
ous phytotoxicity problems. brates. They are, in fact, related to nerve gases by
structure and mode of action. Organophosphorus
pesticides (OPs) are less persistent in the environ-
Organosulfurs ment than the organochlorines such as DDT.
Organophosphorus pesticides (such as azin-
Tetradifon (Tedion) and propargite (Omite, Komite) phosmethyl, parathion, ethion, demeton, dimethoate)
are organosulfurs. These products contain sulfur as function by inhibiting important enzymes (cholin-
a central atom with two phenyl rings. Tedion is par- esterases) in the nervous system. Acetylcholine is the
ticularly toxic to mites, but has very low toxicity to chemical signal that is carried across synapses (where
insects. Organosulfurs are often ovicidal as well as the electrical signal is transmitted across a gap to a
toxic to active stages. Propargite was used for many muscle or another neuron. After the electrical signal
years (more than 20) and appeared to some to be (nerve impulse) has been conducted across the gap
immune to the development of resistance in spider by acetylcholine, the cholinesterase enzyme removes
mite populations. However, propargite resistance the acetylcholine so the circuit wont be kept on.
has now developed in many populations of spider When OPs poison an organism, the OP attaches to
mites around the world. Propargite is less toxic to the cholinesterase so it cannot remove the acetylcho-
beneficial phytoseiid predators than to pest spider line. The circuits then remain on because acetylcholine
mites, and thus could be used in integrated mite accumulates. This gives rise to rapid twitching of the
management programs, although at high concen- voluntary muscles and to paralysis, which is can be
trations it also is toxic to phytoseiid predators. lethal if it persists in the vertebrate respiratory system.
A Acaricides or Miticides

Not all OPs are highly toxic to vertebrates; Pyrethroids

if the phosphorus is modified by esterification
(adding oxygen, carbon, sulfur and nitrogen), six Many of the pyrethroids have acaricidal activity.
different classes of OPs can be produced. Some of Some (such as bioresmethrin, fenpropathrin and
these are relatively safe to vertebrates, such as mala- bifenthrin) are considered effective acaricides.
thion. The use of most OPs is being eliminated in Unfortunately, pyrethroids usually are very toxic
the USA due to the Food Quality Protection Act. to beneficial arthropods, including phytoseiid
predators. These detrimental effects can be very
long lasting because the residues persist a long
Carbamates time. Few have been found useful for integrated
mite management programs for this reason. Labo-
Carbamates (aldicarb, carbofuran, methomyl, ratory selection of phytoseiids ( Amblyseius falla-
propoxur) are derivatives of carbamic acid. The cis, Metaseiulus occidentalis, Typhlodromus pyri)
mode of action of carbamates is to inhibit cholin- for resistance to two pyrethroid insecticides has
esterase. The carbamates were introduced in the been successful. The pyrethroid-resistant strains
1950s. Carbaryl (Sevin) is one of the most popular were developed for use in apple pest management
products available to home gardeners for control- programs using both laboratory and field selec-
ling a variety of insect pests and has low mamma- tion methods.
lian oral and dermal toxicity. Methomyl (Lannate)
and aldicarb (Temik) are examples of other
carbamates. Pyrroles
Sevin is well known to induce outbreaks of spi-
der mites after applications are made to control Pyridaben is a novel pyrrole pesticide that works as
other pests. The outbreaks are due to two factors; (i) a mitochondrial electron transport inhibitor to
Sevin kills phytoseiid predators and other natural block cellular respiration, causing pests to become
enemies of spider mites, and (ii) it stimulates repro- uncoordinated and die. Can be used on both insects
duction of spider mites, a process called hormoligo- and mites.
sis. Even very low doses of Sevin appears to act like a
hormone to stimulate reproduction of the two-spot-
ted spider mite Tetranychus urticae. It is likely that Other Acaricides
the use of carbamates also will be eliminated or
greatly reduced in the USA due to the Food Quality Azadirachtin
Protection Act.
This is a triterpenoid extracted from the seeds
of the neem tree Azadirachta indica. Extracts
Formamides include a combination of compounds, the pro-
portion of which vary from tree to tree. Such
Formamides include chlorodimeform (Galecron or variability in this natural product makes it diffi-
Fundal), amitraz, and formetanate (Carzol). These cult to predict the precise effect of the product
products are effective against the eggs of Lepi- when extracted by local people. Commercial
doptera and also against most stages of mites and products may be more consistent in their effect
ticks. The mode of action of these products is because they have been tested to confirm their
unclear, but thought to be due to the inhibition of quality and are blended to achieve a consistent
monoamine oxidase, which results in the accumu- product. Azadirachtin blocks the action of the
lation of compounds called biogenic amines. molting hormone ecdysone.
Acaricides or Miticides
A 19

Avermectin in spider mite populations in several locations

around the world, including Europe and Australia.
Avermectin is a natural product containing a macro-
cyclic lactone glycoside that is a fermentation product
of Streptomyces avermitilus, which was isolated from
soil.Avermectin is actually a mixture of two homologs,
This is a phenoxypyrazole and has been evaluated
both of which have biological activity. Avermectin has
under the trade name Pyranica in Australia, where
insecticidal and acaricidal properties and is closely
it was shown to be useful in integrated mite
related to ivermectin, which kills nematodes.
management programs in apples because it is selec-
At appropriate rates, abamectin is less toxic to
tive (relatively nontoxic) to phytoseiid predators.
beneficial phytoseiids than to spider mites; it para-
lyzes active spider mite stages, but is not toxic to
eggs. Avermectin has translaminar activity (mean- Acaricides and Fungicides
ing it is taken up by the plant tissue and subsequently
by spider mites feeding on the planttissues), but has Benomyl is a carbamate that has been used pri-
a short residual toxicity to phytoseiids. marily as a fungicide, but also has acaricidal prop-
Resistance to this product has been reported in erties. Benomyl is interesting because it acts as a
some populations of spider mites. A resistant strain sterilant of phytoseiid predators. Adult phytoseiid
of M. occidentalis was obtained after laboratory females treated with benomyl survive, but they do
selection, suggesting that resistance mechanisms not deposit eggs. This product apparently disrupts
may be present in field populations. spindle fiber formation in cells and interferes in
The mode of action of avermectin involves the synthesis of DNA, resulting in females that are
blocking the neurotransmitter gamma-aminobutyric unable to reproduce.
acid (GABA) at the neuromuscular junction. Mites
that are exposed to abamectin become paralyzed
and, although they do not die immediately, the para- Resistance in Mites
lyzed mites do stop feeding.
Resistance to pesticides is an increasingly serious
problem around the world. Resistance to one or
Clofentezine and Hexythiazox more pesticides has been documented in more
than 440 species of insects and mites. Spider mite
These are very interesting growth regulators of and tick species have readily developed resistance
mites; they kill eggs (ovicides) of spider mites, but to all classes of pesticides.
not the active stages of spider mites. The products Resistance is a decreased response of a popula-
have different chemistries, but both are nontoxic to tion of animal to a pesticide or control agent as a
phytoseiid mite eggs or active stages! In fact, the result of their application. It is an evolutionary or
phytoseiid mite Metaseiulus occidentalis can be genetic response to selection. Tolerance is an innate
fed a diet consisting solely of spider mite eggs ability to survive a given toxicant dose without prior
thathave been killed with these products and the exposure and evolutionary change. Cross resistance
predator females reproduce and their progeny is a genetic response to selection with compound A
develop normally. This selectivity makes the products that generates resistance to both compound A and
particularly useful for integrated mite management other compounds (B and C). Multiple resistance is
programs because predators can be maintained resistance to different compounds due to the coex-
while suppressing spider mite populations. Unfor- istence of different resistance mechanisms in the
tunately, resistance to these products has developed same individuals. Multiple resistances usually are
A Acaricides or Miticides

generated by sequential or simultaneous selection Slide Dip Bioassays

by more than one type of pesticide. Slide dip bioassays of adult female spider mites and
phytoseiids have been proposed as a standard
method for assessing resistance or tolerance. This
Methods for Evaluating Resistance method involves placing adult female on their backs
on to double-sided sticky tape applied to glass
There are a variety of methods available for assess- microscope slides and dipping the slides into a spe-
ing resistance to pesticides in mites. The test method cific pesticide concentration. This method has the
chosen will depend upon the goals of theresearcher. virtue of being relatively rapid and easy to conduct.
Each method has strengths and weaknesses. However, measuring toxicity to adult females after
Resistance is a genetically-determined change 24 or 48 h is not an appropriate assay for many pes-
in the ability to tolerate a pesticide. Therefore, one ticide types (for example ovicides, growth regula-
must have at least two different populations to test tors). Also, the results probably bear little relation to
one that is putatively resistant and one that exhib- the field toxicity of the product. It is very likely that
its the normal, wild type response. Unless these two many products are much more toxic to the mites
populations can be compared under identical labo- using this assay than they would be under field con-
ratory conditions, it is difficult to document resis- ditions, where mites can feed and move around and
tance because historical data are of questionable coverage is rarely complete, so this method may give
value in assessing whether a population is resistant. no information about whether the resistance level
This is because it is very difficult to conduct identi- induced is relevant to field concentrations used.
cal bioassays in two different laboratories, even
when attempts are made to use the same methods.
Leaf Dip or Leaf Spray Bioassays
Small differences in techniques can result in very
Leaf dip or leaf spray bioassays involve placing
large differences in toxicity data. For example, spi-
mites on leaf disks, which are then sprayed or
der mites tested on smooth leaves may respond
dipped into a specific concentration of pesticide.
very differently than spider mites tested on the same
This type of bioassay provides an exposure that is
plant species but on a variety with hairy leaves.
more similar that the mites would experience
Small differences in formulations and temperature
under field conditions and it is possible to mea-
also influence responses of mites to pesticides. Small
sure survival, fecundity, and ability to successfully
differences in age or feeding status also influence
develop on pesticide residues.
toxicity responses. Most conclusions about resis-
tance should be based on comparative data obtained
Whole Plant Bioassays
by the same researcher under identical conditions.
This approach, which involves spraying the entire
The apparent failure of a product to control a
plant, is very realistic, unless the plants (and pesticide
mite population under field conditions is NOT ade-
residues) are not exposed to sunlight or rain.
quate evidence for resistance. Field failure is a reason
to investigate further, but field failures can occur
for a variety of reasons that have nothing to do with Field Tests
resistance. Failures could occur because the pesti- Field trials are the most realistic method for assess-
cide applicator may have mixed the product improp- ing resistance, but it can be difficult to determine
erly, coverage may have been inadequate, the pH of why the predators or spider mites died (did other
the water used to mix the pesticide could have tolerant predators fly in and eliminate the pest?). If
altered the toxicity of the product, and the product adequately replicated over time and space, field trials
could have been old or degraded due to improper provide very relevant information. The relevance of
storage. application method (high or low volume), coverage,
Accessory Gland
A 21

and droplet size can be assessed. Unfortunately, field the University of California-Davis, Oregon State, Michi-
trials are the most expensive to carry out so the gan State, Cornell and the University of Idaho.
Roush RT, Tabashnik BE (1990) Pesticide resistance in
methods described above are often used to save time arthropods. Chapman and Hall, New York, NY
and funds.
Insecticide Toxicity Accessory Cell
Insecticide Formulation
Detoxification Mechanisms in Insects A wing cell not normally present in the taxon.
Pesticide Resistance Management
Pesticide Application Accessory Circulatory Organ
Although the dorsal vessel (heart) is normally con-
References sidered to be the organ responsible for blood circu-
lation in insects, sometimes small sac-like structures
Cranham JE, Helle W (1985) Pesticide resistance in Tet-
ranychidae. In: Helle W, Sabelis MW (eds) Spider
are located at the base of appendages (antennae, legs,
mites: Their biology, natural enemies and control, vol. wings). These structures are capable of contractions
1B. Elsevier, Dordrecht, The Netherlands, pp. 405421 independent of the dorsal vessel, and assist in circu-
Croft BA (1990) Arthropod biological control agents and pes- lation. This is also called accessory pulsatile organ.
ticides. Wiley-Interscience, New York, NY
Davidson NA, Dibble JE, Flint ML, Marer PJ, Guye A
(1991) Managing insects and mites with spray oils. IPM
Education and Publications, Statewide IPM Project, Accessory Gland
Publication 3347. University of California, Division of
Agricultural Natural Resources, Oakland, California. A gland associated with the male or female repro-
EXTOXNET: This site provides infor-
mation about pesticides, including concepts in toxicology
ductive system, and producing substances associ-
and environmental chemistry. It is a cooperative effort of ated with the sperm or eggs, respectively (Fig. 7).
Male accessory glands produce such substances to

lateral oviduct
median oviduct

Accessory Gland, Figure 7 Diagram of the female reproductive system, as found in Rhagoletis (Diptera)
(adapted from Chapman, The insects: structure and function).
A Accessory Vein

facilitate sperm transfer, as a barrier to further Sampling Arthropods

insemination, as a means of altering female behav-
ior, and as a means of providing nutrition to the
female. Females produce substances for packaging Acephalous
their eggs, adhering them to a substrate, and provid-
ing a protective coating over the eggs. The condition of lacking an apparent head. This
term is usually applied to certain flies and wasps
that lack a well-defined head.
Accessory Vein
An extra branch of a longitudinal vein. Such veins Acerentomidae
normally are designated by a subscript of a.
Wings of Insects A family of proturans (order Protura).
Accessory Hearts
Pulsatile, sac-like organs that assist in circulation of
the hemolymph into appendages such as the anten- The synaptic transmitter substance found in the
nae, wings, and legs. insect central nervous system. When released into
the synaptic cleft, it is bound to a receptor, depo-
Accidental Host larizing the postsynaptic membrane and stimulat-
ing nervous excitation.
A host in which the pathogenic microorganism
(or parasite) is not commonly found.
Acetylcholine Esterase
Accidental Species An enzyme that breaks down acetylcholine after it
is released into the synaptic cleft of insect neurons.
Species that occur with a low degree of consistency Interference with acetyl choline esterase, as by
in a community type. Such species are not useful exposure to some insecticides, results in prolonged
for community definition. stimulation of the nerves.

The adaptation of an organism s physiological
A family of insects in the superfamily Fulgoroidae
responses to existing environmental conditions. A
(order Hemiptera). They sometimes are called
nearly equivalent term is acclimatization, though
acclimation may be a more rapid or laboratory-based
phenomenon, whereas acclimatization is a long
term, field-based phenomenon.
A family of insects in the superfamily Coccoidae
A measure of the closeness of an estimate to the (order Hemiptera).
true mean or variance of a population. Bugs
Acoustical Communication in Heteroptera (Hemiptera: Heteroptera)
A 23

Acoustical Communication in a stationary portion are needed. The movable

Heteroptera (Hemiptera: part is called the plectrum or scraper. The sta-
Heteroptera) tionary portion may be called the stridulitrum,
file, strigil (strigile, strigilis) or lima (pl., limae).
marta goula The stridulitrum is typically striated or finely
University of Barcelona, Barcelona, Spain tuberculated, and the plectrum is a structure with
a well-defined lip or ridge, tubercles, or provided
Acoustic signaling is found in many hemipteran with spines.
families. It serves a variety of purposes, particu- Different parts of the body may be involved
larly defensive behavior such as repelling poten- to function as the stridulatory device. The fore-
tial predators and signaling alarm or distress, wing edge is most commonly used as a stridu-
but also for species spacing within a particular litrum (file), while the hind femur is the most
habitat, reproduction, and coordination of group usual structure used as a plectrum (scraper).
actions. Vibratory signals for reproductive pur- Other stridulitrum may be located in the head,
poses may be produced by males and/or females, associated with the mouth (labium, maxillary
leading toaggregation to mate attraction, court- plate), the thorax (propleurum, metapleuron,
ship and copulation. Non-receptive females prosternal groove), the wings (metathoracic wing
may sing to reject copulation (male-deterring vein, hypocostal lamina or articulatory sclerite,
stridulation), as in the subfamily Triatominae underside of clavus), the legs (forecoxa, mesotro-
(Reduviidae) and in Pentatomidae (e.g., Nezara chanter, tibia, femora), and the abdomen (sternum,
viridula). the connexival margin, posterior margin of the
The vibratory signals produced by many pygophore). There are also a variety of locations
insect species cannot be heard by the human ear for the plectrum or scraper: the rostrum, the legs
because the sounds are low frequency and gener- (forecoxal cavity, coxal peg, hind tibia, and fore-,
ally transmitted by mechanical vibrations through middle or hind femur) or the abdomen. Some
the substrate, and not by the air. The study of stridulatory devices are present in several fami-
acoustical communication has greatly progressed lies, whereas others are only known from a few
in accordance with improvement of recording and genera or even a single species.
analyzing equipment, including the necessary Examples of stridulation devices are that of
computer software. the Corixidae (spinose area inside the front femur
against the clypeus, genitalia against abdomen
segments), Scutelleridae (wart-like, toothed tuber-
cles in the hind tibia against the femur), Reduvii-
Production of Vibrational Signals dae (tip of the labium against a cross-striated
and Songs furrow (Fig. 8) in the prosternal groove), some
Pentatomoidea and Lygaeiodea (dorsal abdominal
Vibrational signals and songs are produced by files against teeth on the under-sides of the hind-
stridulation (stridulatory device, stridulatory wings), some Miridae, Lygaeidae, Largidae and
organs), by body vibration, or by a simple tymbal Alydidae (hind femur against forewing edge) and
mechanism. Stridulation occurs widely in Het- some other Pentatomoidea (tubercles on the hind
eroptera, and is the act of producing sound or femora against strigose regions on abdominal
vibration by rubbing together certain body parts. sterna).
The first systematic survey of sound-producing Morphological differences in the stridulatory
devices in the Heteroptera was that of Handlirsh device may, in some cases, be related to differences
in 1900.To stridulate, usually both a movable and in the songs emitted by either males or females, as
A Acoustical Communication in Heteroptera (Hemiptera: Heteroptera)

A tymbal is formed by abdominal tergal

plates fused together and which vibrate over a
hollow chamber within the abdomen. The tymbal
is activated by muscular contractions and pro-
duces body vibrations that are low frequency.
Tymbals have been found in Piesmatidae, Pen-
tatomidae, Acanthosomatidae, Cydnidae, Lygaei-
dae, Coreidae and possibly Reduviidae, and
similar vibration-producing mechanisms have
been found in Plataspidae and Rhopalidae. Dif-
ferences exist about precise abdominal parts
and muscle contraction mechanisms among the
tymbals of different families. For example, the des
Acoustical Communication in Heteroptera cription of the N. viridula tymbal follows: The
(Hemiptera: Heteroptera), Figure 8 The stridulatory first and second abdominal tergites are fused into
apparatus in Reduviidae: the tip of the labium a forward-backward movable tymbal-like plate
is rubbed against a cross-striated furrow in the that is loosely fixed, anterior and posterior, to the
prosternal groove. thorax and to the third abdominal tergum, by a
chitinous membrane, and more firmly, laterally,
occurs in the burrower bugs (Fig. 9) Scaptocoris to the pleurites. Longitudinal and lateral com-
castanea and S. carvalhoi. In S. castanea, males pressor muscles contract synchronously and in
have a longer stridulitrum than females. However, phase with these vibratory waves.
in S. carvalhoi the stridulitrum length does not dif- All Heteroptera species investigated so far
fer between sexes. Instead, in S. carvalhoi the male emit low frequency narrow-band signals by body
stridulitrum has more teeth than in the female stri- vibration, and/or broadband signals produced by
dulitrum. There are no intersexual differences for stridulation. For example, in Cydnidae and Pen-
this latter morphological trait in S. castanea. tatomidae, vibratory mechanisms produce a low
Differences in the stridulatory apparatus frequency vibration (around 100 Hz), and the
maybe related to interspecific differences, with a stridulatory vibration extends up to 10 kHz.
specific diagnostic value. In the genus Triatoma
(Reduviidae), it is possible to distinguish between
T. guazy and T. jurbergi at any nymph stage or in Reception of Vibrational Signals
the adults by studying the stridulatory sulcus (stri- and Songs
dulitrum). As T. jurbergi is naturally infected with
Trypanosoma cruzi, the causal protozoan of Cha- Although sound production is found quite widely
gas disease (American trypanosomiasis), identifi- in Heteroptera, it is not common to find structures
cation of specimens along their whole life cycle is specialized for sound reception. Sound reception
of great medical importance. in Heteroptera is possible due to the presence of
In other instances, morphological differences either scolopophorous organs or tympanal organs.
in the stridulatory device do not cause differences Scolopophorous organs are mechanoceptors,
in their song patterns. For example, in Reduviidae and they occur widely in insects. They are com-
of the subfamily Triatominae, individuals of the posed of sensory sensilla (scolopodia), which may
same species have stridulatory grooves with differ- be arranged in groups, and are distally attached to a
ent inter-ridge distances, though the frequency membrane in the body wall or to the body wall
spectra and repetition rates are similar. itself. Scolopophorous organs may be located in
Acoustical Communication in Heteroptera (Hemiptera: Heteroptera)
A 25

antennae (Johnstons organ), legs (subgenual are four scolopodial organs: femoral, tibial, tarso
organ, joint chordotonal organ), thoracic pleura, or pretarsal and subgenual organs. The receptor neu-
abdominal terga. rons may be low frequency (most sensitive between
Legs are the site of sensory organs that detect 50 and 100 Hz) or high frequency sensitive (there
vibratory signals with highest sensitivity. For exam- are two types: middle frequency neurons being
ple, at the dorsal side of each leg of N. viridula there sensitive around 200 Hz, and higher frequency

a b

c d

Acoustical Communication in Heteroptera (Hemiptera: Heteroptera), Figure 9 Interspecific

differences in the stridulatory devices found in females of the burrower bugs Scaptocoris castanea
and S. carvalhoi (Hemiptera: Cydnidae). Images a and b are S. castanea; images c and d are
S. carvalhoi. Images a and c show the stridulitrum in the postcubital vein of the hind wings (scale
bar=100 ); images b and d show the central section of the stridulitrum, showing details of the teeth
(scale bar=10 ) (adapted from okl et al (2006) Physiol Entomol 31:371381).
A Acoustical Communication in Heteroptera (Hemiptera: Heteroptera)

neurons sensitive around 7001000 Hz). Also, the Substrate-borne vibrations are less costly to
Johnstons organ of N. viridula has several vibra- the emitter. Also, substrate-borne vibrations are
tory sensitive organs which respond in the fre- more far-reaching signals for intraspecific com-
quency range between 30 and 100 Hz. When munication and not easily perceived by a potential
standing on its host plant, the male of N. viridula, predator or parasitoid. Usual substrates to trans-
by different combinations of legs and antennae, mit vibrational signals are plants or soil.
may compare the vibratory signals on two branches The characteristics of plants as transmission
of the host plant, and choose the best one in order media for insect-produced vibrations have been
to locate the singing female. The most probable described, and in many respects they determine
mechanism underlying resolution of direction by signal production and the mode of reception.
vibratory cues (vibratory directionality) may be Depending on the physical properties of the host
time-of-arrival differences (perception of vibratory plant, the vibratory signals are transmitted effec-
signals by two different receptors in the insect). For tively or not. Vibrations can be transmitted all along
example, when legs of the receptor bug are sepa- the stem, but the physical properties of a plant (e.g.,
rated by 2 cm, a time-of-arrival difference between elasticity, water content) affect resonance of insect
0.125 and 0.250 ms is created, very close to that vibrations. For transmission through plants, insects
found in scorpions, where vibrational directional- commonly emit broadbanded-mixed stridulatory
ity is well known. Reduviidae also receive vibratory and vibratory signals. Higher-frequency signals
signals via legs and antennae. Fewer data are avail- produced by stridulation are less relevant for long-
able on leg vibratory receptor organs in other land distance communication through plants. However,
bug species. Among land bugs, no sensory organs narrow-band and low frequency songs are efficient
for airborne sound have been found. in long-distance communication when well tuned
Tympanal organs have been found in the to the resonant spectra of their host plants. The
mesothorax of the Corixidae, and are in contact vibrational pulse reflects when attaining both the
with the physical gill air bubble. They are able to root area and top of the plant, and reflected waves
catch airborne sounds, and to respond to stridula- travel up and down the stem several times. Reflec-
tion frequencies produced by conspecific bugs. tions change the patterns of the input signal. Abiotic
factors (temperature, rain, wind) may significantly
modify plant resonance, masking insect vibratory
Transmission of Vibratory Signals signals and thus the effectiveness of the signal.
and Songs Plant-borne vibrations seem to be important in the
success of group-living, herbivorous insects for
Independent of their mode of production, vibra- locating and remaining in a group of conspecifics,
tional signals may be transmitted by the substrate for locating food resources, and to avoid predation.
(substrate-borne vibrations) or by the air (air- Also, small insects that are not able to emit airborne
borne vibrations). The signals may travel a short sounds efficiently at low frequencies in many cases
or long distance, or travel at a low or high speed. communicate with vibratory signals transmitted
Low-frequency components are more suitable for through plants.
longer-range communication through plants. Low In the stink bug Nezara viridula (Pentatomidae)
frequency signals travel longer distances, but (Figs.10 and 11), a species which has become a
slowly; high frequency signals travel shorter dis- model for all Pentatomorpha in relationship to
tances, but quickly. Long-range vibratory songs acoustical communication, its vibratory signals
are associated with pre-mating calling and vibra- were recorded and described first as airborne
tional orientation, and close-range vibratory songs sound. However, further investigations showed that
are associated with courting rivalry and repelling. their most important mode of transmission is as
Acoustical Communication in Heteroptera (Hemiptera: Heteroptera)
A 27

substrate-borne vibrations. In N. viridula, a male of soil transmitted signals varied between 1.5 and
could perceive a female calling in a Cyperus stem 2 12.9 ms-1 at a distance of 0.5 cm.
m away from him, mechanically coupled only by
roots and the surrounding earth. Below the leg
of the singing bug, the intensity of signals was Acoustic Characteristics of
about 4 mm s1. On the bottom of the same stem (a Vibrational Signals
distance of 80 cm) it decreased to around 3 mm s1,
and at the place of the receiving bug, 200 cm away, A vibrational signal may be characterized by its
it was approximately 0.5 mm s1. temporal (pulse train duration, repetition times,
When transmitted through the soil, signals inter-pulse intervals) (Fig. 12) and spectral char-
travel a shorter distance and are more attenuated acteristics (dominant frequency). All of these
than when transmitted through a plant stem. For characteristics may be measured by the receptor
example, in Scaptocoris species (Cydnidae) the velocity insect, who may modify its behavior in response
to the message. The dominant frequency of signals
produced by the vibratory mechanism lies between
50 and 200 Hz in most Heteroptera. Between spe-
cies, songs differ in their time structure and ampli-
tude modulation of their units. On the other hand,
spectrally and temporally different pulse trains
trigger the same male behavior in N. viridula.
In Rhodnius prolixus (Reduviidae), the male-
deterring call has a main carrier frequency of about
1500 Hz, and the disturbance stridulation has a main
carrier frequency of about 2200 Hz. In Rhinocoris
iracundus (Reduviidae), low-frequency components
of carrier frequency below 200 Hz are exchanged
with frequency-modulated stridulatory components
whose dominant frequency lies between 1 and
2 kHz. In Triatoma infestans (Reduviidae), distress
songs have a peak of frequency between 700 and
800 Hz, although in other reduviids the carrier fre-
quency may reach about 2000 Hz.
In N. viridula, dominant frequencies between
80 and 150 Hz were found in songs either as air-
borne sounds, or substrate or body vibrations.
Body vibrations are around 100 Hz, and lie close
to the range of best frequency sensitivity of low
frequency receptor cells.

Acoustical Communication in Heteroptera

(Hemiptera: Heteroptera), Figure 10 Oscillograms of Specificity and Variability of
songs emitted by males and females of the southern Vibratory Signals
green stink bug, Nezara viridula (Pentatomidae)
(adapted from okl, Virant-Doberlet (2003) Annu Sounds, especially those involved in the reproduc-
Rev Entomol 48: 2950). tive process (attraction, courtship, copulation),
A Acoustical Communication in Heteroptera (Hemiptera: Heteroptera)

Acoustical Communication in Heteroptera (Hemiptera: Heteroptera), Figure 11 Laser vibrometer

recordings taken from a plant fed upon by Nezara viridula showing the pattern of recordings at
various distances (in cm) from the point of stimulation (adapted from Miklas et al (2001) J Insect Behav

can be very complex and highly species-specific,

and may be used as taxonomic characters of land
bugs. In contrast, signals are much less specific
when they provide information about enemies,
rival mates, or serve as distress (disturbance or
alarm) signals.
Females of N. viridula sing to trigger the male
approach, and to evoke emission of the male
Inter pulse trains courtship song. Females coming from popula-
Duration interval (IPI)
tions of different geographic origin emit different
calling songs, which can be differentiated by
Repetition time (RT)
males. Females of N. viridula may emit a song that
rejects copulatory attempts of males and stops
Acoustical Communication in Heteroptera their courting, andthis is also known in the redu-
(Hemiptera: Heteroptera), Figure 12 Temporal viid Rhodnius prolixus. The courtship songs of
parameters of a vibratory signal (adapted from both males and females in different populations
Miklas et al (2003) Behav Process 61: 131142). are not markedly different, but the calling songs
Acoustical Communication in Heteroptera (Hemiptera: Heteroptera)
A 29

may differ in some features and may be the

source of reproductive isolation among popula-
tions. Nezara viridula produces four different Oscilloscope
Video camera VCR
species and sex-specific songs, and two of them
play a vital role in mate location. There is song
variability within populations (inter-individual
variability). To assess those differences, the tem-
poral song (pulse train duration, repetition
times, inter-pulse intervals) and spectral song
characteristics (dominant frequency) may be Accelerometer Amplifier
measured. Males usually show a preference for
the females of their own population, although Acoustical Communication in Heteroptera
they may recognize females from other popula- (Hemiptera: Heteroptera), Figure 13Experimental
tions as potential partners. setupused to study substrate-borne signals
In Tritoma infestans, stridulation songs dif- produced by stridulation in Rhodnius prolixus
fer in their syllable durations, repetition rate and (Hemiptera: Reduviidae). The accelerometer
main carrier frequency, depending on the song generated a signal with voltage proportional to the
function. Differences come from rubbing their instantaneous acceleration of the moving object,
rostrum (scraper) at different speeds on the pros- electrical signals were amplified and monitored
ternal file. Also in Rhodnius prolixus, the different by an oscilloscope, then this information was
frequency between deterrent and disturbing stored in the sound track of a videotape. Also,
signals can be explained on the basis of a differ- the behavior of the bugs was simultaneously
ent rubbing velocity by the proboscis against the videotaped (adapted from Manrique, Schilman
prosternal stridulatory organ (Fig. 13). (2000) Acta Trop 77:271278).
In Scaptocoris carvalhoi and S. castanea
(Cydnidae), two sympatric burrower bugs, high
individual variation of the dominant frequency Corixidae
was observed in both male and female emissions
(Fig. 14). In Corixidae, males or both sexes use species-
specific sound for mate attraction and in court-
ship. The sounds are produced by stridulation,
Vibratory Signaling in the Families i.e., rubbing together specially modified parts of
of Heteroptera the body, or the partners body.

Vibratory signaling has been reported in several

Dipsocoromorpha and Leptodomorpha, but has Notonectidae
been better studied in the following families:
Veliidae (Gerromorpha), Nepidae, Corixidae, In Notonectidae, males produce species-specific
Notonectidae (Nepomorpha), Reduviidae, Miridae, courtship sounds by rubbing roughened parts of
Tingidae, Nabidae (Cimicomorpha), Aradidae, their front tibiae and femora against a special
Acanthosomatidae, Cydnidae, Pentatomidae, stridulatory region at the base of the rostrum
Scutelleridae, Tessaratomidae, Thaumastellidae, (Fig. 17). In genus Buenoa, the sound can be heard
Colobathristidae, Lygaeidae, Piesmatidae, Largi- at a distance of several meters. While next to the
dae, Alydidae, Coreidae, Rhopalidae (Pentatomor- female, but before clasping her, the sound pattern
pha) (Figs.15 and 16). Selected examples follow: can change.
A Acoustical Communication in Heteroptera (Hemiptera: Heteroptera)

Scaptocoris carvalhoi
Male Female


0.2s B
A 0.2s

Scaptocoris castancea
Male Female


0.2s 0.2s

Acoustical Communication in Heteroptera (Hemiptera: Heteroptera), Figure 14 Vibratory emissions of male

and female burrower bugs Scaptocoris carvalhoi (above) and S. castanea (below) (Hemiptera: Cydnidae).
A and B designate the two types of female song found in S. carvalhoi. Time scales are marked below
oscillograms (adapted from okl et al. (2006) Physiol Entomol 31:371381).

Reduviidae silent environment, the sound is audible at about

1 m away. A disturbance call hasbeen described
Distress (disturbance or alarm) signals in Reduvii- in the following triatomine species: Dipetalogaster
dae may be produced either as nymphs or adults maxima, Triatoma infestans, T. guasayana, T. sordida,
(males and/or females). In Panstrongylus rufotuber- Panstrongylus megistus and Rhodnius prolixus.
culatus (Reduviidae), stridulation occurs only under In the spined assassin bug, Sinea diadema,
conditions of extreme provocation. Its sound is agonistic interactions between adult females may
audible by the human ear, which is unusual among be resolved by stridulation in 33% of the cases.
stridulating Triatominae, and is similar to the Stridulating individuals retreated more often than
sound of sandpaper scraping wood. The tip of the their non-stridulating opponents, indicating that
rostrum is rubbed along the transversely ridged stridulation may be a startle mechanism employed
prosternal groove with an anterior-posterior move- by temporarily disadvantaged individuals to
ment; the return stroke (posterior-anterior) is silent. escape from encounters. Together with other signs,
Stridulation lasts for about 5 min, although the insect stridulations provide information on the identity
remains immobile when held for a longer time. In a and relative fitness of the opponent.
Acoustical Communication in Heteroptera (Hemiptera: Heteroptera)
A 31

Acoustical Communication in Heteroptera

(Hemiptera: Heteroptera), Figure 16 Phyllomorpha
laciniata (Hemiptera: Coreidae): (above) dorsal
view of pronotum with scraper (plectrum) at its
Acoustical Communication in Heteroptera
margin, (below) detail of spines of scraper (adapted
(Hemiptera: Heteroptera), Figure 15 Artabanus
from Moulet 1995 Faune de France 81).
lativentris (Hemiptera: Aradidae): (above) ventral
view of abdomen, with file (stridulitrum), (below)
hind leg with detail of scraper (plectrum) in the Tingidae
interior surface of the hind tibia (adapted from
Schuh and Slater 1995 True bugs of the world In the tingid Corythuca hewitti, vibrational sig-
(Hemiptera-Heteroptera): classification and natural naling during group movements may occur as
history. Cornell University Press, Ithaca, NY, 335 pp). groups of nymphs are attended by a female. It has
been reported that a disturbance of the leaf where
The acoustic repertoire of the ambush bug, C. hewitti aggregated caused feeding to stop and
Phymata crassipes, is quite large and may be displayed dispersal by the bugs, with occasional stopping of
by females, males or nymphs. Its vibrational songs the bugs to vibrate the abdomen in a vertical
may be emitted in response to, and alternating with, plane, a behavior followed by conspecifics.
calls from conspecifics, or even human speech or
whistle. Sound emission is related to disturbance,
interaction with other males and females, or court- Cydnidae
ship. Signals are produced by locomotory, stridula-
tory and/or vibratory mechanisms. Airborne signals In cydnids, the low species and sex specificity of
directly or indirectly stimulate vibrational receptors. pure stridulatory signals indicates that these
Bugs within a group respond to each other only via vibratory emissions may play a role in distur-
substrate, even in close proximity. bance (defensive) behavior, as in Tritomegas
bicolor. Stridulatory signals are also related to
aggregation or some other unspecific behavioral
Miridae context. Cydnid bugs engage in rival singing and
also distress (disturbance or alarm) signals, either
Although a stridulatory device has been described in as nymphs or adults (males and/or females).
several tribes and subfamilies, the functions of acous- Courtship, acceptance and rivalry songs
tical communication in Miridae are still unknown. show higher specificity and in most cases are
A Acoustical Communication in Heteroptera (Hemiptera: Heteroptera)

of the emitted signals and the uniformity of

songsindicate that calling and courtship may be
mediated by signals of other modalities. The lack
of low frequency signals may be explained by the
direct contact of the bug with soil, which mechan-
ically prevents free vibration of the abdomen.


Pentatomid bugs engage in rival singing. For

example, Nezara viridula and Rhapigaster nebu-
losa may alternate rival songs until one or both
stop singing, and in P. lituratus, males perform
rival singing. Vibrational directionality has been
demonstrated in host or prey searching in the
predatory stink bug Podisus maculiventris.
The general pattern of singing during pre-
mating behavior is similar for all Pentatomoidea.
Calling starts with the emission of the female calling
song, which triggers males to respond with calling
Acoustical Communication in Heteroptera and courtship songs, activates them to walk on the
(Hemiptera: Heteroptera), Figure 17 Male Anisops plant, and enables directional movement toward
megalops (Hemiptera: Notonectidae): (above) the female. Alternation of male and female songs
foreleg with detail of scraper (plectrum) in the may result in more or less complex duets, as is the
interior surface of the fore-tibia, (below) lateral case in N. viridula. Nezara viridula vibrates its body
view of the head with file (stridulitrum) on the as part of intersex communications (courtship,
labium (adapted from Schuh and Slater 1995 directional cue for locating the mate, mate recogni-
True bugs of the world (Hemiptera-Heteroptera): tion), which implies that substrate-borne signals
classification and natural history. Cornell arehighly species-specific. The female song causes
University Press, Ithaca, NY, 335 pp). the male to walk, to respond with the calling and
courtship songs, and to approach the source of the
roduced by low frequency body vibration and/or
p song with characteristic search behavior. In con-
by stridulation. Tritomegas bicolor produces court- trast, females show no reaction to vibratory stimu-
ship, mating, and male rivalry calls by stridulation lation and no vibrational directionality.
and body vibration. In Sehirus luctuosus, the males Insect Acoustics
courtship call is produced by body vibration, Cicada Acoustics
giving a drumming song. Two types of species- Vibrational Communication
specific male courtship songs, produced by stridu-
latory and vibratory mechanisms, have been
described. The first type triggers female response,
a species-specific agreement song. The second type
stimulates pair formation.
okl A, Virant-Doberlet M (2003) Communication with
In the group-living species of genus Scapto- substrate-borne signals in small plant-dwelling insects.
coris, the absence of low frequency components Annu Rev Entomol 48:2950
Acoustic Communication in Insects
A 33

okl A, Nardi C, Bento JMS, Hirose E, Panizzi AR (2006) predators of an unpleasant experience if predation is
Transmission of stridulatory signals of the burrower
attempted, but it is also possible that clicks function
bugs Scaptocoris castanea and Scaptocoris carvalhoi
(Heteroptera: Cydnidae) through the soil and soybean. as startle sounds, allowing escape. This latter expla-
Physiol Entomol 31:371381 nation seems unlikely, however, as caterpillars usu-
McGavin GC (1993) Bugs of the world. Blandford, London, ally move very slowly, so escape is not very likely.
UK, 192 pp
Miklas N, okl A, Renou M, Virant-Doberlet M (2003)
Acoustic Communication in Insects
Variability of vibratory signals and mate choice selectiv- Vibrational Communication
ity in the southern green stink bug. Behav Process
Reyes-Lugo M, Daz-Bello Z, Abate T, Aviln A (2006) Stridu- Reference
latory sound emission of Panstrongylus rufotuberculatus
Champion, 1899 (Hemiptera: Reduviidae: Triatominae).
Braz J Biol 66(2A):443446 Brown SG, Beottner GH, Yack JE (2007) Clicking caterpillars:
Schuh RT, Slater JA (1995) True bugs of the world (Hemiptera acoustic aposematism in Antheraea polyphemus and
Heteroptera): classification and natural history. Cornell other Bombycoidea. J Exp Biol 210:9931005
University Press, Ithaca, NY, 335 pp
Wheeler AG Jr (2001) Biology of the plant bugs. Cornell
University Press, Ithaca, NY, 507 pp Acoustic Communication in
Acoustic Aposematism (Clicking) allen sanborn
by Caterpillars Barry University, Miami Shores, FL, USA

Adult Lepidoptera are well known to perceive sound, Sound is used by a wide variety of insects for
such as the ultrasonic cries of insectivorous bats. diverse purposes. It is difficult to evolve an acous-
Some even produce sounds that are used for social tic communication system. There must be modifi-
communication. Less well known is the sound pro- cations to produce sound, transmit the sound in
duction and reception of larvae. Some caterpillars the environment and modify the sound to specific
employ vibrational signals with ants (e.g., Lycaenidae biological purposes, as well as the evolution of
and Riodinidae), communicate about space with structures that will receive and decipher the sig-
conspecifics (e.g., Gracillariidae), or detect insect nals. Arthropods are one of only two major groups
predators or parasitoids (e.g., Noctuoidea and of animals (along with the vertebrates) that have
Gracillariidae). However, clicking sounds are an evolved acoustic communication and insects are
audible sound produced by caterpillars of silk moth the primary group of arthropods to exhibit acous-
(Saturniidae) and hawk moth (Sphingidae). This tic behavior. At least ten orders of insects possess
noise has also been described as squeaking or species that produce acoustic signals.
crackling, and originates with the mandibles.
Defensive sounds are usually categorized as startle
or warning behaviors; startle sounds warn a poten- Sound
tial predator, causing momentary hesitation and
escape of the potential prey, whereas warning sounds Sound is generated by causing the repeated com-
alert a potential predator that it is inadvisable to pression and rarefaction of an elastic medium. The
attack. Associated with the clicking sound is regurgi- waves of compression produced will then travel
tation behavior, and both actions follow disturbance through the medium to the receptor or target. The
of the larva. Regurgitant usually is adverse to preda- medium can be air, water or a substrate so sound
tors, and is a widespread defensive behavior among also encompasses vibrations. As the sound energy
insects. Thus, clicking is thought to warn potential travels through the environment, it is modified
A Acoustic Communication in Insects

by various interactions with the components of to decrease the acoustic short circuiting of a small
that environment. In addition, sound waves are dipole sound source, mole crickets (Gryllotalpidae)
reflected as they move through any environment. dig burrows which act as loudspeakers that amplify
This bending of sound waves can initiate interac- and direct their calls skyward, and the hollow
tions among the waves. The signal begins to dete- abdomen of male cicadas (Cicadidae) acts as a res-
riorate as a result of these wave interactions and onating structure to amplify the acoustic signals
with the variations in the signal initiated due to produced by the tymbals.
temperature and humidity. Acoustic energy is also The sound producing systems of insects are
lost as it is absorbed by structures in the environ- generally vibrating structures. These structures
ment and as a result of spherical spreading from are necessary because muscles cannot contract
the sound source. The loss of signal integrity due rapidly enough to produce high frequency sounds.
to these environmental interactions is termed the The sound generating structures vibrate multiple
excess attenuation of the signal. The amount of times for each muscle contraction, so the sound
excess attenuation varies with the habitat and the producing systems act as frequency multipliers.
original signal properties. Once the vibrations are initiated, the sound can
To complicate matters, the small size of insects be modified by attached resonant structures.
requires that they use high frequencies (Fig. 18) to There are many different schemes that can be
transfer energy efficiently to the signal. This will employed to describe insect calls. The variety of
limit the range of a given signal because higher fre- sound production mechanisms has led to a variety
quencies attenuate more rapidly in the environment. of terminologies. No one terminology has been suc-
To use lower frequencies, insects must modify their cessful in describing the different sound production
bodies or behaviors to increase the efficiency of mechanisms or phylogenetic relationships of insects.
sound production. For example, tree crickets (Gryl- When sounds are recorded and analyzed in any
lidae) create a baffle by inserting their body in a leaf acoustic work, each analysis of acoustic signals in



0.5 sec

B 50 msec

Acoustic Communication in Insects, Figure 18 Acoustic signal produced by the cicada Beameria venosa
(Uhler). (a) Sonogram of calling song illustrating energy distribution of the call. Middle trace is an
oscillogram illustrating the temporal pattern of the call. (b) Expanded oscillogram to show individual
sound pulses within the call.
Acoustic Communication in Insects
A 35

insects should provide a detailed description of

the terminology used to describe the signals,
and illustration of the terminology on any oscillo-
gram or sonogram in the work.

Sound Production

There are several different types of sound produc-

tion mechanisms that have evolved in insects. The
relatively low muscle contraction frequency means
that additional structures had to evolve in order to Acoustic Communication in Insects,
produce the high frequency sounds that small Figure19Stridulatory apparatus of the cicada
body size will require to generate acoustic signals Tettigades undata Torres. The file (illustrated) is
efficiently. located on the mesothorax in this group of cicadas.
The primary mechanism used by insects to The plectrum is located on the tegmina. Sound
produce sound is a stridulatory apparatus. The chi- pulses are p
roduced as the tegmina is rubbed over
tinous exoskeleton and jointed appendages of the series of m
esothoracic ridges.
insects are preadapted for modification into stridu-
latory apparati. Each stridulatory apparatus is com-
posed of a file and plectrum or scraper. The file is muscle (Fig. 20). The tymbal buckles as the tymbal
generally a row of small cuticular teeth that is muscle contracts. Sound pulses are produced when
rubbed against a ridge or blade (the plectrum) on the tymbal buckles, when individual ribs buckle
some other body part (Fig. 19). The teeth are bent and potentially when the tymbal returns to its
as they catch on the plectrum and pop forward as relaxed position. The unbuckling of the tymbal is
they release from the plectrum. The release causes assisted by the resilin within the tymbal. The reso-
the teeth to vibrate alternately compressing and nant frequency of the tymbal determines the fre-
expanding air molecules producing sound. In quency of the sound pulses produced. Additional
general, these vibrations will occur in a single structures such as the opercula, tymbal covers, var-
plane, resulting in a dipole being formed that pro- ious muscles and the abdominal air sacs can mod-
duces an asymmetrical sound field at close range. ify the sound produced. The abdominal air sacs can
The joints and exoskeleton of insects have permit- become rather large as in the bladder cicada Cysto-
ted stridulatory apparati to evolve in many loca- soma saundersii Westwood whose abdomen is so
tions on the body. There are file and scrapers found large, to resonate at a low frequency, that the male
between antennal segments (Phasmatodea), sepa- has difficulty flying. The tymbal is a common organ
rate mouthparts (Orthoptera), head and thorax in the Hemiptera and acts as the ultrasonic pulse
(Coleoptera), abdominal segments (Hymenoptera), generator in the Lepidoptera.
wings and thorax (Lepidoptera, Hemiptera), body Percussion is another mechanism of sound pro-
parts and legs (Hemiptera, Orthoptera), legs and duction used by insects. Crepitation, a clicking sound
wings (Orthoptera, Lepidoptera, Coleoptera), legs produced by the wings, is another percussion mecha-
and legs (Thysanoptera, Hemiptera), between nism in insects. The wings are clapped together or
wings (Orthoptera), and between segments on the banged on the substrate to produce a sound pulse in
genitalia (Lepidoptera, Hemiptera). A tymbal organ some Hemiptera. It is relatively rare for airborne sig-
is a specialized sound production organ. It is a nals which are generally produced as a by-product of
ribbed, chitinous membrane attached to a tymbal flight. The wings may strike each other or the legs
A Acoustic Communication in Insects

production in insects. This method of sound pro-

duction has been described in a number ofinsects,
most notably the Madagascan hissing cockroach
(Gromphadorhina portentosa [Schaum]), the
Deaths Head hawk moth (Acherontia atropos [L.])
and some African Sphingids (Lepidoptera).
In addition to the airborne signals produced by
acoustic insects, it has now been shown that vibra-
tional signals are also produced by sending infor-
mation through the substrate. These vibrational
signals are primarily produced by tremulation but
also can be important in deciphering anairborne
signal, particularly at close range to the sound
source. Vibrational signals have been identified in
members of the Neuroptera, Diptera, Hemiptera,
Plecoptera, Coleoptera, Orthoptera, Mecoptera,
Raphidioptera, Lepidoptera and Hymenoptera.

Acoustic Communication in Insects, Figure 20 Sound Reception

Tymbal organ of the cicada Beameria venosa
(Uhler). The internally attached tymbal muscle A receptor for acoustic signals is necessary for a
buckles the tymbal to produce a sound pulse. communication channel to exist. Because sound is
produced by changing air pressure, a modified
during flight-producing sound pulses as in the mechanoreceptor is needed to sense the acoustic
acridid grasshoppers (Orthoptera). However, spe- information. The receptors can be classified as
cialized percussive sound production systems have either a pressure detector or a particle detector.
evolved. A castanet is found on the costal margin of Pressure detectors are membranes that bend when
the tegmina in some moths (Lepidoptera) which pressure is unequal on the two sides of the detec-
produces a pure tone when struck. The acridid grass- tor. Particle detectors are long structures that move
hopper Paratylotropidia brunneri Scudder can snap when impacted by many particles moving in the
their mandibles together to generate sound. Percus- same direction. The movement of the particle
sion has been observed in Hemiptera, Isoptera, Ple- detector alternately stretches and compresses sen-
coptera, Lepidoptera, and Orthoptera. sory cells at the base of the organ.
Tremulation is the vibration of unspecialized The primary type of pressure receptor organ is
body parts to generate sound. The abdomen is a tympanum, which has evolved independently in at
often moved dorso-ventrally or laterally to send least eight orders of insects. Tympana are generally a
vibrations through the legs to the substrate. This thin membrane system stretched over a closed cav-
mechanism has been well studied in the lacewings ity. The tympanum bends away from the side of
(Neuroptera). Wing vibrations can also be used to higher pressure setting up oscillations as the sound
send information over short distances. This type waves strike the membrane. Each tympanum has a
of signal is produced by members of the Diptera resonant frequency based on its thickness, size and
(flies and mosquitoes) and Hymenoptera. shape. The resonant frequency is generally tuned to
The expulsion of tracheal air is the final the carrier frequencies of the communication chan-
and relatively rare mechanism of insect sound nel to increase the efficiency of information transfer.
Acoustic Communication in Insects
A 37

The tympana are associated with other mechanore- conditions are not optimal for other signal pathways.
ceptor organs to transduce the signal for the nervous However, sound is energetically costly to produce
system. As the tympana oscillate, vibrations are sent and advertises your position to potential predators
to various types of receptors, generally a specialized as well as potential mates. Even with these potential
chordotonal organ called a scolopidial organ, which problems, insects have evolved diverse functions for
act as the input site to the central nervous system. acoustic signals. Acoustic signals are used for a vari-
There can be elaborate structures associated with the ety of purposes in insects including sexual signaling,
tympana to transduce energy into the central ner- courtship signals, aggression, social recruitment and
vous systems such as the crista acoustica of the crick- defense.
ets and katydids (Orthoptera) and Mllers organ of A primary function of acoustic signals is as an
locusts (Orthoptera), which provides frequency dis- intraspecific communication channel. Sounds are
criminating abilities. These pressure receptors are used to attract mates and to isolate species repro-
generally found in pairs, one on each side of the body. ductively. Each species has a characteristic call that
This provides an animal with a means to determine can prevent related species from cross mating. The
the direction from which the sound originated. calls produced by individual signalers can be com-
The sensory structures for vibrational signals pared by receivers providing the opportunity to
are trichoid sensilla generally found on the feet or select a mate who is producing a call that exhibits
cerci or through specialized subgenual and metatar- specific characteristics. This is particularly true
sal organs located in the legs. The subgenual organs when callers have congregated into localized areas,
are located just distal to the femoral-tibial joint in all which is another function of acoustic signaling.
six legs to promote detection and directional hear- Mates may be selected based on the number of
ing. They are similar to tympana in that they are calls, the temporal patterns, loudness, etc. The spe-
chordotonal organs but lack the tympanal mem- cific characteristics used by a given species are
branes and tracheal air sacs of the tympanal system. usually chosen based on their ability to demon-
Particle receptors are generally specialized strate the viability of the caller. Duets between
structures found in specific groups of insects. For individuals (either intrasexual or intersexual) can
example, female fruit flies (Diptera) sense the also act as mechanisms to determine mate choice.
courtship signals produced by the male with a spe- The signals often change as potential mates
cial antennal segment called an arista. Male mos- approach. These courtship sounds are modified
quitoes and midges (Diptera) have plumose advertisement calls which provide further oppor-
antennae which detect the species-specific wing tunities for mate assessment. Acoustic signals can
beat frequencies produced by females. Johnstons also be used intraspecifically to space individuals
organ is a more complex chordotonal organ found in the environment, as aggression signals, or as
in the antennal pedicel of some Diptera and competitive signals to jam the signal of a neighbor.
Hymenoptera which is stimulated by the vibration Eusocial insects use acoustic signals as warnings,
of the antennal flagella. The long bristles and anten- to recruit a defensive response within the colony,
nae provide a mechanical advantage to the sensory and to recruit foragers to specific food sources.
cells increasing the sensitivity of the receptor. Predator deterrence is another significant
function of acoustic signals. The loud sounds pro-
duced by many insects (e.g., cicadas [Hemiptera]),
Functions of Acoustic Signals can startle a potential predator giving the insect a
chance to escape. The percussion sounds pro-
Sound has many potential benefits in that the sig- duced by acridid grasshoppers (Orthoptera) are
nals can be used day or night, can be modified thought to be defensive displays. Sound produc-
quickly, and can travel a significant distance even if tion has evolved specifically for this anti-predator
A Acrididae

function in click beetles (Coleoptera). The Greenfield MD (2002) Signalers and receivers: mechanisms
and evolution of arthropod communication. Oxford
acoustic systems that use air movement in the
University Press, New York, NY, 414 pp
Deaths Head moth (Lepidoptera), cockroaches
(Blattodea) and grasshoppers (Orthoptera) also
have anti-predator functions. Moths (Lepi- Acrididae
doptera) have evolved a sound production system
that is used to jam the ultrasonic signals of the A family of grasshoppers (order Orthoptera). They
bats that prey upon them. The arctiid moths emit commonly are known as shorthorned grasshoppers.
ultrasonic pulses as bats approach which act to Grasshoppers, Katydids and Crickets
confuse the bat as to the exact location of its tar-
get. This gives the moth a chance to escape while
the bat circles around for another attempt at Acriology
capturing the insect. These pulses may also have
an aposomatic function warning bats of a poten- The study of grasshoppers, katydids, crickets and
tially distasteful meal. their relatives (Orthoptera). This is sometimes
Insects have also evolved specialized acous- expanded to include the orders of insects related
tic receptors as a means of avoiding predation. to Orthoptera (the Orthopteroids) such as cock-
Bat-detectors have evolved independently in roaches (Blattodea), mantids (Mantodea), stick
geometrid, noctuid and hawk moths (Lepi- insects (Phasmatodea), earwigs (Dermaptera), and
doptera), lacewings, (Neuroptera), praying man- gladiators (Mantophasmatodea).
tises (Mantodea), beetles (Coleoptera), crickets, Classification
locusts and katydids (Orthoptera). These bat- Grasshoppers, Katydids and Crickets
detectors provide an early warning system for the
insects that a bat is nearby giving the insect a
chance to escape before the bat can sense an Acrobat Ants (Hymenoptera:
insect in the vicinity. Formicidae)
Sound Production in the Cicadoidea
Acoustic Aposematism A term applied to ants of the genus Crematogaster.
Drumming Communication and Intersexual Typically they are small, shiny brown or black, and
Searching Behavior of Stoneflies (Plecoptera) possess a pedicel with two nodes. They elevate the
gaster (tip of the abdomen) over the thorax or
head when alarmed. Acrobat ants usually are
found in association with wood or trees, including
the tunnels of termites and wood boring beetles.
Bailey WJ (1991) Acoustic behavior of insects, an evolution-
ary perspective. Chapman and Hall, London, UK, 225pp They are omnivorous and tend aphids. Their bite
Drosopoulos S, Claridge MF (eds) (2006) Insect sounds and can be painful.
communication: physiology, behavior, ecology and evo- Ants (Hymenoptera: Formicidae)
lution. CRC Press, Boca Raton, FL, 532 pp
Elliot L, Hersberger W (2007) The songs of insects. Hough-
Ant-Plant Interactions
ton-Mifflin Company, Boston, MA, 228 pp
Ewing AW (1989) Arthropod bioacoustics: neurobiology and
behavior. Comstock Publishing Associates, Ithaca, NY, Acroceridae
260 pp
Gerhardt HC, Huber F (2002) Acoustic communication
A family of flies (order Diptera). They commonly
in insects and anurans: common problems and
diverse solutions. University of Chicago Press, Chicago, are known as small-headed flies.
IL,531 pp Flies
A 39

Acrolepiidae Active Ingredient (A.I.)

A family of moths (order Lepidoptera). They com- The toxic component of an formulated pesticide. It
monly are known as false diamondback moths. also is known as the toxicant.
False Diamondback Moths
Butterflies and Moths
Active Space
Acrolophidae The area or space in which the concentration of
pheromone (or other behavioral chemical) is at or
A family of moths (order Lepidoptera). They com-
above threshold concentration necessary to elicit a
monly are known as tube moths.
response from the receiver.
Tube Moths
Butterflies and Moths

Active Ventilation
Although most gas exchange in insects occurs
A preoral, unsegmented portion of the body, ante- through diffusion (passive ventilation), in some
rior to the first true body segment. This is also cases it is inadequate to meet the oxygen needs
known as the prostomium. of insects, particularly large or flying insects.
Thus, muscular contractions acting via hydro-
static pressure (pressure on the hemolymph)
Acrosternite compress the trachea and air sacs to force carbon
dioxide out and allow more rapid intake of air.
The narrow marginal region at the anterior edge The exact mechanism varies among taxa, but
of a sternite. It appears to be the posterior edge of usually involves contraction of the abdomen, and
the preceding sternite, and includes the interseg- synchronization of opening and closing of the
mental fold. It is found on the abdominal sterna, spiracles.
but absent from the thoracic sterna. Abdominal Pumping

Action Threshold
A level of pest abundance that stimulates action to
protect plants from serious damage. Small spines on the wing membrane of Lepidoptera.
Economic Injury Level (EIL) and Economic
Threshold (ET) Concepts in Pest Management
Active Dispersal
This term has several meanings depending on
the taxon of insects under consideration. In
The redistribution of animals caused by their own Hymenoptera, it is synonymous with the sting,
actions such as flying or walking. The wings of an eversible hollow cylindrical structure at the
insects allow active dispersal frequently (contrast tip of the abdomen used to deliver venom.
with passive dispersal). Though derived from the ovipositor, it is not used
A Acute

to deliver eggs. Among Diptera, this term is Adaptation of Indigenous Insects

sometimes used to refer to a pointed, sclerotized to Introduced Crops
structure associated with the reproductive sys-
tem in males. Among Lepidoptera, this term wendell. l. morrill
refers to hair-like structures on the body and Montana State University, Bozeman, MT, USA
wings of primitive moths; these also are known
as microtrichia. Host range expansion, or adaptation of insects to
new crops, is a world-wide phenomenon that has
been observed repeatedly and extensively. It is
Acute particularly well documented in North America,
where forests and prairies consisting of indige-
Of short duration, characterized by sharpness or nous plants were planted extensively to intro-
severity. duced cultivated crops only after European
emigrants arrived in the eighteenth century.
Although many new insect pests were also acci-
Acute Bee Paralysis dentally introduced from Europe into Canada
and the United States, many species of native
A disease of honey bees caused by a picornavirus. insects adapted to the new crops and became eco-
Symptoms include trembling, sprawled append- nomically important pests (Table 2).
ages, and sometimes hairlessness (contrast with Prior to widespread introduction of cultivated
chronic bee paralysis). crops, some species of native insects fed on a wide
range of plants and therefore might be expected to
accept the new crops readily. Although polypha-
Aculeate gous insects such as grasshoppers (Orthoptera:
Acrididae), wireworms (Coleoptera: Elateridae),
Pertaining to the stinging Hymenoptera (suborder
and cutworms (Lepidoptera: Noctuidae) readily
Aculeata), a group including the bees, ants, and
accepted corn, wheat and other crops, not all of
many wasps.
the species within these groups became agricul-
Wasps, Ants, Bees and Sawflies (Hymenoptera)
tural pests. For example, several hundred species
of cutworms and grasshoppers are present, but
Acute Toxicity only about a dozen species in each group have
achieved regular pest status.
The toxicity of a pesticide determined after 24 h. The Other native insect species had a narrower
toxicity resulting from a single dose or exposure. host range, and therefore adapted to a more nar-
Insecticide Toxicity row range of crops, or perhaps only a single crop.
Insecticides In the south the boll weevil, Anthonomus grandis
grandis Boheman (Coleoptera: Curcullionidae),
originally fed on native plants in Mexico related to
Adaptation cultivated cotton, and dispersed northward into
the new cotton belt of the southeastern United
This term has at least two meanings: changes in States as cotton was planted extensively. In the
the form or behavior of an organism during its life, north, the wheat stem sawfly, Cephus cinctus Nor-
and natural selection of organisms in evolutionary ton (Hymenoptera: Cephidae), fed on hollow-
time. In entomology, usually the latter definition is stemmed wild grasses, and within 10 years after
intended. tillage began in Alberta, wheat was damaged from
Adaptation of Indigenous Insects to Introduced Crops
A 41

Adaptation of Indegenous Insects to introduced Crops, Table 2 Examples of American insect pests that
have adapted to introduced crops
Insects with wide host ranges Old host plant New host plant
Apple maggot, Rhagoletis pomo- Hawthorne Apple
nella (Walsh) Diptera: Tephritidae
Chinch bug, Blissus leucopterus Grasses Corn
leucopterus Heteroptera:
Western corn rootworm, Diabrot- Grasses Corn
ica virgifera virgifera LeConte
Coleoptera: Chrysomelidae
Western bean cutworm, Loxagro- Solanaceous weeds Corn, beans
tis albicosta (Smith) Coleoptera:
Colorado potato beetle, Leptino- Buffalo burr Potato
tarsa decemlineata (Say)
Coleoptera: Chrysomeliodae
Carrot weevil, Listronotus oregon- Umbelliferous weeds Carrots, etc.
ensis (LeConte) Coleoptera:
California red scale, Aonidiella Shrubs and trees Citrus
aurantii (Maskell) Hemiptera:
Boll weevil, Anthonomus grandis Malvaceous weeds Cotton
Boheman Coleoptera:
Insects with wide host ranges Old host plant New host plant
Sugar beet wireworm, Limonius Weeds, grasses Field and vegetable crops
californicus Coleoptera:
White grubs, Phyllophaga spp. Weeds, grasses Field and vegetable crops
Coleoptera: Scarabidae
False chinch bug, Nysius raphanus Weeds, grasses Field and vegetable crops
Howard Hemiptera: Lygaeidae
Tarnished plant bug, Lygus line- Weeds, grasses Field and vegetable crops
olaris (Palisot de Beauvois)
Hemiptera: Miridae
Army cutworm, Euxoa auxiliaris Weeds, grasses Field and vegetable crops
(Grote) Lepidoptera: Noctuidae
Yellow striped armyworm, Weeds, grasses Field and vegetable crops
Spodoptera ornithogalli (Guene)
Lepidoptera: Noctuidae
Redlegged grasshopper, Melano- Weeds, grasses Field and vegetable crops
plus femurrubrum (de Geer)
Orthoptera: Acrididae
A Adaptation of Indigenous Insects to Introduced Crops

Adaptation of Indegenous Insects to introduced Crops, Table 2 (Continued)

Insects with wide host ranges Old host plant New host plant
Migratory grasshopper, Melano- Weeds, grasses Field and vegetable crops
plus sanguinipes (Fabricius)
Orthoptera: Acrididae

the Canadian prairie provinces south into Mon- feeding on a plant (species richness) does not
tana and North Dakota. In the eastern United increase indefinitely, leveling off after less than 300
States the apple maggot, Rhagoletis pomonella years if there is not an increase in crop acreage.
(Walsh) (Diptera: Tep hritidae), expanded its host We can observe insects with both broad and
range to included newly introduced cultivated narrow host selection behavior expanding their
fruits, especially apples. The Colorado potato bee- host range to include introduced crop plants. This is
tle, Leptinotarsa decemlineata Say (Coleoptera: not surprising for generalist species, which feed
Chrysomelidae) originally fed on solanaceous broadly on many plants, but it is quite interesting
weeds in Mexico or the southwestern United when insects with a narrow host range adopt new
States, and quickly spread across the United States hosts. In such cases the species with a narrow host
on potatoes once it gained access to potato acre- range usually are pre-adapted to accept the foreign
age. These are just a few examples of insects accept- crops because they feed on plants in the same fam-
ing new hosts, and the same phenomenon is well ily as the introduced crop. North America possesses
documented for important tropical crops such as close relatives to nearly all the introduced crops
sugarcane and cacao, and other crops, as they were among its indigenous flora, so it is not surprising
introduced and cultivated in various locations that insects associated with the native plants would
around the world. adapt to the introduced crops. The presence of sec-
Some of the native insect populations origi- ondary plant metabolites (allelochemicals) such as
nally occurred at low levels because their host plants alkaloids, terpenoids, and cyanogenic glycosides
were relatively sparse. Large acreage of newmono- often serves to keep non-adapted insects from feed-
crop habitats therefore resulted in an abundant food ing extensively on plants, but may serve as chemical
supply, excellent survival, and eventually in pest cues or stimulants for insects that are adapted. Thus,
population outbreaks. Also, the native environ- insects that specialize on cruciferous weeds and
ments were relatively stable, and supported a wide crops are attracted to allylisothiocyanate, and
range of beneficial insects suppressed herbivorous insects that feed on cucurbitaceous weeds and crops
insects. However, soil tillage within the agricultural are attracted to cucurbitacin.
environments produced highly disturbed systems, Host selection behavior by insects is not a
and pests with high fecundity were not effectively static situation, nor is it as simple as the single-
suppressed by predators and parasitoids. chemical scenario presented above. It is constantly
Americas native insects displayed consider- evolving in response to various biotic characteris-
able plasticity in acquiring new hosts. This trend tics such as herbivory, and even to crop cultural
has been noted everywhere agriculture is practiced, practices. Some natural selection of insect strains
and we can expect the number of pests to increase may have occurred during the adaptation from
with time, and especially with the area planted to native to introduced plants. In the northern Great
each particular crop, as indigenous species adapt to Plains, wheat matures earlier in the season than
imported host plants or crops are exposed to addi- wild grasses. Therefore, after a century, adult wheat
tional potential pests in new geographic areas. stem sawflies are now active nearly a month earlier
However, species accrual occurs most rapidly soon than previously, and now are more effective in
after plant introduction, and the number of species utilizing wheat.
A 43

Changes in farming practices that have also in a continuing stream of new pest problems for
impacted populations of native insect pests in crop plants.
croplands. Originally, horses were used for farm- Host Plant Selection by Insects
ing, and oats were needed for their feed. Later, the Allelochemicals
horses were replaced by tractors, and the need for Plant Resistance to Insects
oats was reduced. Oats are resistant to wheat stem
sawflies, and when oats was eliminated from the
cropping system, the vast acreages of wheat resulted References
in a population explosion of the sawflies. More
recently, canola has been included in the Canadian Bernays EA, Chapman RF (1994) Host-plant selection by phy-
tophagous insects. Chapman and Hall, New York, NY
Prairie Provinces, and populations of grass-feeding Connor EF, McCoy ED (1979) The statistics and biology of
insects are somewhat disrupted by the presence of the species-area relationship. Am Nat 113:791833
a non-host, cruciferous crop. Other water and Kim KC (1993) Insect pests and evolution. In: Kim KC,
McPheron BA (eds) Evolution of insect pests; patterns
soil conservation practices such as alternate-year
of variation. Wiley, New York, NY, pp 325
summer fallow, strip cropping, and chemical fallow Morrill WL, Kushnak GD (1996) Wheat stem sawfly
have affected the prevalence of both pest and (Hymenoptera: Cephidae) adaptation to winter wheat.
beneficial insects. Environ Entomol 25:11281132
Strong DR (1979) Biogeographic dynamics of insect-host
Beneficial insect populations were also plant communities. Annu Rev Entomol 24:89119
impacted by tillage and cultural practices, and
changes in the chemical constituency of crop
plants. Parasitoids have complex host searching
Adaxial Surface
behavior that begins with finding plant envi-
ronments in which their hosts could occur.
The upper surface of a leaf (contrast with abaxial
Therefore, it was necessary for the parasitoids
to learn that the new crops could be sources of
hosts. In the case of the wheat stem sawfly, only
two of the known parasitoid species have cur-
rently adapted from wild grasses to wheat. Par-
asitoids may be more favored by one plant
A family of thrips (order Thysanoptera).
cultivar than another, or less favored by a crop
than a similar weed. The availability of food for
the adult parasitoid or predator, either nectar
from blossoms or extra floral nectaries or pol-
len from blossoms, is often implicated in dif-
ferential survival of beneficial insects among
A family of insects in the order Hemiptera. They
different plants.
sometimes are called pine and spruce aphids.
Overall, adaption by herbivorous insects to
new host plants is a dynamic and widespread phe-
nomenon. Though sometimes it is difficult to
determine whether it is the change in the constitu-
ency of the host plant that accounts for insect
acceptance, or it is some other factor such as wide- Adelidae
spread host plant availability that accounts for
insect abundance, it is clear that the relationship A family of moths (order Lepidoptera). They
between insects and plants is not static, resulting commonly are known as long horned fairy moths.
A Adephaga

Long horned Fairy Moths Adipohemocyte

Butterflies and Moths
A type of hemocyte, ovoid in shape and likely
secretory in function.
Adephaga Hemocytes of Insects: Their Morphology and
One of four suborders of beetles (Coleoptera), and
one of two suborders that contain numerous and
important beetles (the other is suborder Polyphaga). Adipokinetic Hormone (AKH)
It is comprised of about nine families, the principal
ones being Carabidae, Gyrinidae, and Dytiscidae. A decapeptide hormone synthesized in neurose-
Nearly all groups are predatory, and many are cretory cells of the corpora cardiaca and impor-
aquatic. tant in the regulation of lipid metabolism, and
Beetles (Coleoptera) sometimes carbohydrate or proline metabolism
and other physiological functions.
Adipokinetic and Hypertrehalosemic
Aderidae Neurohormones

A family of beetles (order Coleoptera). They com-

monly are known as antlike leaf beetles. Adipokinetic and
Beetles Hypertrehalosemic

gerd gde, heather g. marco

Adfrontal Areas University of Cape Town, Rondebosch, Republic
of South Africa
A pair of narrow oblique sclerites on the head of a
caterpillar. The adfrontal areas border the front,
The adipokinetic hormones and hypertrehalosemic
which normally is triangular, so the adfrontal areas
hormones of insects comprise a family of peptide
take on the shape of an inverted V.
hormones that primarily regulate the levels of
Head of Hexapods
energy metabolites, such as trehalose, diacylglycerol
and proline that circulate in the hemolymph. These
peptide hormones are products of neurosecretory
Adelphoparasitism neurons located in the corpora cardiaca, neuroen-
docrine glands attached to the brain. The structural
A type of hyperparasitism occurring in organization of the insect corpora cardiaca is simi-
Hymenoptera: Aphelinidae in which the males are lar to the hypothalamus-neurohypophysis of the
parasitoids of females of their own species (the vertebrate endocrine system.
females parasitize Hemiptera).

Historical Perspective
The existence of hypertrehalosemic hormones
The ability of a material such as a pesticide to stick was discovered with the observation that injec-
to a surface. tions of extracts of corpora cardiaca elevated the
Adipokinetic and Hypertrehalosemic Neurohormones
A 45

concentration of trehalose in the hemolymph of missing in S. gregaria. Subsequently, three research

cockroaches (hypertrehalosemia). Unlike verte- groups ultimately reported, in the same year, the
brates that use glucose as the major blood carbo- presence of two octapeptides from the corpora
hydrate, the hemolymph of insects generally cardiaca of the American cockroach, Periplaneta
contains the disaccharide trehalose, an -1-1- americana, that were structurally related to the
gluco-glucoside, as its major circulating carbohy- locust AKHs and the crustacean RPCH. These two
drate. In addition, the enzyme glycogen peptides were isolated on the basis of myotropic
phosphorylase in the fat body of cockroaches was or heartbeat acceleration bioassays and are referred
demonstrated to be activated when these insects to as cardio acceleratory hormones (Peram-CAH-I
were injected with an extract from the corpora and Peram-CAH-II), but they also produced
cardiaca. Subsequently, studies in locusts showed hypertrehalosemia in the cockroach and represent
that injections of corpora cardiaca extracts ele- the hypertrehalosemic hormones. These pioneer-
vated hemolymph diacylglycerols, instead of tre- ing studies, along with numerous subsequent stud-
halose, and this action was referred to as an ies, demonstrated that there are, so far, about forty
adipokinetic or hyperlipemic effect. Injections of structurally related, but distinct, peptides with
locust corpora cardiaca extracts into cockroaches adipokinetic and hypertrehalosemic effects in the
produced the hypertrehalosemic response, and insects and one in crustacean (Table 3). The name
vice versa. Hence, it was likely that the adipoki- adipokinetic hormone/red pigment-concentrating
netic hormone (AKH) of locusts and the hyper- hormone (AKH/RPCH) family was coined for this
trehalosemic hormone (HrTH) of cockroaches general family of peptides, which likely encom-
were related, or identical, peptides. passes the arthropods.
The locust adipokinetic hormone was isolated
and characterized first. It was obtained from the
migratory locust, Locusta migratoria, and its pri- Chemistry of the AKH/RPCH Family
mary structure consisted of ten amino acids. It was
designated Locmi-AKH-I according to the newest The members of the adipokinetic hormone/red
nomenclature for naming insect neurohormones. pigment-concentrating hormone family share
The amino acid composition and sequence of Loc- numerous structural features. They consist either
mi-AKH-I had a remarkable similarity to a previ- of eight, nine or ten amino acids, depending on the
ously reported red pigment-concentrating hormone insect species from which they are isolated. They
(Panbo-RPCH) obtained from the shrimp Pandalus are blocked by pyro-glutamate at the N-terminus
borealis and later found in various crustaceans. It and by an amide moiety at the C-terminus. Pre-
was shown that Locmi-AKH-I was also present in sumably, blocked termini prevent degradation of
the desert locust, Schistocerca gregaria. the neuropeptides by amino- and carboxypepti-
Subsequently, both L. migratoria and S. gregaria dase enzymes while circulating in the hemolymph.
were shown to contain a second adipokinetic hor- Aromatic amino acids, usually phenylalanine and
mone (Locmi-AKH-II and Schgr-AKH-II, respec- tryptophan, always occupy positions 4 and 8,
tively) that differed from each other by the amino respectively, but aromatic amino acids can also
acids in position 6. The two locust AKH-IIs were occupy other positions. The peptides are usually
octapeptides with sequences similar to those of neutral under physiological conditions, but a few
Locmi-AKH-I and Panbo-RPCH. A third octa- have a negatively charged aspartate at position 7.
peptide AKH (Locmi-AKH-III) was also found in Glycine is always present at position 9 as deduced
L. migratoria and a similar octapeptide (desig- from cDNA analysis of the precursor. The terminal
nated Phymo-AKH-III) occurs in pyrgomorphid glycine is converted to the amide moiety on the
and pamphagid grasshoppers, but an AKH-III is tryptophan in the octapeptides.
Adipokinetic and Hypertrehalosemic Neurohormones, Table 3 Representative sequences of adipokinetic/hypertrehalosemic peptides from vari-
ous insect orders
Order Peptide name Genus Structure
1 2 3 4 5 6 7 8 9 10
Odonata Libau-AKH Libellula, Pantala, pGlu Val Asn Phe Thr Pro Ser Trp NH2
Anaim-AKH Anax, Aeshna pGlu Val Asn Phe Ser Pro Ser Trp NH2
Psein-AKH Pseudagrion, pGlu Val Asn Phe Thr Pro Gly Trp NH2
Blattodea Peram-CAH-Ia Periplaneta, Blatta pGlu Val Asn Phe Ser Pro Asn Trp NH2
Blattidae a
Peram-CAH-II Periplaneta, Blatta pGlu Leu Thr Phe Thr Pro Asn Trp NH2
Bladi-HrTH Blaberus, pGlu Val Asn Phe Ser Pro Gly Trp Gly Thr NH2
Adipokinetic and Hypertrehalosemic Neurohormones

Mantodea Emppe-AKH Empusa, pGlu Val Asn Phe Thr Pro Asn Trp NH2
Phasmatodea Carmo-HrTH Carausius, Sipy- pGlu Leu Thr Phe Thr Pro Asn Trp Gly Thr NH2
loidea, Extatosoma
Mantophasmatodea Manto-AKH Not known pGlu Val Asn Phe Ser Pro Gly Trp NH2
Orthoptera, Caelifera Locmi-AKH-I Locusta, pGlu Leu Asn Phe Thr Pro Asn Trp Gly Thr NH2
Locmi-AKH-II Locusta pGlu Leu Asn Phe Ser Ala Gly Trp NH2
Schgr-AKH-II Schistocerca, pGlu Leu Asn Phe Ser Thr Gly Trp NH2
Locmi-AKH-III Locusta pGlu Leu Asn Phe Thr Pro Trp Trp NH2
Phymo-AKH-III Phymateus pGlu Ile Asn Phe Thr Pro Trp Trp NH2
Orthoptera, Ensifera Grybi-AKH Gryllus, Acheta, pGlu Val Asn Phe Ser Thr Gly Trp NH2
=Schgr-AKH-II Tettigonia, Decticus pGlu Leu Asn Phe Ser Thr Gly Trp NH2
Order Peptide name Genus Structure
1 2 3 4 5 6 7 8 9 10
Isoptera Micvi-CC Microhodotermes pGlu Ile Asn Phe Thr Pro Asn Trp NH2
Dermaptera =Grybi-AKH Labidura, Forficula pGlu Val Asn Phe Ser Thr Gly Trp NH2
Hemiptera/Homoptera Placa-HrTH Platypleura, Munza, pGlu Val Asn Phe Ser Pro Ser Trp Gly Asn NH2
Cacama, Magici-
cada, Diceroprocta
Hemiptera, Pyrap-AKH Pyrrhocoris, pGlu Leu Asn Phe Thr Pro Asn Trp NH2
Heteroptera Disdercus
=Peram-CAH-II Pyrrhocoris, pGlu Leu Thr Phe Thr Pro Asn Trp NH2
Corpu-AKH Corixa pGlu Leu Asn Phe Ser Pro Ser Trp NH2
Letin-AKH Lethocerus pGlu Val Asn Phe Ser Pro Tyr Trp NH2
Nepci-AKH Nepa pGlu Leu Asn Phe Ser Ser Gly Trp NH2
Neuroptera =Grybi-AKH Palpares pGlu Val Asn Phe Ser Thr Gly Trp NH2
Coleoptera Scade-CC-I Scarabaeus, Gareta, pGlu Phe Asn Tyr Ser Pro Asp Trp NH2
Scade-CC-II Scarabaeus, Gareta pGlu Phe Asn Tyr Ser Pro Val Trp NH2
Oniay-CC Onitis pGlu Tyr Asn Phe Ser Thr Gly Trp NH2
=Peram-CAH-I Leptinotarsa pGlu Val Asn Phe Ser Pro Asn Trp NH2
=Peram-CAH-II Leptinotarsa pGlu Leu Thr Phe Thr Pro Asn Trp NH2
Lepidoptera Manse-AKH Manduca, Vanessa, pGlu Leu Thr Phe Thr Ser Ser Trp Gly NH2
Bombyx, Heliothisb
Helze-HrTH Heliothisb pGlu Leu Thr Phe Ser Ser Gly Trp Gly Asn NH2
Hymenoptera Tenar-HrTH Tenthredo pGlu Leu Asn Phe Ser Thr Gly Trp Gly Gly NH2
=Schgr-AKH-II Xylocopa, Bombus pGlu Leu Asn Phe Ser Thr Gly Trp NH2
=Grybi-AKH Vespula, Vespa pGlu Val Asn Phe Ser Thr Gly Trp NH2
=Manse-AKH Apis pGlu Leu Thr Phe Thr Ser Ser Trp Gly NH2
Adipokinetic and Hypertrehalosemic Neurohormones

Adipokinetic and Hypertrehalosemic Neurohormones, Table 3 Representative sequences of adipokinetic/hypertrehalosemic peptides from vari-
ous insect orders (Continued)
Order Peptide name Genus Structure
1 2 3 4 5 6 7 8 9 10
Diptera Phote-HrTH Phormia, pGlu Leu Thr Phe Ser Pro Asp Trp NH2
Adipokinetic and Hypertrehalosemic Neurohormones

Anoga-AKH Anopheles pGlu Leu Thr Phe Thr Pro Ala Trp NH2
Tabat-AKH Tabanus pGlu Leu Thr Phe Thr Pro Gly Trp NH2
Crustacea Panbo-RPCH Pandalus pGlu Leu Asn Phe Ser Pro Gly Trp NH2

Note that the peptide in certain orders is identical. For example: Peram-CAH-I and -II of the Blattodea, Blattidae is also present in Coleoptera (Leptinotarsa); Schgr-AKH-II
of Orthoptera, Caelifera is present in Orthoptera, Ensifera and in Hymenoptera (Xylocopa, Bombus); Grybi-AKH of Orthoptera, Ensifera is also present in Dermaptera, Neu-
roptera and Hymenoptera, etc.
Heliothis is revised to Helicoverpa.
Adipokinetic and Hypertrehalosemic Neurohormones
A 49

Some of the members of this peptide family anbo-RPCH peptide which has a chromatophoro-
have additional post-translational modifications tropic effect. Panbo-RPCH has also been found in
besides the blocked termini. For example, two an insect species, the heteropteran stinkbug, Nezara
HrTH decapeptides are present in the corpora viridula, where it has an adipokinetic effect.
cardiaca of the stick insect, Carausius morosus; Phylogenetic relationships of the HrTHs have
one of these decapeptides is glycosylated and has a been proposed for the cockroaches based on mor-
unique C-glycosylation where the sugar is linked phological, behavioral and physiological charac-
to the C-2 atom of the indole ring of tryptophan. ters congruent with the distribution of the various
Another unusual modification has been found in structures of the HrTHs within the order.
an AKH of the protea beetle, Trichostetha fascicu-
laris: the corpora cardiaca contain two AKHs, one
of which is an octapeptide with a phosphothreo- Physiological Actions
nine at position 6.
The relationships between individual AKH The general physiological action of the adipoki-
and HrTH peptides and insect species are complex. netic and hypertrehalosemic hormones in insects
There are no clear rules concerning which peptide is to elevate the hemolymph metabolites that are
occurs in which order of insects. Several species used by the muscles and other tissues as a source
within an order may share the same peptide and of energy, regardless of the nature of the metab-
have other species-specific sequences, and the same olites. This is accomplished by stimulating the
peptide may be present in species of different fat body, which is the hormones target tissue, to
orders. As described above, the two locust species, convert its stores of triacylglycerides or glycogen
S. gregaria and L. migratoria, share an identical to diacylglycerides or trehalose, respectively, or
decapeptide (Locmi-AKH-I); each species pos- to synthesize proline. The diacylglycerides, tre-
sesses a second, unique octapeptide (Locmi- halose or proline are released from the fat body
AKH-II; Schgr-AKH-II); and L. migratoria contains to increase their respective levels in the hemo-
a third octapeptide (Locmi-AKH-III) that does not lymph. The same peptides that elevate diacylg-
have a complement in S. gregaria. Cockroach spe- lycerides in locusts elevate trehalose when
cies of the families Blattellidae and Blaberidae administered to cockroaches, and vice versa, and
share a single hypertrehalosemic decapeptide hor- the hormones of locusts and cockroaches elevate
mone (Bladi-HrTH), whereas cockroaches of the proline in the Colorado potato beetle. The deci-
family Blattidae contain two octapeptide hormones sion as to whether lipid-, carbohydrate-, or pro-
(Peram-CAH-I and -II). In addition, there is over- line-mobilizing pathways are activated is a
lap between orders. Grybi-AKH is present in cer- species-related function of the enzyme composi-
tain crickets and in species of Neuroptera, tion in the fat body.
Dermaptera and Heteroptera. Peram-CAH-I and Muscular activity for animal locomotion can
-II of the blattid cockroaches are also found in the involve either long-term or short-term events.
Colorado potato beetle, Leptinotarsa decemlineata, Long-term activities might entail sustained, non-
and Peram-CAH-II is shared with the heteropteran emergency actions such as migration or persistent
bug, Pyrrhocoris apterus. Whereas Peram-CAH-I searching for food, mates or shelter. Short-term
and -II mobilize glycogen from the fat body of the activities might consist of local searching activi-
cockroach to increase hemolymph trehalose, the ties but may also require immediate, brief, emer-
same or similar peptides increase hemolymph pro- gency responses such as evading attack by a
line in beetles to serve as the major flight substrate. predator or defending a breeding territory. Long-
Unlike the complex situation in insects, the term events require a steady supply of energy
crustaceans apparently possess only the single metabolites, whereas short-term events may be
A Adipokinetic and Hypertrehalosemic Neurohormones

brief but intense, and, if successful, they can be By contrast, insects such as cockroaches,
followed by a period of recovery to replenish bees and flies use only carbohydrate (trehalose)
exhausted metabolites. as the primary source of energy for muscular
The adipokinetic hormones are often involved activity and locomotion. These species do not
in prolonged, constant muscular activity such as migrate and lack the adipokinetic response, but
migration. This is characteristically true for the they are faced with emergency situations of pred-
locusts whose migratory behavior has been ator evasion, and in such cases, the hypertreha-
described since biblical times. Migration is a sus- losemic hormone mobilizes trehalose in response
tained flight activity that uses muscular oxidation to the emergency. However, injections of hyper-
of fatty acids to produce energy, since fatty acids trehalosemic hormone show that significant ele-
deliver more energy per mole than carbohydrates. vation of the hemolymph trehalose may take as
However, carbohydrate serves as the major source long as 1030 min. This delay in elevating hemo-
of muscular energy during initial flight, and lipid lymph carbohydrate is too long to significantly
becomes the major source for energy as flight per- assist the insect in evading capture. Furthermore,
sists and becomes sustained. Based on differing the open circulatory system of insects does not
physiological effects, it is speculated that the three efficiently direct circulating metabolites to the
AKHs may exert different regulatory actions on muscles in the manner of the closed circulatory
metabolite mobilization and use during the differ- system of vertebrate animals. Energy metabolites,
ent stages of migration. Locmi-AKH-II is likely to such as trehalose, must constantly be maintained
be the major carbohydrate-mobilizing hormone at high levels in the hemolymph to meet urgent,
that provides trehalose for initial flight; Locmi- immediate demands. Hence, the role of the
AKH-I is the major hormone responsible for fat hypertrehalosemic hormone appears to be to
mobilization during sustained flight and Locmi- replace depleted hemolymph trehalose and main-
AKH-III may be responsible for regulating energy tain it at high levels. Maintenance of high treha-
metabolism during rest. Furthermore, during lipid lose levels allows the insect to make quick
mobilization, AKH performs several distinct but responses to elude capture that may require only
related actions. In the fat body, AKH activates seconds, or at most, several minutes to conclude.
lipase for triacylglyceride degradation; this is If the insect is successful at escape, the hormone
achieved by binding of the AKH to a G-protein stimulates the degradation of fat body glycogen
coupled receptor at the cell membrane, activation to restore the high trehalose levels by activating
of adenylate cyclase resulting in the second mes- specifically the enzyme glycogen phosphorylase
senger cAMP which, in conjunction with Ca2+, is after the hormone has bound to a G-protein cou-
responsible for lipase activation. In the hemo- pled receptor on the membrane of a fat cell and
lymph, AKH increases the lipid-carrying capacity had activated a phospholipase C, resulting in the
of lipophorins (proteins) resulting in increased production of inositol trisphosphate and the
amounts of low-density lipophorin for shuttling release of Ca2+ from internal stores (influx of
lipids from the fat body to the muscles. At the flight external Ca2+ is also activated by HrTH) which
muscle level, AKH increases the rate of lipid oxi- sets in motion a cascade of activation of kinases
dation. Recent research on a number of terrestrial and, finally, glycogen phosphorylase. Removal of
and aqueous heteropteran bugs that have various the hypertrehalosemic hormone does not affect
feeding patterns (plant sap sucking, predators, the ability of such insects to be active for the
obligatory hematophagous), also established a lip- short term (several minutes), but after exhaus-
id-based activity (flight and/or swimming) metab- tion, lengthens their recovery time.
olism that is regulated by the respective AKHs of Tsetse flies and various beetle species fuel
these insects. their flight metabolism by the partial oxidation of
A 51

proline and the production of alanine. For con- individual biology of the insect species in which
tinuous flight or replenishment of proline reserves they are found. They display a unique relation-
in the fat body a unique system exists in these ship with their target tissue in that the hormone
insects to synthesis proline: the respective AKHs carries the endocrine message to the target tissue
activate a lipase in the fat body and the fatty acids (fat body) to mobilize energy stores, but the tar-
that are liberated from triacylglyerols undergo get tissue determines which metabolic pathways
-oxidation, and the resulting acetyl CoA units are are activated depending on the biology of the
used in conjunction with alanine to synthesize species. It is this biology that determines the
proline. Alanine, which is derived from the partial nature of the muscular activity (prolonged
oxidation of proline, is re-used for proline synthe- migration; briefpredator evasion) and its meta-
sis and can be viewed as a shuttle system for the bolic need for consuming carbohydrates, lipids
transport of acetyl units. or proline as a source of energy.
Although the mobilization of energy for
flight and other metabolically intense situations
is likely the major function for the adipokinetic
and hypertrehalosemic hormones, the hormones
exhibit pleiotropic actions. The hypertreha-
Beenakkers AMT (1969) The influence of corpus allatum and
losemic hormones were isolated originally based cardiacum on lipid metabolism in Locusta. Gen Comp
on their cardioacceleratory action on the heart. Endocr 13:492
This is a logical action for the hormone since ele- Fernlund P, Josefsson L (1972) Crustacean color-change hor-
mone: amino acid sequence and chemical synthesis.
vated heartbeat rate would facilitate distribution Science 177:173175
of the energy metabolites throughout the body Gde G (1989) The hypertrehalosaemic peptides of cock-
and assure their ready access to the muscles. In roaches: a phylogenetic study. Gen Comp Endocr
keeping with the stimulatory action of AKH on 75:287300
Gde G (2004) Regulation of intermediary metabolism and
lipid degradation in locusts, lipid synthesis is water balance of insects by neuropeptides. Annu Rev
inhibited by AKH. Other, less well characterized Entomol 49:93113
actions include: inhibition of RNA and protein Mayer RJ, Candy DJ (1969) Control of haemolymph lipid
concentration during locust flight: an adipokinetic hor-
synthesis related to vitellogenesis in locusts and
mone from the corpora cardiaca. J Insect Physiol
crickets, and the stimulation in cockroaches of 15:611620
the oxidative capacity of mitochondria during fat Steele JE (1961) Occurrence of a hyperglycemic factor in the
body maturation, and of gene expression for a fat corpus cardiacum of an insect. Nature 192:680681
Stone JV, Mordue W, Batley KE, Morris HR (1976) Structure of
body cytochrome P450 related to lipid oxidation. locust adipokinetic hormone, a neurohormone that regu-
These latter actions by the hormones may be lates lipid utilization during flight. Nature 263:207221
equally important as their effects on mobilization Vroemen SF, van der Horst DJ, van Marrewijk WJA (1998)
of energy metabolites, but they are poorly eluci- New insights into adipokinetic hormone signalling. Mol
Cell Endocrinol 141:712
dated because of insufficient research, and they
cannot yet be placed into perspective as to their
physiological significance. Other actions in which
AKHs seem to be involved are an enhanced acti- Adjuvants
vation of the locust immune system and, possi-
bly, in the activation of an antioxidant protection Chemicals added to insecticides to improve their
mechanism in potato beetles. effectiveness. Examples of adjuvants include toxic-
In summary, the adipokinetic-hypertreha- ity, stability, and adhesion.
losemic-hyperprolinemic hormones constitute Insecticide Formulations
a family of peptides that are adapted to the Insecticides
A Adoption Substance

Adoption Substance Adventive

A secretion presented by a social parasite that An organism that has arrived in an area from else-
induces the host insects to accept the parasites as where. It is not native, and likely arrived as an
members of their colony. invader or accidental introduction. It is also known
Social Insect Pheromones as nonindigenous.
Invasive Species

The sexually mature stage of an animal. The adult
is usually the winged stage in insects. With rare The intromittent (copulatory) organ of the male;
exceptions, the adult does not molt again. the distal portion of the phallus. Sometimes
Metamorphosis referred to as the penis.
Abdomen of Hexapods

A pesticide used to kill adult insects. This term
often is used to describe products used to kill adult
A family of bugs (order Hemiptera).
mosquitoes. (contrast with larvicide)

Adultoid Reproductive Aeolothripidae

In higher termites, a supplementary reproductive A family of thrips (order Thysanoptera). They
that in indistinguishable morphologically from commonly are known as broad-winged thrips or
the reproductive. banded thrips.

Adult Transport
A behavior in which social insects (usually ants)
drag or carry their nestmates to a new location.
A family of bugs (order Hemiptera). The some-
This normally occurs during colony emigration.
times are called marine bugs.

Adventitious Veins
In some insects, additional wing veins are present Aeshnidae
which are neither secondary nor intercalary veins.
They usually are the result of the lining up of cross A family of dragonflies (order Odonata). They
veins. commonly are known as darners.
Wings of Insects Dragonflies and Damselflies
African Armyworm, Spodoptera exempta (Walker) (Lepidoptera: Noctuidae)
A 53

Aetalionid Treehoppers even though it causes no economic loss. The

presence of some insects in homes or on orna-
Members of the family Aetalionidae (order mental plants are examples of aesthetic pests,
Hemiptera). though in the latter case if they inhibit the ability
Bugs to market plants then the same insect can be
come an economic pest.
Economic Injury Level (EIL) and Economic
Aerial Photography Threshold (ET) Concepts in Pest Management

In pest management, photographs taken by an air-

plane or satellite that are used to identify varia- Aestivation
tions within fields/crops to help make management
decisions. A state of inactivity or curtailment of normal
activity during the summer months. Diapause

The opening in the egg surface (chorion) through
which air enters. A family of insects in the order Hemiptera. They
sometimes are called aetalionid treehoppers.
Aerosol Treehoppers

The air suspension of liquid or solid particles of

small diameter. This is a common formulation for African Armyworm, Spodoptera
flying insects or for household use where noaddi- exempta (Walker) (Lepidoptera:
tional formulation or preparation is desired. Noctuidae)
Insecticide Formulations joe c. b. kabissa
Tanzania Cotton Lint and Seed Board, Dar Es
Salaam, Tanzania
Aesthetic Injury Level
The African armyworm (Fig. 21) is a larva of a
The level of pest abundance above which aesthetic, nocturnal moth, Spodoptera exempta (Walker).
emotional, or sociological considerations require This species, although commonly referred to as
pest control actions. Economic considerations are the African armyworm, occurs rather widely in
not relevant. the grasslands of tropical and subtropical Africa
Economic Injury Level (EIL) and Economic and Asia. In Africa, where S. exempta is of major
Threshold (ET) Concepts in Pest Management economic importance, its occurrence is confined
to countries south of the Sahara: Tanzania, Kenya,
Uganda, Ethiopia, Somalia, Malawi, Zimbabwe,
Aesthetic Pest Zambia and South Africa. Outside Africa, S.
exempta has been reported from southwest Saudi
A pest which, through its presence or actions, is Arabia in the republic of Yemen, southeast Asia,
deemed objectionable and in need of elimination Australia, New Zealand and Hawaii.
A African Armyworm, Spodoptera exempta (Walker) (Lepidoptera: Noctuidae)

African Armyworm, Spodoptera exempta (Walker) (Lepidoptera: Noctuidae), Figure 21 African armyworm:
top left, adult female, top right, adult male; middle left, solitary form of larva; middle right, gregarious form
of larva; lower left, pupae in soil; lower right, eggs on foliage.

During an armyworm outbreak, larvae of exceed 1,000 per m2 over areas covering tens or
S. exempta march together in long columns, akin even hundreds of square km.
to army columns, in search of susceptible plant In Africa, S. exempta is adapted for survival on
material. This is the basis for the name army- seasonal grasslands by combining a high intrinsic
worm. When susceptible plant material is found, rate of increase with migration to places of rainfall
it is often voraciously devoured to ground level. In where grasses are suitable for survival of its cater-
typical armyworm outbreaks, larval density may pillars. Because of its capability to move over long
African Armyworm, Spodoptera exempta (Walker) (Lepidoptera: Noctuidae)
A 55

distances, hundreds and sometimes thousands of Adults emerge within 712 days after pupa-
km across national boundaries freely, S. exempta is tion and can live up to 14 days if appropriately
truly an international pest. Often times it appears fed. In their lifetime, females can lay up to 1,000
sporadically and suddenly in dense outbreaks capa- eggs. Spodoptera exempta is not known to enter
ble of causing extensive and enormous damage to into any type of diapause. This probably explains
susceptible rangeland grasses, cereal crops and sug- why this species has to migrate soon after
arcane. Because of its ability to appear suddenly and emergence.
then disappear equally suddenly, the African army- Spodoptera exempta exhibits a phenomenon
worm has sometimes been referred to by farmers as called polyphenism, or phase polymorphism (i.e.,
the mystery worm. The scale of devastation to the occurrence in a population of two or more
crops and pastures by armyworm is comparable phenotypes due to exposure to different environ-
only to that caused by locusts. Thus, the armyworm mental conditions). For example, up until the
is greatly feared wherever it occurs. third molt, all larvae of S. exempta remain green
in body color. However, at this stage, depending
on whether there are many larvae or just a few,
Biology and Ecology they will turn black or remain in various shades
of green or brown. If there are large numbers of
Adult moths of S. exempta have a wing span in the larvae present as in a typical outbreak situation,
range of 2037 mm. The forewings are character- larvae tend to be characteristically velvety black
ized by an overall dull gray-brown appearance. The on top with pale lines on each side and green-
hind wings are whitish with dark veins. The two ish-yellow underside; this phenotype is called
sexes can be distinguished by examining the num- the gregarious phase. It is during this phase that
ber of bristles on the frenulum (the mechanism S. exempta is most devastating to crops. Larvae
that couples the fore and hind wings during flight), in the gregarious phase tend to be very active
which are single in the males and multiple in the and often march on the soil in one direction only
females. A characteristic feature of the African looking for fresh food. They also feed high on the
armyworm is the presence of racquet-shaped scales plant during the day.
at the tip of the abdomen of the males and black However, if not crowded, the developing lar-
scales on the tip of the body of the females. vae remain one of the many shades of green, pink
Females of S. exempta lay between 100 and or brown color until they pupate. In contrast to
400 eggs per night in a mass covered by black black gregarious larvae, they are sluggish, living
scales from the tip of their abdomen. Eggs are mostly at the bases of plants and are not as
small, 0.5 mm in diameter, whitish in color, but destructive to crops. Although their appearance is
then turn black prior to hatching. Eggs are often so different, they are the same insect and one may
laid on the lower side of leaves and hatch in about easily be converted into the other. Nevertheless,
24 days after oviposition. There are six larval because moths derived from gregarious and soli-
instars extending over a larval period of between tary larvae exhibit the same level of readiness to
14 and 22 days depending on the temperature and fly, phase change in S. exempta is construed as
the host plant on which the larvae have been being merely a stress phenomenon associated
reared. Fully grown sixth instar larvae are often with crowding. It is unknown whether this aspect
2535 mm long. Pupation occurs 23 cm below of phase polymorphism is of any evolutionary
the soil surface. This process is often preceded by a significance to S.exempta. In the case of locusts,
sudden and synchronized disappearance of larvae it is thought that the solitary form is the one
that quickly burrow into the ground, particularly that enables the populations to persist at a low
if soil conditions are moist enough. level during the dry season when there are no
A African Armyworm, Spodoptera exempta (Walker) (Lepidoptera: Noctuidae)

outbreaks occurring. It is worth speculating, in temporary loss of contact with the main popula-
the case of S. exempta, that at such low densities, tion in remote and perhaps uninhabited areas.
its populations continue to breed during the dry This continuity implies that the first outbreaks to
season in areas where grasses remain green, such appear in East Africa are due to the migration of
as in the cool highland areas and, more especially, parent moths from the north at the end of the sea-
the coastal areas where it is hot and there are peri- son. Although there is ample evidence of adult
odic showers during the dry season. This form dispersal on the wind over distances up to several
may, therefore, be of some critical survival value hundred km in a few successive nights, there is lit-
for this pest. tle evidence in support of a southward dispersal at
the start of an armyworm outbreak in East Africa.
The concentration hypothesis, on the other
Seasonal Movements and hand, postulates that because of occasional cap-
Armyworm Outbreaks ture of moths in traps during the off-season, as
well as the finding of rare caterpillars after con-
Upon emergence, adult moths of S. exempta are certed searches, S. exempta persists during such
fully capable of movement from their breeding times of year as uncrowded populations (the soli-
sites to new areas. Such flights can be very short, or tary phase, in which caterpillars remain green and
very long, depending on whether they are carried unreported), and that the first outbreaks of the
in a downwind direction or not. Nevertheless, season are due to concentration of moths before
because they are one-way journeys, they cannot, the synchronized mating and egg laying.
therefore, be regarded as migration in its strict Thus, there seem to be two types of armyworm
ethological sense where there is invariably a outbreaks: primary and secondary outbreaks. Dur-
return flight to breeding sites. ing primary outbreaks, sources of outbreaks are the
The question of how armyworm outbreaks low density populations that survive and breed
start has baffled scientists and farmers alike for a during dry seasons in green areas of the coast and
long time. Because moths emerge over a period of the highlands. Secondary outbreaks occur down-
up to 12 days, and can also fly off on migration at wind from the coast near the first highlands. Dur-
different times, they become widely dispersed and ing years of serious armyworm outbreaks, the first
do not form swarms as occurs with locusts. More- outbreaks often start in Tanzania, Zimbabwe and
over, the moths are weak fliers and are often carried Malawi at about the beginning of the wet season in
in a downwind direction. Thus, moths disperse in December, and are followed by a progression of
space and time downwind. For purposes of this outbreaks at about one generation time-intervals
narrative, an armyworm outbreak is simply from Tanzania through Kenya, Uganda, Ethiopia,
described as the sudden appearance of larval infes- Somalia to the Yemen and from Zimbabwe to
tations, often simultaneously on many farms in South Africa. Wind convergence plus localized
one region. weather and moth behavior provide the mecha-
Two hypotheses, the continuity and the con- nism for transporting and concentrating moths
centration hypotheses, have been put forward in emerging from primary outbreaks.
order to explain how outbreaks begin during an Flight mechanisms of S. exempta prior to out-
armyworm season. The continuity hypothesis, breaks have been the subject of extensive studies.
which is based on biogeographical analyses of past After drying and hardening their wings, moths first
outbreaks, proposes that much of the armyworm move up into the trees. Then, when they are ready,
population is always at crowded, outbreak densi- they fly up several hundreds of meters into the air,
ties, and that the seasonal absence of reports rep- where if caught up by prevailing wind, are carried
resents not a real absence of outbreaks, but a away downwind. When dawn arrives, the moths
African Armyworm, Spodoptera exempta (Walker) (Lepidoptera: Noctuidae)
A 57

descend and hide on the ground in the grass, but at the dry season. But since they have no diapause,
dusk they take off again. They will continue to do they must be living and breeding somewhere. It
this for several days, either until they die, or until has now been established that they survive in quite
they come to an area where rain is falling. Rain low numbers along the coastal area, where some
causes the moth to descend to the ground. The rain falls in every month of the year. When the
winds coming out of the rainstorm have the effect rains begin again in central Tanzania, small num-
of concentrating the moths, rather as though they bers of moths migrate inland from the coast and it
were being swept together with a brush. This is the is these that cause the first outbreaks.
reason why armyworms occurring during out-
breaks are not evenly distributed. Upon descent to
the ground, moths tend to drink water if this is Economic Importance of
available, mate and then lay their eggs. At this stage Outbreaks of S. exempta
dispersal will have come to an end.
In Africa, seasonal rains are brought by the During armyworm outbreaks, feeding damage by
meeting of large scale winds from the northeast S. exempta to cultivated and wild host plants is
and the southwest at the Inter-Tropical Conver- almost entirely restricted to the leaves, although
gence Zone (ITCZ). The position of the ITCZ when food is scarce, the young stems or flowers,
moves with the sun across the tropical zone twice particularly of wild grasses, may also be eaten up.
each year, from north to south between July and The young larvae at first eat the upper and lower
December and from south to north between Janu- surface tissue of the leaves, which results in the
ary and June. The first outbreaks, designated as skeletonization, or windowing, of the leaves. As a
primary outbreaks in East Africa, usually occur in rule, armyworm larvae tend to prefer young plants
central Tanzania in November or December. and recently germinated crops, often defoliating
Occasionally they occur further south, in Mbeya, them to ground level. It is estimated that two lar-
Mtwara and Lindi regions of Tanzania. More vae can completely destroy a 10-day-old maize
rarely, they occur in southeastern Kenya. plant with 67 leaves and a single larva can con-
Moths produced by these primary outbreaks sume 200 mg of dry mass of maize leaves in the
are then carried by the wind towards the ITCZ, course of the sixth instar.
which has meanwhile moved north. The moths are Destruction of cereal crops such as maize,
thus concentrated in areas where the rains are just rice, wheat or sorghum often necessitates replant-
beginning, and so they breed and multiply yet again. ing of the entire affected crop. S. exempta larvae
If conditions are suitable, they will increase at an are most damaging to cultivated and wild host
enormous rate. Although the first outbreaks are only plants during outbreaks when gregarious bands of
a few hectares in extent, by February and March larvae travel together on the ground. Such army-
there may be outbreaks of hundreds of square km. worm outbreaks are often capable of causing total
Thus, as the ITCZ takes the rains northwards, so crop loss within hours at the local level. Thus, crop
the moths, carried by the prevailing wind, move with losses as a result of armyworm outbreaks can be
it, bringing new armyworm outbreaks, designated potentially devastating to local and national econ-
secondary outbreaks, to northern Tanzania, Kenya, omies as they occur in some of Africas most
Somalia, Ethiopia, Sudan and eventually Yemen. impoverished economies. Losses are sometimes
Armyworm outbreak seasons vary greatly in sever- difficult to assess quantitatively because of hidden
ity, extent and timing in each of these countries. costs such as effects of forage destruction, damage
From September to November, there are sel- to subsistence farms, the expense of additional
dom armyworm outbreaks in any part of eastern seed and, most importantly, food aid from interna-
Africa. The armyworm seems to disappear during tional donors.
A African Armyworm, Spodoptera exempta (Walker) (Lepidoptera: Noctuidae)

Survey and Control associated with armyworm monitoring, forecast-

ing and control in countries most ravaged by this
Circumstantial evidence from light and phero- pest is the inadequacy of funding for these
mone traps shows that long distance migrations operations.
occur between moth emergence and the next Because elimination of all primary outbreaks
breeding areas which are in the vicinity of seasonal is almost impossible to achieve, management
passages of low-level wind convergence such as strategies for S. exempta tend to focus more on
Inter-tropical Convergence Zone or African Rift suppression of secondary outbreaks. This approach
Convergence Zones. Furthermore, increasing lev- has invariably been dependent on the application
els of moth catches in light and pheromone traps of pesticide sprays. To date, a series of low toxicity
have been found to be followed by increased prob- pesticides are often used (e.g., synthetic pyre-
ability of infestations of larvae occurring 24 throids, carbamates and organophosphates). Pre-
weeks later at up to 200 km from the trap. viously, highly toxic, persistent and broad spectrum
Because primary outbreaks of S. exempta orig- pesticides such as DDT, BHC, dieldrin and a series
inate from moths taken downwind from sources of others were used due to lack of alternative, safer
nearest the coasts of eastern Africa, and concen- products. However, due to the need for rapid
trate where heavy rains are falling, control of this intervention and coverage of fairly extensive areas
pest involves accurately detecting primary out- during outbreaks, newer oil-based pesticides are
breaks and eliminating them before the subsequent being applied as ultra-low volume (ulv) formula-
moths emerge and move downwind causing sec- tions using hand held, battery driven applicators.
ondary outbreaks in another region or country. Nevertheless, suppression tactics are only cost-
The major objective is to kill as many armyworms effective when applied in a timely manner on
as possible on pasture or crops before subsequent farmland. Outbreaks occurring on uncultivated
moths emerge and move downwind. Normally, the land often remain unchecked.
largest and densest outbreaks are attacked first. Unfortunately, control of armyworms by less
This approach is referred to as strategic control. toxic means has received far less attention than in
However, because primary outbreaks often remain the case of other international migratory pests
unnoticed until after secondary outbreaks have such as locusts, which have been the targets for
occurred, direct elimination of the latter becomes tests involving fungi, protozoa and even viruses.
the main objective in multiple outbreak situations. Extensive studies in eastern Africa have con-
Monitoring possible sources of the moths that firmed that all life stages of the armyworm are
cause primary outbreaks has involved deployment subject to attack by a diversity of natural enemies.
of a network of light and pheromone traps, as well For example, up to 90% of armyworm caterpillars
as ground searches for low-density larvae in the in the last instar can be killed by a nuclear polyhe-
off-season in areas historically known to be sites drosis virus (NPV). Similarly, pupal and prepupal
of primary outbreaks. In addition, for all primary stages can be killed by a cytoplasmic virus. The
outbreak areas, rainfall stations must report on a only fungus known to attack armyworms during
weekly or daily basis the amount of rainfall during conditions of high humidity and temperature is
the first month of the rainy season. Recently, there Normuraea rileyi. Armyworms infected by this
have been attempts to introduce predictive models fungi typically climb to the top of grass blades
that integrate African Real Time Environmental where they die amidst masses of mycelia.
Monitoring and Information System (ARTEMIS) Armyworm larvae are also subject to parasit-
and satellite imagery and synoptic weather data ism by some 28 species of tachnid flies (Diptera:
that has been applicable for detection and elimina- Tachinidae). In cases of parasitism from wasps
tion of gregarious locusts. The major challenge (Hymenoptera), some 25 parasitoids have been
African Honey Bee, Africanized Honey Bee, or Killer Bee, Apis mellifera scutellata Lepeletier (Hymenoptera: Apidae)
A 59

isolated from eggs, larvae and pupae of S. exempta. 2

Florida Department of Agriculture and Con-
Apart from attack by parasitoids, there are several sumer Services, DPI, Gainesville, FL, USA
arthropod predators that prey on armyworms,
including ants (Hymenoptera: Formicidae) and The African honey bee (Apis mellifera scutellata
beetles (Coleoptera), which often prey on eggs and Lepeletier) is a subspecies (or race) of western
early larval stages of S. exempta. To date, there has honey bee (A. mellifera L.) that occurs naturally in
been no effort to study the potential of some of sub-Saharan Africa but has been introduced into
these natural enemies for commercialization. the Americas. More than 10 subspecies of western
Armyworm outbreaks also attract flocks of avian honey bees exist in Africa and all justifiably are
predators, notably storks such as Marabou storks, called African honey bees. However, the term
white (European) storks and Abdims storks. Occa- African (Africanized) honey bee refers exclu-
sionally, such assemblages of predators help to sively to A.m. scutellata in the bees introduced
eliminate small primary or secondary outbreaks range (Fig. 22).
of S. exempta. Subspecies of western honey bees are native
For the more foreseeable future, it is evident to Europe and Africa but have been spread widely
that preventive control of armyworms will con- outside their native range due to their economic
tinue to rely on diligent surveillance during reces- importance as pollinators and producers of honey.
sions and, as outbreaks occur, intensified scouting Initially, only European subspecies of honey bees
will be necessary to locate and then eliminate (hereafter referred to as European bees) were
pockets of solitary morphs before outbreaks. introduced into the Americas, where they were
found to be productive in temperate North America
but less so in Central and South America where
tropical/subtropical climates dominate. In response
References to the poor performance of European bees in
Brazil, Warwick Kerr, a Brazilian scientist, traveled
Regional Armyworm Programme of Desert Locust Control to southern Africa to screen African honey bee
Organization for Eastern Africa (1992) The African
armyworm DLCO-EA, Addis Ababa, Ethiopia, 19 pp
subspecies for productivity and viability. His visit
Meinzingen WF (ed) (1993) African armyworm. In: A guide resulted in the importation of A.m. scutellata into
to migrant pest management in Africa. FAO AGP, Rome, Brazil in the late 1950s.
Italy, pp 7185 Dr. Kerr hoped that through experimentation
Rose DJW (1979) The significance of low-density popula-
tions of the African armyworm, Spodoptera exempta. and selective breeding, the African bee could be
(Walker). Philos Trans R Entomol Soc London B made manageable and available for use by Brazilian
287:393402 beekeepers. As such, he initiated efforts to breed
Odiyo PO (1979) Forecasting infestation of a migrant pest:
gentleness into the African stock while amplifying
the African armyworm, Spodoptera exempta. (Walker).
Philos Trans R Entomol Soc London B 287:403413 its many positive traits. The breeding effort was
not carried to completion because the African
bees swarmed accidentally, ending their initial
quarantine. Following this, the bees began to
African Honey Bee, Africanized spread throughout Brazil and into other parts of
Honey Bee, or Killer Bee, Apis South America.
mellifera scutellata Lepeletier All subspecies of Apis mellifera can interbreed
(Hymenoptera: Apidae) or hybridize. Consequently, African bee hybrid-
ization with European bees became frequent as
jamie ellis1, amanda ellis2 African bees moved into areas previously occu-
University of Florida, Gainesville, FL, USA pied by European bees. It is this hybridization with
A African Honey Bee, Africanized Honey Bee, or Killer Bee, Apis mellifera scutellata Lepeletier (Hymenoptera: Apidae)

African Honey Bee, Africanized Honey Bee, or Killer Bee, Apis mellifera scutellata Lepeletier
(Hymenoptera: Apidae), Figure 22 The natural distribution of Apis mellifera scutellata in Africa (modified
from Hepburn HR, Radloff SE (1998) Honeybees of Africa. Springer-Verlag, Berlin, 370 pp), and its
distribution in the Americas.

European honey bees that earned them the name ovement through South and Central America
Africanized honey bees. Traditionally, African was rapid and largely unassisted by humans, African
and Africanized have been used interchangeably bees earned the reputation of being the most suc-
although the former really refers to the pure race cessful biologically invasive species of all time. In
and the latter to the hybrid. 1990, populations of African honey bees had satu-
The spread of African bees throughout South rated South and Central America and begun to
and Central America, fueled by rapid hybridization move into the USA. As of 2006, African honey bees
with European subspecies and the dominance of were established in the southernmost USA: Texas,
African alleles over European ones, occurred at a California, New Mexico, Arizona, Oklahoma, Lou-
rate of 200300 miles per year. Because their isiana, Arkansas, Alabama, and Florida.
African Honey Bee, Africanized Honey Bee, or Killer Bee, Apis mellifera scutellata Lepeletier (Hymenoptera: Apidae)
A 61

The spread of African bees in the U.S. contin- of honey bee colonies by humans) is more com-
ues, albeit at a much slower rate than what occurred mon in Europe, where the native honey bees have
throughout South and Central America. This been bred for gentleness and ease of management.
slowed rate of territory expansion appears due to In contrast, honey hunting (near-complete
climatic limitations. African bees do not survive in destruction of hive to harvest contents) is more
temperate climates as well as European bees do. common in Africa, resulting in a bee that is more
Therefore, they have failed to establish populations defensive of its nest. Other selection pressures
below about 32 latitude in the southern hemi- that led to a heightened defensiveness in African
sphere. Although they have expanded beyond bees include climatic stresses, resource availabil-
this parallel in the northern hemisphere, African ity, and predation by birds, mammals, and various
bee expansion northward also appears limited reptiles. These selection pressures resulted in an
climatically, being found only below about 34 African race of bee that can be 10+times more
latitude currently. defensive than any of the various European races
of bee.
All honey bees readily defend their nests, and
Description and Behavior an attack usually means that the victim is too close
to the nest. While European races of bees may
African honey bees cannot be distinguished from attack a nest intruder with >10 bees, African bees
European honey bees easily, although they are may attack the same intruder with >1,000 bees.
slightly smaller than the various European races. Further, African bees defend a larger radius around
Laboratory personnel use morphometric analy- their nest and require lower levels of stimuli to ini-
ses to determine the likelihood that a given col- tiate an attack. Because of these characteristics,
ony is Africanized or fully African. With honey African bees are capable of killing large mammals,
bees, the measurement of wing venation patterns including man. This defensiveness has earned
and the size and coloration of various body parts them the nickname killer bee. It is important to
(morphometry) are important determinants of note that their ability to kill humans has nothing
identification at the subspecific level. Morphom- to do with their size or the potency of their venom.
etry has been used to differentiate honey bee African bees are smaller than European bees and
races since the 1960s and remains the first round probably deliver a comparatively smaller dose of
of identification when suspect colonies are dis- venom to their victim than do European bees.
covered. Morphometric analyses were first used Because both bees use the same type of venom,
to differentiate Africanized and European honey human deaths are a result of the number of stings
bees in South America in 1978. A more rigorous they receive rather than an increased potency of
identification is achieved by genetic analysis and African bee venom.
often is necessary when the suspect bees are a Another behavioral difference between Afri-
hybrid between African bees and the European can and European bees concerns colony level
subspecies. reproduction and nest abandonment. African
Other differences between African and Euro- honey bees swarm and abscond in greater frequen-
pean bees manifest themselves behaviorally. To cies than their European counterparts. Swarming,
the casual bystander, the primary identifying bee reproduction at the colony level, occurs when a
behavioral characteristic of Africanized bees is single colony splits into two colonies, thus ensuring
their heightened defensiveness compared to that survival of the species. European colonies com-
of European subspecies. Selection pressures monly swarm 13 times per year. African colonies
induced by man are, in part, responsible for this may swarm >10 times per year. African swarms
increased defensiveness.Beekeeping (management tend to be smaller than European ones, but the
A African Honey Bee, Africanized Honey Bee, or Killer Bee, Apis mellifera scutellata Lepeletier (Hymenoptera: Apidae)

African Honey Bee, Africanized Honey Bee, or Killer Bee, Apis mellifera scutellata Lepeletier
(Hymenoptera: Apidae), Figure 23 (a) African bees swarm readily and nest in unusual locations,
including (b) exposed on tree limbs, (c) within cavities in the soil, (d) within discarded furniture.

swarming bees are docile in both races. Regardless, volume than European honey bees and have been
African colonies reproduce in greater numbers found in water meter boxes, cement blocks, old
than European colonies, quickly saturating an area tires, house eaves, barbecue grills, cavities in the
with African bees. Further, African bees abscond ground, and hanging exposed from tree limbs, just
frequently (completely abandon the nest) during to name a few places. One rarely finds European
times of dearth or repeated nest disturbance, while colonies in any of these locations because they
this behavior is atypical in European bees. prefer to nest in larger cavities like those provided
Another common difference between African by tree hollows, chimneys, etc. As one can imagine,
and European honey bees is their choice of nest humans inadvertently provide multiple nesting
locations. African honey bees are less selective sites for African bees. Therein lies the primary rea-
when considering a potential nesting site than are son African bees are encountered frequently by
European bees. They will nest in a much smaller humans (Fig. 23).
African Honey Bee, Africanized Honey Bee, or Killer Bee, Apis mellifera scutellata Lepeletier (Hymenoptera: Apidae)
A 63

A final behavioral curiosity of African bees bees, but key differences confer adaptive benefits
concerns nest usurpation (or colony takeover) to the former.
of European colonies. Small African swarms Virgin queens of all western honey bees
containing a queen often land on the outside emerge from peanut hull-shaped waxen cells. After
infrastructure of a European colony (a wall, a short time of further maturation, a virgin queen
beekeeper-managed hive, etc.). As time passes, will leave the colony to mate with drones. All mat-
the worker bees in the African swarm begin to ing occurs in the air, with the fastest drones being
exchange food/pheromones with the European the most successful suitors. Queens will mate mul-
workers from the colony. This gradually ensures tiple times over the course of 710 days and during
the adoption of the African bees into the European this time they will mate with an average of 1220
colony. Somewhere during this process, the drones. Queen bees store semen in an organ called
European queen is lost (perhaps killed by the a spermatheca. African colonies produce more
African bees her fate remains uncertain at this drones/colony so drone populations in an area
point) and the African queen is introduced into tend to favor African bees. As such, virgin European
the colony, thus becoming the reigning matri- queens are more likely to mate with African drones
arch. European bees do not display this behavior rather than European ones. Further, flight time and
but often fall victim to it, thus creating an African distances of mating flight from the colony tend to
colony from a preexisting European one. result in European queens encountering African
Other behavioral differences between African drones more often than European drones, thus
and European races exist and are worth discuss- setting the stage for hybridization.
ing briefly. For example, African bees are more All honey bees undergo complete metamor-
flighty than European bees, meaning that when phosis but the time from egg to adult varies by
a colony is disturbed, more of the bees leave the subspecies. The newly-mated queen bee oviposits
nest rather than remain in the hive. African bees in wax cells constructed by worker bees. Fertilized
use more propolis (a derivative of saps and resins eggs result in female offspring, either workers or
collected from various trees/plants) than do queens. If fed a diet rich in royal jelly, the female
European bees. Propolis is used to weather-proof larva will develop into a queen, with the reciprocal
the nest and has various antibiotic properties. true for the development of workers. Drones result
African colonies produce proportionally more from unfertilized eggs and consequently only
drones (male bees) than European bees. Their inherit genetic material from their mother (they
colonies grow faster and tend to be smaller than have no father).
European colonies. Finally, they tend to store Developmental time varies by caste member
proportionately less food (honey) than European (see Table 4) and favor African honey bees because
bees, likely a remnant of being native to an envi- they generally develop faster than European bees.
ronment where food resources are available When bee colonies decide to make a new queen,
throughout the year. newly emerged female larvae are fed royal jelly
constantly. Because Africanized offspring, includ-
ing queens, develop faster than European off-
Life Cycle and Genetic Dominance spring, a queen having an African genotype is
more likely to emerge earlier than a queen with a
Mating biology and developmental time play an European genotype. The first queen to emerge
important role in the success of African bee colo- kills all of her queen sisters that have not yet
nies in replacing European colonies in an area. emerged from their cells. The Africanized virgin
For the most part, mating and developmental proceeds to mate in an area having higher den-
biology are similar for African and European sities of African drones. Over time, this results
A African Honey Bee, Africanized Honey Bee, or Killer Bee, Apis mellifera scutellata Lepeletier (Hymenoptera: Apidae)

in the colony becoming more African with the Africanized honey bees have been established.
European phenotype being replaced almost alto- These precautions are not suggested to make peo-
gether. This process is exacerbated further due to ple fearful of honey bees but only to encourage
the dominance of African genetic traits over Euro- caution and respect of honey bees. The precau-
pean ones. tions include remaining alert for honey bees flying
Finally, African bees are more resistant to into or out of an area (suggesting they are nesting
many honey bee pests/pathogens than are Euro- nearby), staying away from a swarm or nest, and
pean bees. Western honey bees face a myriad of having wild colonies removed from places that
pests and diseases, the most severe of which humans frequent. The latter is perhaps the most
include varroa mites (Varroa destructor), tracheal important advice one can heed when dealing with
mites (Acarapis woodi), small hive beetles (Aethina African bees. In the USA, a large percentage of
tumida), and American foulbrood (Paenabacilis African bee attacks occur on people who know a
larvae). These bee pests almost eliminated all wild nest is present but elect not to have it removed (or
colonies of European honey bees in North America. try to do it themselves).
Because African bees are resistant to many of these If an attack occurs, remembering a few simple
pests/diseases, their survivorship in the wild is recommendations will increase ones chances of
favored over that of European bees. minimizing the effects and severity of the attack. If
attacked, a victim should run away from the area
using his shirt to cover his head and especially
Public Risks airways. Running through tall grass or small trees
will help to disrupt the attacking bees. The victim
Due to their heightened defensive behavior, should not stand and swat at the bees. The bees are
African honey bees can be a risk to humans. Chil- defending their nest, and the victim needs to get
dren, the elderly, and handicapped individuals are away from that nest as quickly as possible. It is
at the highest risk of a deadly attack due to their important that the victim get cover in a bee-proof
inability or hampered ability to escape an attack. vehicle or structure if either is available. One
African honey bees are agitated by vibrations like should not jump into the water or hide in bushes.
those caused by power equipment, tractors, lawn The bees can remain defensive and in the area for
mowers, etc. Further, their nesting habits often put some period of time, thus increasing the risk to
them in close proximity to humans. Because of the victim. If stung, the victim should remove the
this, precautions should be taken in an area where stinger quickly by scraping it rather than by pull-
ing it. One should see a doctor immediately if
breathing is affected.
African Honey Bee, Africanized Honey Many African bee attacks can be prevented by
Bee, or Killer Bee, Apis mellifera scutellata limiting the number of nesting sites that are avail-
Lepeletier ( Hymenoptera: Apidae), Table 4 The able to the bees. A homeowner, school worker, etc.
developmental time in days (from egg to adult) of can bee proof his or her property by eliminating
European and African honey bees possible nesting sites. This can be accomplished by
removing any unnecessary debris from an area
European honey African honey
bees bees
and closing off wall, chimney, electrical and
plumbing-related gaps that are >30 mm using a
Queen 16 14
small-mesh hardware cloth or caulking. This will
Worker 21 1920 limit bee access to potential nesting sites. Finally,
Drone 24 24 one should check walls and eaves of structures
regularly, looking for bee activity.
African Honey Bee, Africanized Honey Bee, or Killer Bee, Apis mellifera scutellata Lepeletier (Hymenoptera: Apidae)
A 65

Managing African Bee Colonies suit, boots, gloves, and a bee veil. Bee veils (protec-
tive headgear) are worn by almost all beekeepers
It is important to remember that African honey worldwide. Traditionally, the veil mesh protecting
bees pollinate crops and produce honey just like the face is colored black to keep down the suns
other races of honey bees. Beekeepers in South glare. African bees (and most honey bees) attack
Africa use African honey bees as the bee of choice dark colors so black-faced veils often get covered
in their operations. So, African bees can be man- with bees. Consequently, beekeepers can use
aged efficiently and safely but the skills required to white-faced veils to keep the bees off of their veils.
manage African bee colonies differ from those Beekeepers managing African colonies often tape
required to manage European bee colonies. their bee suits to their boots and gloves to limit the
In general, the management of African bee possibility of bee access.
colonies has been discouraged in the US while Finally, some beekeepers in areas with African
accepted in Central and South America. This may bees try to requeen African bee colonies with
have to do with the public perception of honey European queens. This is not a common practice
bees, particularly African bees, in the USA and the in sub-Saharan Africa. Most African beekeepers
robust legal system in place in the USA. On the in areas having African bees gladly use the bee in
other hand, beekeepers in Central and South their operations, paying little attention to the bees
America routinely use African bees in their opera- defensiveness.
tions with slight management modifications. In
fact, some South American countries are among
the leading honey producers in the world, due
largely to the presence of African bees in the Conclusion
Beekeepers in South and Central America The economic impact of African bees in an area
utilize a number of management practices in order can be substantial. Keepers of European bees
to keep African bees. First, they keep single bee often notice a decrease in resource availability
colonies on individual hive stands rather than for their bees because of the density of African
using one hive stand for multiple colonies. This bee colonies in an area, and thus the demand on
limits the management activity to one colony at a the available resources is high. Furthermore, cit-
time rather than aggravating other colonies while ies, municipalities, etc., often initiate eradication
working only one. programs, with much futility. Finally, the loss of
Secondly, beekeepers in South and Central animal and human lives is a tragic occurrence,
America use ample amounts of smoke when work- being beyond measurable cost.
ing African bee colonies. It is believed that smoke African bees also may affect the environment
masks the alarm pheromone of the bees, thus less- negatively. Colony densities as high as 300 African
ening the defensive response of the colony. Most bee colonies per square mile have been suggested.
South and Central American beekeepers agree If true, African bees may have a substantial impact
that copious amounts of smoke should be used on the native flora and fauna in an area. While this
when working African bee colonies. It is impor- impact often is not reported and largely is not
tant to smoke the colonies well before any work is understood, it could be significant considering the
done, for once bees from a colony are agitated, potential number of colonies and their need for
smoke may fail to calm them down. resources. Thus, the worlds most infamous honey
Beekeepers managing African bees wear bee is among natures most enigmatic creatures.
appropriate protective gear. A typical beekeeper Apiculture (Beekeeping)
working an African colony would wear a full bee Honey Bee
A African Horse Sickness Viruses

References midge involved in the transmission cycle. Most

species of equines can develop viremias sufficiently
Caron DM (2001) Africanized honey bees in the Americas. high to infect midges. Some tick species have been
The A.I. Root Co., Medina, OH, 228 pp shown to be able to become infected and transmit
Hepburn HR, Radloff SE (1998) Honeybees of Africa.
Springer-Verlag, Berlin, 370 pp
the virus in the laboratory, but the importance of
Winston ML (1992) Killer bees: the Africanized honey bee in this transmission route in nature is unknown.
the Americas. Harvard University Press, Cambridge, In endemic areas, African horse sickness
MA, 176 pp viruses circulate primarily between midges and
zebra, and frequently multiple serotypes are pres-
ent. Transmission rates can be very high. For
African Horse Sickness Viruses example, in the Kruger National Park, South
Africa, zebra foals typically are exposed to all nine
cynthia c. lord serotypes by the time they are 1 year old.
University of Florida, Florida Medical Entomol- The first outbreak outside the sub-Saharan
ogy Laboratory, Vero Beach, FL, USA zone began in 1959 in Saudi Arabia and Iran,
spreading to involve Afghanistan, Pakistan, Syria,
African horse sickness is a highly fatal, nonconta- Lebanon, Jordan, Iraq, Turkey, Cyprus and parts
gious disease of equines, particularly horses. The of India before being controlled by vaccination
African horse sickness virus group consists of nine campaigns and the loss of most susceptible horses
serotypes, in the genus Orbivirus, family Reoviridae. by the end of 1961. Another outbreak occurred in
It is closely related to the bluetongue viruses, which North Africa in 1965, crossing into Spain in 1966.
cause disease in cattle and sheep. Infection with any An outbreak of African horse sickness serotype 4
of the serotypes of African horse sickness virus usu- virus began in central Spain in 1987 and ulti-
ally results in severe disease and high mortality in mately encompassed a large part of Spain, along
horses. Donkeys and mules generally exhibit less with parts of Portugal and Morocco. This out-
severe disease and lower mortality, while wild equids break was the first recorded instance of an African
such as zebra generally show no signs of disease or horse sickness virus overwintering outside of
mortality after infection. The serotypes are differen- Africa. African horse sickness cases occurred for
tiated based on the host immune response, and there four subsequent years in Spain, and it was not
is some cross reaction between serotypes. eradicated until 1990. The most likely route of
All nine serotypes are endemic to sub-Saharan introduction was via zebra imported from
Africa, and have caused serious epidemics when Namibia. Control and eradication of the virus
introduced outside this area. African horse sick- was achieved only by extensive vaccination cam-
ness has had a significant impact on the history of paigns and slaughter of infected or exposed
some parts of Africa, as horses could not be used in equines. It is estimated that 2,000 horses died and
exploration and farming. Outbreaks of African over 350,000 were vaccinated during this out-
horse sickness have a significant economic impact, break. In 1989, an outbreak of serotype 9 occurred
resulting from the direct loss of animals, the costs in Saudi Arabia.
of control programs, and trade regulations and Spain was divided into African horse sick-
quarantines restricting movement of equines from ness-free and infected regions, in order to allow
infected areas. movement of horses for the 1992 Olympics held in
The virus is transmitted by biting midges in Barcelona. No vaccination was allowed in the Afri-
the genus Culicoides. Culicoides imicola has been can horse sickness-free region, so that any trans-
implicated in most outbreaks, while in endemic mission activity would be observed. Equine
areas there may be several species of Culicoides movement out of the infected region was prohibited.
African Horse Sickness Viruses
A 67

A similar strategy is used in South Africa, with an horses, against all serotypes, is practiced in
African horse sickness-free zone in the Western endemic areas. Elsewhere, vaccination generally is
Cape Province, based on historically lowincidence not used routinely or is not allowed. In outbreak
of African horse sickness, and that C. imicola is situations, the virus is first typed to determine the
rare. Surrounding this zone is a surveillance zone serotype involved, then vaccination is targeted
and a protection zone. No vaccination is allowed against that serotype only.
in the free and surveillance zones, and strict move- Most countries restrict importation of equids
ment controls are in place for equines moving from endemic countries. An extended quarantine
from other areas of the country. The zoning cre- period usually is imposed, thus restricting the
ates an area where animals can be held prior to movement of horses for competition. Because it is
exportation. In 1999, there was an outbreak of difficult to differentiate between vaccinated and
African horse sickness in the surveillance zone, infected animals, generally there are restrictions
opening debate about the effectiveness of the on importing vaccinated animals.
movement restrictions and the vector species Antibodies to African horse sickness viruses
involved. Currently, the zoning is still in place and have been found in other animals such as elephants,
there have not been further outbreaks in the camels, and bovines, but there is no apparent ill-
African horse sickness-free zone. ness and their impact on the transmission cycle is
Clinical signs of African horse sickness in not known. Dogs can become infected and die by
horses generally begin with fever. There are three eating meat from the carcass of an infected animal.
forms of African horse sickness disease in horses: Antibodies to African horse sickness have been
pulmonary, cardiac, and febrile. The febrile form, found in wild canids and other carnivores, most
often referred to as horse sickness fever, does not likely also via feeding on infected carcasses.
progress beyond fever and generally resolves. The Humans have been infected only rarely, generally
pulmonary form begins with fever and progresses through laboratory accidents with vaccine strains.
to respiratory difficulty, coughing and nasal dis- African horse sickness has never been found
charge. Death is due to pulmonary edema and in the New World. However, there are species of
cardiac failure. The cardiac form also begins with Culicoides, particularly the C. variipennis complex,
fever, and subsequently edemas develop around present throughout the U.S. Some members of this
the head, neck and chest. Death results from car- complex are competent vectors of African horse
diac insufficiency and progressive pulmonary sickness viruses in the laboratory, and will be com-
edema. Mortality rates can be as high as 95% in petent vectors in the field should an African horse
horses, although donkeys and mules are much less sickness virus be introduced. Vector competence
susceptible and mortality rates are much lower. for the C. variipennis complex varies considerably
Movement of donkeys may be important in the for the closely related bluetongue viruses, but we
spread of these viruses outside the endemic area, lack information on similar variation for African
as they are rarely clinically ill and so infected ani- horse sickness viruses. The implications of this for
mals are not noticed. There is variation in virulence an introduction of any of the African horse sickness
between the nine African horse sickness serotypes, viruses is unknown, but requires further study.
along with differences between breeds of horse and
individual immune responses to the virus.
Vaccines have been developed against these
viruses, but due to the immunological differentia-
tion between the serotypes, cross protection is not
Bram RA, George JE, Reichard R, Tabachnick WJ (2002)
complete and full protection requires vaccination Threat of foreign arthropod-borne pathogens to live-
against each serotype. Routine vaccination of stock in the United States. J Med Entomol 39:405416
A Africanized Bees

Holbrook FR, Tabachnick WJ, Schmidtmann ET, McKinnon C, with peaks from May to August, i.e., during the
Bobian RJ, Grogan WL (2000) Sympatry in the Culicoides cool, dry season. Egg clusters are located mostly
variipennis complex (Diptera: Ceratopogonidae): a taxo-
nomic reassessment. J Med Entomol 37:6576 on the lower trunks of trees and consist of 24130
House JA (1993) Recommendations for African horse sick- glossy, globular or dimpled spheres, each about
ness vaccines for use in nonendemic areas. Rev lev 1mm in diameter. They hatch 610 days after ovi-
Md Vt Pays Trop 46:7781
Mellor PS, Boorman J (1995) The transmission and geo-
position, depending on the season (Fig. 24).
graphical spread of African horse sickness and blue- Caterpillars of various instars are found any-
tongue viruses. Ann Trop Med Parasitol 89:115 time between early March to the end of November,
Mellor PS, Boorman J, Baylis M (2000) biting midges: their role with periods of greatest abundance from June to
as arbovirus vectors. Annu Rev Entomol 45:307340
September. All larval instars are hairy and last
instars occur in two color phases. One of them is
Africanized Bees milky-green; the other, more common one, a highly
camouflaged, brown to greyish-white mottled bark
Honeybees in the Western Hemisphere that are pattern including up to three, more or less distinct
derived from hybridization of African and dorsal saddles. Caterpillars feed solitarily, and move
European subspecies of Apis mellifera. The degree with great speed and agility when disturbed, includ-
of hybridization is unresolved. ing a hopping and ballooning response during the
Bees first four instars. Early instars skeletonize Khaya
Honeybee leaflets at night and spend the day motionless on or
African Honey Bee underneath leaflets. Older instars are free feeders
and rest on the lower trunk during the day. All
instars shun the foliage of just-expanding shoots.
African Mahogany-Feeding There are five instars in males and six in females,
Caterpillar, Heteronygmia each of them lasting from 56 days.
dissimilis aurivillius (Lepidoptera: Pupae are found from late February to early
Lymantriidae) December, abundantly so from June to September.
They often are cradled in loose leaf shelters tied
hans g. schabel together by sparse strands of silk or in other hid-
University of Wisconsin, Stevens Point, WI, USA ing places, such as under bark scales.
The moths are present from February to
Several genera and species of mahoganies in various mid-November, most abundantly so from May to
parts of the tropics are highly valuable timber spe- September. They rest during the day and are
cies, among them African mahogany (Khaya spp.). attracted to lights at night. Slender male moths are
Relatively few defoliators are known to target this light to dark or reddish brown with substantial plu-
genus, among them caterpillars of several silkmoths, mose antennae, whereas the white to cream-colored
the nymphalid Charaxes and the lymantriid Hetero- females are bigger and more robust with smaller
nygmia. Only the latter, presumably monophagous antennae. Males have wingspans of about 40 mm
on Khaya, appears to have pest potential as indicated and are able fliers, while females with wingspans of
by small-scale outbreaks observed in Morogoro, about 50 mm, are reluctant to take to the air and
Tanzania, in the 1980s. then only as poor fliers. Both sexes have faint to
A succession of four generations per year more pronounced grey line markings and a small
allow Heteronygmia dissimilis to be active most of black dot on each front wing. Before oviposition, the
the year, except for a period of estivation during greatly distended abdomen of the female is greenish.
the hottest season, i.e., from November to February. Male moths reach adulthood after an average of 41
Eggs are found from early March to late October days (September/October), females after about 45
African Mahogany-Feeding Caterpillar, Heteronygmia dissimilis aurivillius (Lepidoptera: Lymantriidae)
A 69

African Mahogany-Feeding Caterpillar, Heteronygmia Dissimilis aurivillius (Lepidoptera: Lymantriidae),

Figure 24 Egg (top left) (2x), last instar larva (top right) (1x), pupa (bottom left) (1.7x) and (mating) adult
stages (bottom right) (2x) of Heteronygmia dissimilis, respectively. Both sexes are represented in the
bottom figures, the larger females being above in each picture. (Photo: H. Schabel et al. 1988; reprinted
with permission from ICIPE, the International Centre of Insect Physiology and Ecology.)

days of development. On the average, each female Numerous arthropod predators of the cater-
produces about 200 eggs, laid in several batches. pillars and pupae are believed to be generalists with
While no field control of Heteronygmia has little impact. On the other hand, four hymenopter-
been undertaken to date, a laboratory study docu- ous and two dipterous parasites affecting various
mented full protection of Khaya leaves from defo- stages of H. dissimilis seem more specific. Season-
liation by H. dissimilis, following application of 1% ally, egg parasites in particular had significant
crude, aqueous seed extracts of the neem tree impacts on this insect and in conjunction with the
(Azadirachta indica). fungus Paecilomyces farinosus, which severely
A African Maiden Moths (Lepidoptera: Thyretidae)

ecimated pupae during the rainy season, were

d References
responsible for serious setbacks in the annual
buildup of H. dissimilis. As a result, natural controls Janse AJT (1945) On the South African species of Metarctia,
seem to be quite effective with this insect. with the description of a new species. J Entomol Soc
South Africa 8:9198
Kiriakoff SG (1949) Over de phylogenie van de Thyretidae fam.
nov. (Lepidoptera). Natuurwetenschappen Tijdschrift
References 30:310
Kiriakoff SG (1953) Les Thyretidae du Muse Royal du
Congo Belge (Lepidoptera Notodontidae). Annales du
Ballard E (1914) Two pests of mahogany in Nyasaland. Bull
Muse Royal du Congo Belge, Sciences Zoologiques
Entomol Res 5:6162
(8) 26:191
Rwamputa AK, Schabel HG (1989) Effects of crude aqueous
Kiriakoff SG (1957) Notes sur les Thyretidae (Lepidoptera:
neem extracts on defoliation of Khaya nyasica by Het-
Notodontidae). Bulletin et Annales de la Socit Royale
eronygmia dissimilis (Lepidoptera: Lymantriidae) in
Entomologique de Belgique 93:121160
East Africa. In: Alfaro RI, Glover SG (eds), Proceedings,
Kiriakoff SG (1960) Lepidoptera. Fam. Thyretidae. In: Genera
IUFRO Working Group on Insects Affecting Afforestation,
Insectorum. Brussels, 214:166
XVIII International Congress of Entomology. Vancouver,
British Columbia, Canada, pp 245250
Schabel HG, Schabel A, Msanga HP (1988) Bioecological
aspects of the mahogany defoliator Heteronygmia dis-
similis in Morogoro, Tanzania. Insect Sci Appl
African Pine-Feeding
Grasshoppers, Plagiotriptus
pinivorus (Descamps) and
P.Hippiscus (Gerst.) (Orthoptera:
African Maiden Moths Eumastacidae)
(Lepidoptera: Thyretidae)
hans g. schabel
john b. heppner University of Wisconsin, Stevens Point, WI, USA
Florida State Collection of Arthropods, Gainesville,
FL, USA These grasshoppers are excellent examples of
indigenous insects that developed a preference for
African maiden moths, family Thyretidae, include an exotic plantation-grown crop, in this case pines.
212 species, all African. The classification remains Plagiotriptus pinivorus attained some prominence
controversial and various specialists also place the after causing persistently severe defoliation of
group within Arctiidae. The family is in the super- exotic pines, especially Pinus patula, in Malawi in
family Noctuoidea, in the section Cossina, subsec- the 1960s, resulting in significant tree mortality.
tion Bombycina, of the division Ditrysia. Adults A smaller scale defoliation of P. patula was
medium-size (2357 mm wingspan). Haustellum observed at Morogoro in Tanzania in the mid-
usually reduced or vestigial; antennae pectinate; 1980s, which was attributable to another, very
wings very elongated, with reduced hindwings closely related species, Plagiotriptus hippiscus.
(some with greatly reduced hindwings). Macula- Both of these grasshoppers are highly polypha-
tion typically dark with white or hyaline patches, gous, including herbaceous hosts, shrubs and both
or more colorful. Adults perhaps mostly diurnal; angiosperm and gymnosperm trees. The prime
often wasp mimics. Larvae are thought to be leaf requirement for P. pinivorus seems to be access to
feeders, but most species remain unknown bio- evergreen or semi-evergreen vegetation in areas of
logically. Host plant records include Thymelae- moderate to heavy rainfall, i.e., mostly at altitudes
aceae and Ulmaceae, for the few species known between 1,5252,135 m, but occasionally as low
biologically. as 490 m.
African Pine-Feeding Grasshoppers, Plagiotriptus pinivorus (Descamps) and P.Hippiscus (Gerst.) (Orthoptera: Eumastacidae
A 71

Plagiotriptus pinivorus in Malawi exhibits of emergence two weeks in advance. Another

three generations every 2 years, and the complete smaller peak in emergence in August, however,
life cycle takes about 1 year. Nymphs and adults cannot be explained by rainfall. The first instar
have been observed on pines throughout the year, nymph is ephemeral (about 12 h), and will molt
except from December to late January. Copulation immediately when reaching the soil surface, before
occurs anytime, but peaks from October to January feeding on ground vegetation for the next 23
and May to June. During copulation, the small weeks. Advanced instars complete their life cycle
male assumes a characteristic dorso-lateral posi- on trees, each instar lasting about one and a half to
tion by clinging to one of the hind femurs of the over two months. Young instars are wasteful feed-
female. Both males and females are promiscuous. ers. There are generally six instars for males and
About 720 days after the last mating, females seek seven for females. Despite the extra instar, females
bare soil and dig a shallow pit to lay a batch of up develop more rapidly and reach adulthood at
to six eggs. They then resume voracious feeding in about the same time as do males.
the trees, before laying other batches of eggs at Adult males (Fig. 25) are about 1.52 cm
1735 day intervals. long, moderately robust grasshoppers. Their
Eggs incubate from 49248 days, with an abdomen, shield-like pronotum and greatly
average of 115 days. The winter population hatches enlarged hind femora are strongly compressed. A
from April to May, maturing in November, while minute set of non-functional wings, not found
the summer population hatches from December on nymphs, is hidden under the pronotum. The
to January and matures from May to July. Within thread-like antennae are about one third the
the same batch, an average of 34 days and a maxi- length of the head. The abdomen is strongly
mum of 88 days may elapse between first and reflexed over the back in the male. The insect is
lasthatch. largely leaf-green, but sports inconspicuous,
Nymphal peak emergence and rainfall are small areas of blue, pink, red and white on various
strongly correlated in February, allowing prediction parts of the legs, wings, antennae and the pronotal

African Pine-Feeding Grasshoppers, Plagiotriptus pinivorus (Descamps) and P. hippiscus (Gest.)

(Orhthoptera: Eumastacidae), Figure 25 Plagiotriptus hippiscus female (above) and male (below) with
size comparison (shaded) (drawing, Paul Schroud).
A African Primitive Ghost Moths (Lepidoptera: Prototheoridae)

ridge. Eyes are golden yellow. Females are about present; labial palpi long and porrect, 3-segmented;
twice the size of males, more robust and gener- maxillary palpi short and 3-segmented; antennae
ally less compressed. They are uniformly leaf- short. Maculation is brown or gray, with various
green, except for the golden yellow eyes and darker spots. Biologies and larvae remain
valves of the ovipositor. Their wings are also min- unknown.
ute and hidden under the pronotal shield.
Numerous invertebrate and vertebrate preda-
tors, including skinks, birds and blue monkeys, as References
well as parasites were documented, but ultimately
they were deemed insufficient by themselves to Davis DR (1996) A revision of the southern African family
reduce populations of the grasshopper to non- Prototheoridae (Lepidoptera: Hepialoidea). Entomol
Scand 27:393439
damaging levels. As a result, sticky bands and Davis DR (2001) A new species of Prototheora from Malawi,
chemical controls were relied on for monitoring with additional notes on the distribution and morphol-
and control purposes, respectively. In the 1960s in ogy of the genus (Lepidoptera: Prototheoridae). Proc
Malawi, gamma-BHC at 0.5% proved the most Entomol Soc Wash 103:452456
Davis DR (2003) Prototheoridae. In: Lepidopterorum Catalogus,
effective insecticide for ground and aerial applica- (n.s.). Fasc. 11. Assoc Trop Lepid Gainesville, 8 pp
tions at ultra-low volume formulations. Spraying Janse AJT (1942) Prototheoridae. In: Janse AJT (ed) The
of road banks was particularly recommended, as moths of South Africa, Pretoria, 4(1): 6574
insects clustered there for oviposition in the bare
African Red Tick, Rhipicephalus
evertsi, (Acarina: Ixodidae)
This important tick, known as African red tick,
Lee RF (1972) A preliminary account of the biology and ecol- affects ungulates in Africa.
ogy of Plagiotriptus spp. (Orthoptera: Eumastacidae). Ticks
Malawi Forest Research Institute Research Record 48,
Forestry Research Institute of Malawi, Zomba, 100 pp
Schabel HG, Hilje L, Nair KSS (1999) Economic entomology
in tropical forest plantations: an update. J Trop For Sci
Xth Anniversary Issue: 303315. African Skipper Moths
(Lepidoptera: Apoprogonidae)

African Primitive Ghost Moths john b. heppner

(Lepidoptera: Prototheoridae) Florida State Collection of Arthropods, Gainesville,
john b. heppner
Florida State Collection of Arthropods, African skipper moths, family Apoprogonidae,
Gainesville, FL, USA includes only a single species from South Africa.
The family is in the superfamily Uranioidea, in
African primitive ghost moths, family Prototheo- the section Cossina, subsection Bombycina, of
ridae, comprise 12 species of small moths from the division Ditrysia. Adults medium size (4656
South Africa. The family is in the superfamily mm wingspan), with head rough scaled and eyes
Hepialoidea, in the infraorder Exoporia. Adults large; haustellum naked; labial palpi porrect;
small (18mm wingspan), with head rough-scaled; maxillary palpi minute, 1-segmented; antennae
haustellum reduced and vestigial mandibles clubbed (hooked at tip). Wings triangular and
African Swine Fever
A 73

African Slug Caterpillar Moths

(Lepidoptera: Chrysopolomidae)

john b. heppner
Florida State Collection of Arthropods, Gainesville,

African slug caterpillar moths, family Chrysopo-

lomidae, are a small African family of about 30
known species. Two subfamilies are known:
Ectropinae and Chrysopolominae. The family is
in the superfamily Cossoidea (series Limacodi-
African Skipper Moths (Lepidoptera: formes) in the section Cossina, subsection Cos-
Apoprogonidae), Figure 26 Example of African sina, of the division Ditrysia. Adults medium
skipper moths (Apoprogonidae), Apoprogones size (2452 mm wingspan), with head scaling
hesperidis Hampson from South Africa. smooth; haustellum and maxillary palpi absent;
antennae short and bipectinate in males. Body
short; body robust. Maculation dark gray with robust. Wings rounded and broad (some with
some pale markings, plus pale discal spot on irregular distal margins). Maculation mostly
fore- and hindwings. Adults presumed diurnal, pale brown, often with subapical wing line and
but nothing is known of the biology or larvae light discal spot; hindwing with forewing line
(Fig. 26). and coloration continued (Fig. 27). Adults noc-
turnal as far as is known. Larvae leaf-feeding
and slug-like, with small spines; often colorful.
References Host plants include Celastraceae. No economic
species are known.
Janse AJT (1932) Family Sematuridae. In: The moths of South
Africa, 1:8789. Pretoria [Apoprogones]
Seitz A (1926) Subfamilie: Apoprogeninae [sic]. In: Seitz A
(ed) Die Gross-Schmetterlinge der Erde. Teil 14. Die References
afrikanischen Spinner und Schwrmer, pl. 1. A. Kernen,
Stuttgart, pp 1617
Aurivillius C (1911) Chrysopolomidae. In Lepidopterorum
catalogus, 1:14. W. Junk, Berlin.
Hering EM (1928) Familie: Chrysopolomidae. In: Seitz A.
African Sleeping Sickness (ed) Die Gross-Schmetterlinge der Erde. Teil 14. Die
afrikanischen Spinner und Schwrmer, pl. 76. A. Kernen,
Stuttgart, pp. 477479
A disease of humans caused by protozoans in the Herring EM (1937) Revision der Chrysopolomidae (Lep.).
genus Trypanosoma. It is also known as human Ann Transvaal Mus 17:233257, pl. 9
sleeping sickness or human trypanosomiasis. The
same disease, when infecting other vertebrate ani-
mals, is called nagana. It is transmitted by tsetse
flies in Africa. African Swine Fever
Sleeping Sickness or African Trypanosomiasis
Trypanosomes A viral disease of hogs, this tick-transmitted
Tsetse Flies disease is found on several continents.
Nagana Ticks
A Agaonidae (Hymenoptera)

opening. The females of some species actively gather

and carry pollen in thoracic pockets or specialized
leg structures. The cultivated fig, Ficus carica, has
many varieties that no longer rely on pollination to
produce edible ripe syconia. However, a few varieties
still require the pollination service provided by the
agaonid partner, Blastophaga psenes.
Wasps, Ants, Bees and Sawflies
Fig Wasps

African Slug Caterpillar Moths (Lepidoptera: References

Chrysopolomidae), Figure 27 Example of African
slug caterpillar moths (Chrysopolomidae), Bronstein JL (1988) Mutualism, antagonism, and the fig-pol-
Chrysopoloma similis Aurivillius from South Africa. linator interaction. Ecology 69:12981302
Galil J, Eisikowitch D (1968) On the pollination ecology of
Ficus sycomorus in East Africa. Ecology 49:259269
Galil J, Zeroni M, Bar Shalom D (Bogoslavski) (1973) Carbon
Agaonidae (Hymenoptera) dioxide and ethylene effects in the co-ordination between
the pollinator Blastophaga quadriceps and the syconium
in Ficus religiosa. New Phytol 72:11131127
hannah nadel Machado CA, Jousselin E, Kjellberg F, Compton SG, Herre EA
USDA-ARSSan Joaquin Valley Science Center, (2001) Phylogenetic relationships, historical biogeography
Parlier, CA, USA and character evolution of fig-pollinating wasps. Proc R
Soc London B 268:685694
Ramirez BW (1969) Fig wasps: mechanisms of pollen trans-
fer. Science 163:580581
A tropical family of about 750 species of miniscule Weiblen GD (2002) How to be a fig wasp. Annu Rev Entomol
wasps (order Hymenoptera) that are mutualistically 47:299330
associated with fig plants (Ficus spp.). The associa- Wiebes JT (1966). Co-evolution of figs and their insect polli-
nators. Annu Rev Ecol Syst 10:112
tions are usually between a unique pair of fig and
wasp species and are crucial for the reproduction of
both. The winged female wasp enters a young floral
receptacle, the flask-like syconium, and lays single Agathiphagidae
eggs in many of the tiny female flowers lining its
inner surface. Carrying pollen from her natal fig, she A family of moths (order Lepidoptera). They com-
deposits it onto the stigmas during oviposition, monly are known as Kauri moths.
ensuring seed production for the fig and food for Kauri Moths
her own offspring. The syconium will generally not Butterflies and Moths
ripen without pollination. The female is trapped
inside the syconium and dies there. After the larvae
develop and pupate in seed-galls, the wingless adult
males emerge first and chew holes into galls to mate Agassiz, Jean Louis Rodolphe
with the quiescent females inside, and then coopera-
tively chew an opening through the syconial wall. Louis Agassiz was born at Mtier-en-Vuly, Swit-
The changed atmosphere inside the syconium wakes zerland, on May 28, 1807. He displayed an early
the females, which chew out of their galls, actively interest in natural history. In 1824 he entered
or passively pick up pollen, and leave through the Universitt Zrich for medical training, then
A 75

moved to Universitt Heidelberg in Germany. In Nordenskild E (1935) The history of biology: a survey. Tudor,
New York, 629 pp.
Heidelberg his interest in natural history increased.
His next move was to Universitt Mnchen and,
while there in 1829 and still only 21 years old, he
published a work on Brazilian fishes, using the Age Polythism
collected materials of von Martius and von Spix.
His next zoological endeavor was to begin research This refers to the division of labor within a colony
on fossil fishes. In 1831, he moved to Paris still of social insects wherein the responsibilities of the
with the ambition of completing his medical individuals change as they mature.
training. However, he spent part of each day
studying fossil fishes and he came under the influ-
ence of Cuvier and adopted the latters views of Aggregation
creation. Thus, according to Agassiz, each species
was the result of separate creation, not of evolu- A group of individuals consisting of more than just
tion. In 18321846 he was a professor at Univer- family members; a coming together of individuals
sit de Neuchtel, Switzerland. In 1836, Louis to form a group. (contrast with colony)
began to study glaciers and their effects, on which Cycloalexy
he published works in 1840, 1846 and 1847. In Allelochemicals
Switzerland, he also published a large catalog
Nomenclator zoologici of the names of animals,
of which some fascicles were on insects. In 1846
Aggregation Pheromone
he moved to the USA, was welcomed as a famous
scientist, and in 1848 was appointed professor of
A pheromone that causes insects to aggregate. This
zoology and geology at Harvard University. At
type of pheromone is used by insects for mating,
Harvard, he clashed with Asa Gray, professor of
feeding, or oviposition.
botany, about evolution, because Gray supported
Darwins theory. However, it was Louis efforts and
influence that led to the foundation of the Museum
of Comparative Zoology at Harvard University,
an institution that became very influential in Aggregative Response
research on insect systematics. He also was a co-
founder of the U.S. National Science Foundation. The response of predators in which they increase
He died in Cambridge, Massachusetts, on December their time spent in areas with more prey, leading to
14, 1873. His son Alexander and two daughters higher predator density, and fewer prey.
of his first marriage accompanied him to the Learning in Insects
USA (his first wife having died in Switzerland). Predation: The Role of Generalist Predators in
Alexander Agassiz (18351910) likewise became Biodiversity and Biological Control
a zoologist. Louis Agassiz remarried in 1850 in
the USA.

References This term is used to refer to insects that lack man-

dibles, which essentially means that they lack
Anon (1998) Jean Louis Rodolphe Agassiz. Encyclopedia of mouth structures.
world biography, 2nd ed. Gale, Detroit, MI Mouthparts of Hexapods
A Aggressive Mimicry

Aggressive Mimicry Agricultural Crop Pests in

Southeast Asia Including
This is a type of mimicry in which a predator mim- Southern China
ics their prey, allowing ready capture and con-
sumption of the victim. emmett r. easton
Mimicry University of Hawaii at Manoa, Honolulu, HI,
Myrmecomorphy USA
Southeast Asia is often called the Oriental faunal
region and includes the provinces of southern
Agonoxenidae China south of the Yangtse river (Anhui, Fujian,
Guangdong, Guangxi, Guizhou, Hainan, Hubei,
A family of moths (order Lepidoptera). They com- Hunan, Jiangshu, Jiangxi, Sichuan, Yunnan and
monly are known as palm moths. Zhejiang) as well as the island of Taiwan, islands
Palm Moths and peninsular area of Hong Kong, Macao and
Butterflies and Moths those countries to the south including Vietnam,
Malaysia, Singapore, Myanmar (Burma), Cambodia
and Laos, as well as portions of Pakistan and India.
The northern provinces of India share similar fau-
Agricolous nal elements as southern China as both are at a
similar latitude. The insect fauna of northern
This refers to species that dwell in agricultural China is more Palearctic in nature and many of
habitats. the northern species will differ from those in the
southern provinces or elsewhere in Southeast Asia.

Agricultural Chemicals
Insects of Rice
Pesticides, adjuvants, and other chemicals, other
than fertilizers, that are used to enhance crop What are believed to be key pests, or those of
production. major importance as opposed to minor can vary
Insecticides from country to country. In Vietnam and south-
Acaricides or Miticides ern China, the rice stemborer complex of lepidop-
terous insects includes the yellow or small rice
borer, Scirpophaga incertulas (Wlk), the striped
Agricultural Consultant rice-stalk borer, Chilo suppressalis (Wlk), the dark-
headed rice borer, C. polychrysus (Meyr) (all
Someone trained in the agricultural and manage- Pyralidae) and the noctuid pink borer, Sesamia
ment sciences who provides plant and animal inferens (Wlk). Although they are considered
production and protection services for a fee. Inde- minor pests in some regions, because the rice
pendent crop consultants sell the advising service plants are able to tolerate some damage and can
only, deriving no income from sale of products, compensate for light infestations, these insects
whereas other crop consultants usually derive some have been ranked as major pests in the countries
income from product sales such as pesticides or of Malaysia and Thailand. The yellow or small rice
fertilizer. borer, Sc. incertulas, has a wide distribution in
Careers in Entomology Southeast Asia and is found in India, Pakistan,
Agricultural Crop Pests in Southeast Asia Including Southern China
A 77

Sri Lanka, Bangladesh, Myanmar (Burma), Viet- Cicadellidae) and Nilaparvata lugens (Stl) and the
nam, Singapore, Taiwan and Hong Kong. brown plant-hopper (Hemiptera: Delphacidae).
The common names of these insects generally These leafhoppers are important in Thailand and
refer to the color of the larval stage such as the pink Malaysia as well as in India and Pakistan, while the
borer, S. inferens, with pinkish-colored larvae, or planthopper is more widely distributed and is
the yellow larvae of the small rice borer. The head found in southern China, India, Taiwan, Japan and
capsule in larvae of C. suppressalis is brown in color some of the Pacific Islands. The planthopper is
while larvae of Sc. incertulas are yellow with brown unusual in that it is able to migrate between land
heads and the pink borer caterpillar, Sc. inferens, is masses and migrates from East China to Japan
pink in color and larger when mature than the annually. This insect also has migrated to Macao
other species. The larvae tunnel as caterpillars (where rice is no longer grown due to urbaniza-
intothe stems of the rice plants. There they feed on tion) from mainland China.
the plant tissues and destroy the growing points of Nephotettix virescens is an important vector
the plant causing wilting of new shoots, eventually of two viral diseases in Malaysia. The first disease
producing a condition known as dead heart. In is similar to yellow dwarf disease and the second is
mature plants, empty panicles appear white in called Tungro disease. Both cause a stunting of
color, and the condition is known as white-head. plant growth, the first disease a general yellowing
Masses of eggs are generally laid on the leaves in and profusion of tillers, while the second causes a
the case of the female dark-headed borer or striped reddening of the leaves. Both diseases decrease
rice borer. Pupation generally occurs in the stem crop yield.
with these species. The female pyralid moths in the The eggs of the rice leafhoppers are laid in
genus Scirpophaga have a scale tuft at the tip of rows within leaf-sheaths. The five nymphal stages
their abdomens, while female moths in the genus are completed in l7 days in Malaysia. Leafhoppers
Chilo have no scale tuft at the end of the abdomen. generally feed on the upper parts of the plant, while
The female Sc. incertulas has a yellow forewing, the planthopper, Nilaparvata, feeds at the base of
which is whitish in Sc. nivella. the plants near the water line. The brown planthop-
The eggs of stemborers are often attacked by per is sometimes considered the most serious pest
the parasitoid wasps Trichogramma sp. (Tricho- of rice in Asia. They cause a scorching of the
grammatidae) and Telenomus rowani (Scelioni- plants, or a condition known locally as hopper-
dae). In Malaysia, granular insecticides have burn, when the number of nymphs and adults per
been used to control rice stem borers when the clump of rice exceeds 900 or more. The eggs are
incidence in stems (tillers) exceeds l0%, but generally laid in plant tissue. The young resemble
this practice is not recommended in southern the parents except for their smaller size and absence
China or in northern Vietnam where the plants of wings. Five nymphal stages generally require
are able to tolerate some damage and compensate two weeks to develop. Predators can include the
for injury. staphylinid beetle Paederus fuscipes and the coc-
cinellid beetle Harmonia octomaculata.
In Thailand and Malaysia, another important
pest of rice is the rice gall midge, Orseolia oryzae
Rice Leafhopper and Planthopper (W.-M.) (Diptera: Cecidomyiidae), which is also
Complex found in Pakistan, India, Bangladesh, Sri Lanka
and parts of Indonesia as well as in southern
The rice leafhopper and planthopper complex China, where it is considered of minor impor-
includes Nephotettix virescens (Dist) and N. nigropictus tance. The larvae of the fly feed between the leaf
(Stl), the green rice leafhoppers (Hemiptera: sheaths and, when reaching the apical buds, can
A Agricultural Crop Pests in Southeast Asia Including Southern China

lacerate tissue and can cause the formation of a Sugar Cane Insects
gall known locally as a silver or onion shoot. The
adults are delicate looking midges, long and brown The lepidopterous sugar cane borer complex in
in color with long legs. Upon hatching, the pale, Thailand includes several pyralids including the
1mm long larva grows to 3 mm and becomes red- yellow top or early shoot borer, Chilo infuscatellus
dish in color. The pupa, when formed, is pinkish (Snellen), the sugar cane stem borer, Chilo sac-
and turns red with age. Grassy vegetation near the chariphagus (Bojer), the white top borer, Scirpoph-
rice fields is often associated with the presence of aga excerptalis (Walker) and the noctuid sugar
the midge. cane stalk borer, Sesamia inferens (Wlk).
The rice leaf folder, Cnaphalocrocis medinalis The larvae of these insects bore into the
(Gn), (Lepidoptera: Pyralidae), is considered one shoots of sugarcane. As the common name sug-
of the more important pests of rice in southern gests, the larvae of the yellow top or early shoot
China as well as in Malaysia and Thailand. It is also borer tunnel into the growing shoots of the plant,
found in India, Pakistan, Sri Lanka and Bangladesh. while the larvae of C. sacchariphagus bore into
The larvae fold the leaf while feeding and trans- the stems. The noctuid moth larvae of purple
parent patches form so that the rice plant appears stalk borer, Sesamia inferens, previously discussed
ragged. The adult moths lay eggs on young, 4 to as a pest of rice, also affects sugar cane, but sugar
6-week-old plants or in nursery stock in Malaysia. cane is not preferred for oviposition as are rice
Early instar caterpillars feed by scraping the epi- and grasses. The larvae are colored purple to pink
dermis from rice leaves, while later instar caterpil- dorsally and white ventrally and have a reddish-
lars fold them. The opposite edges of the leaf, or orange head capsule. The adult moth is fawn-
one edge of the leaf, is attached to the midrib by colored with dark brown streaks on its forewings
silken threads produced by the larvae. Pupation and whitish hindwings.
occurs inside a silken cocoon within the folded In Malaysia, the white sugar cane aphid, Cera-
leaf. Parasitoid wasps, such as Apanteles opacus tovacuna lanigera Zehntner, a mealybug-like insect
(Braconidae) or Temelucha philippinensis (Ichneu- (Hemiptera), causes injury. The non-winged
monidae), often keep populations in check in females and nymphs are covered by a waxy layer,
Malaysia. while the winged adults are bluish-green in color
The lepidopterous armyworm and cutworm and are not covered with a layer of wax.
complex, including the rice armyworm, Mythimna
loreyi (Duponchel), and the rice ear-cutting cater-
pillar or paddy armyworm, Mythimna separata Fruit Tree Insects:
(Wlk), are important pests in Thailand and southern Mango-Citrus-Banana-Litchi
China. The paddy armyworm affects rice in
Pakistan, Sri Lanka, India and Bangladesh. The One of the most important fruit tree insects is con-
larvae feed on leaves and stems and can defoliate sidered to be the Oriental fruit fly, Batrocera
the plants. dorsalis Hendel (Tephritidae). It is found in Hawaii
The rice skipper, Parnara guttata (Bremer & as well as in other Pacific islands and the southeast
Grey) (Lepidoptera: Hesperiidae), was formerly Asian countries of Thailand and Malaysia where it
considered a major pest in southern China, but is a serious mango pest. In southern China, where
recent mass production and release of parasitic there are fewer mangoes grown, it is considered a
wasps has probably lowered its status to a minor minor pest. In addition to mangoes, the guava and
pest. The larvae roll the apical portion of the leaves, carambola are affected in Malaysia. The larvae or fly
web the sides and cut off the apex, forming long, maggots feeding inside the skin of the fruit cause it
conspicuous tubes. to decay. Female flies puncture the skin of the fruit
Agricultural Crop Pests in Southeast Asia Including Southern China
A 79

with their ovipositors, laying several eggs inside. i ncreasingly important, particularly in the Pacific
The larvae can hatch in one day and develop through island area. The first instar in these Papilionid but-
three instars in about a week. When mature, they terflies are colored differently than older larvae,
are able to leave the fruit by flipping themselves in which may be an adaptation that protects them
the air and dispersing to enter the ground to pupate. from predators such as birds or lizards. The early
Fruits can be protected from these flies by bagging stage larva resembles a bird or lizard dropping as it
them using paper bags. The use of traps treated with is dark brown in color with white markings that
methyl eugenol as an attractant has met with some may appear to be unappetizing to the predator.
success, but as only the males are attracted, it is not When the caterpillar is older, its color changes to
totally effective as a control. green with grey and white markings. Hand picking
In Malaysia, a longhorn beetle known as the the larvae is probably an adequate control in young
mango shoot borer, Rhytidodera stimulans (White), plants.
tunnels into the young growing shoots of the tree. The orange spiny whitefly, Aleurocanthus
Eventually it kills the outer branches, which often spinifera (Quaintance) (Hemiptera: Aleyrodidae),
break off during subsequent wind storms. In is an important insect in southern China as well
southern China, the citrus long-horn beetle, Ano- as India, Sri Lanka, Bangladesh, Malaysia and
plophora chinensis (Forst.) (Cerambycidae), is Thailand. The adults are l mm in length and lay
considered one of the most important pests of cit- eggs on the undersides of leaves. There are three
rus trees. The larvae of these beetles tunnel under nymphal stages and the third-stage nymph appears
the bark of young trees and sometimes into the blackish in color with waxy secretions on the outer
heartwood, which can cause the death of the edge of the body so it looks superficially like an
plants. The adult beetles have striking black and insect pupa. Both the nymphs and the adults
white body coloration. Parasitoid wasps have dif- remove plant nutrients when feeding, and the
ficulty reaching the larvae that bore into young honeydew produced by the nymphs encourages a
healthy trees. sooty moldto grow on the upper surfaces of the
Butterflies (Lepidoptera) are not considered leaves and the fruits. Heavy infestations of this
to be serious pests of agricultural crops in the insect cause fruit production to fall off. A parasitic
northern hemisphere with the exception of the wasp, Eretmocerus serius (Aphelinidae), has been
small white, Pieris rapae. However, in the semi effective in regulating the orange spiny whitefly in
and tropical regions of the world, cold climatic Malaysia.
conditions are not as severe and the insects often Aphids, such as the black citrus aphid, Tox-
do not have to enter diapause (hibernation), so optera aurantii (Bayer de Fonscolombe), and the
there can be continuous generations in some brown citrus aphid, Toxoptera citricida (Kirkaldy),
regions almost throughout the year. Papilionid are important in southern China and also range
butterflies, as well as several species of skippers into India, Sri Lanka and Bangladesh.
(Hesperiidae), cause injury to plants because they Fruit-piercing moths, such as Othreis fullo-
are able to oviposit in both the spring and the fall. nia (Cl.) (Noctuidae), pierce the ripening fruits
In Southeast Asia, the lemon or lime butterfly, of citrus, mango, papaya and guava or banana in
Papilio demoleus (L.), and another species known order to obtain sap. A short, stout proboscis with
by the common names of the common Mormon a barbed tip enables the moth to puncture the
swallowtail or the white-banded swallowtail, skin of the fruit and can permit the entry of plant
Papilio polytes L., lay their eggs on the undersides pathogens such as viruses or secondary rots that
of leaves of citrus plants and are considered impor- can cause premature fruit drop. Fruit-piercing
tant in southern China as pests of citrus. Another moths are considered of major importance
species, Papilio xuthus L., is also becoming today in southern China. Management of their
A Agricultural Crop Pests in Southeast Asia Including Southern China

populations is difficult as the immature of O. ful- Vegetable Insects

lonia do not feed on citrus trees. Instead, the cat-
erpillars feed on the foliage of the Erythrina The diamondback moth, Plutella xylostella (L.)
species of shade trees. On the Pacific island of (Lepidoptera: Plutellidae), is currently considered
Guam, the eggs can be laid on the foliage all year among the top 25 most important arthropod pests
round and the insects are considered to be major in southern China. It was the first agricultural pest
pests as they feed on ripe banana, mango, papaya, in Malaysia to be reported resistant to pesticides,
pomegranate and guava as well as tough-skinned and it is also an important pest in Thailand as well
citrus fruits. as in India. Its distribution has been considered cos-
On banana plants, there are two species of mopolitan. Cruciferous plants, such as the cabbages,
leaf rollers or banana skippers in Southeast Asia, are affected. The caterpillars penetrate the epider-
one of which is Erionota thrax (Hesperiidae), which mis of leaves, mining the tissue and making win-
rolls the banana leaves in Malaysia, Thailand and dows or holes in it. The adult is recognized by the
in southern China. The caterpillars cut and roll pale triangular or diamond-shaped marks seen on
strips of banana leaf, then hide in the roll that is the midline of the back when the wings are closed.
held together by silken threads. They emerge at The caterpillar is pale green in color, and it wrig-
night to feed and are often covered with a white gles violently when disturbed, sometimes falling off
powdery secretion. the edge of the leaf. A microbial insecticide, Bacillus
thuringiensis, is effective in the control of the dia-
mondback moth, but farmers in some areas of Malay-
Litchi and Longan Fruit Insects sia have not accepted it because this method takes
longer to kill the caterpillars than other insecticides.
The litchie stink bug, Tessaratoma papillosa The green stink bug, Nezara viridula (L.)
(Drury) (Hemiptera: Tessaratomidae), has been (Hemiptera: Pentatomidae), is a cosmopolitan
considered the most important pest of litchi insect in Southeast Asia that damages developing
(Litchi chinensis) and longan (Euphoria longan) vegetables, such as potato, sweet potato, tomato
fruit trees in the Guangdong region of southern and cotton, by their feeding punctures. Three color
China, including Hong Kong and Macao as well varieties or subspecies of this insect are recognized
as Vietnam and Thailand. The adults are mostly in southern China. An all-green form, known as
brown dorsally and whitish underneath. Imma- N. viridula smaragdula, is the most common,
ture bugs, more brightly colored than the adults, accounting for 7580% of vegetable bugs observed
have red markings dorsally often with a white in Macao in l996. A second form with yellow on
waxy secretion underneath. Plant sap is taken the head and pronotum, N. viridula torquata,
from the stems of fruit trees. The saliva of the bug makes up about 10% of the stink bug population,
can stain the clothing of fruit tree workers. Also, while the least common form is mostly all yellow
the fluid is extremely irritable if it gets in the eyes. with green spotting on the hemelytra and abdo-
An effective biological control of the litchie stink men, and is called N. viridula aurantiaca (1.0%).
bug has been developed in southern China by the The small white butterfly, Pieris rapae (L.) (Lep-
Guangdong Entomological Institute in which the idoptera: Pieridae), along with two other species, is
egg parasitic wasp, Anastatus japonicus (Eupelmi- still considered important in southern China and
dae), has been mass-reared and released to achieve Southeast Asia. The small cabbage butterfly, Pieris
control in the Fujian, Guangdong and Guangzi canidia (Sparrman), also damages nasturtium. In
provinces. Biological control of the litchie stink both species, larvae feed singly in the cabbage heart,
bug has also been reported from the northern make holes in the leaves and cause frass accumulation.
highlands of Thailand. The insects are also found in India, Taiwan and the
A 81

Philippines, and breeding can be continuous with Agroecology is the application of ecological prin-
up to eight generations annually, which is not the ciples to agricultural production systems and the
case in northerly regions, where overwintering resources needed to sustain them. A convenient
occurs in the pupal stage. unit of study is the agroecosystem, often a single
The Asian corn borer, Ostrinia furnacalis agricultural field. A major difference between an
(Guenee) (Lepidoptera: Pyralidae, Pyraustinae), is agroecosystem and a natural ecosystem is in the
more important in the northern countries of level of human intervention and management
Southeast Asia including southern China where involved. Like natural ecosystems, agroecosystems
corn is grown. The Guangdong Entomological consist of living (biological) and nonliving (chem-
Institute rates it as a highly important pest there. ical, physical) portions. The science of agroecology
The larvae bore into the stalks and the ears of corn, examines the living organisms (collectively called
and can also survive on foxtail millet, Setaria ital- the community) in the system, their interactions
ica, and on Panicum grasses. The eggs are laid in with one another, and the environmental factors
clusters of 1040 underneath leaves about a week that influence them.
before the plant forms its inflorescence. The young
larvae can scarify the leaves and later, bore into the
stem. Pupation generally occurs within the stalk, Nutrient Cycling
but can occur within the ear. The Asian corn bor-
ers range includes India, SriLanka, Korea, China, The organisms within an ecological community
Hong Kong, Taiwan, Vietnam, Japan, Malaysia, depend on one another for energy and materials.
Thailand, Singapore and Indonesia. Green plants are referred to as producers since they
Tropical Fruit Pests and Their Management are at the base of the food chain in ecosystems. Ini-
Sugarcane Pests and Their Management tially, carbon and energy are stored in plant tissues
Vegetable Pests and Their Management through the process of photosynthesis. Consumers
must obtain their carbon and energy by eating
References plants or other organisms. Therefore carbon and
other materials move from green plants to herbi-
Denton GWR, Muniappan R, Austin L, Diambra OH (1999) vores to carnivores, including predators and para-
Fruit-piercing moths of Micronesia. Agricultural Experi- sites. The different levels of energy production and
ment Station, University of Guam. Tech Report #217, 26 pp
Hirose Y (ed) (1992) Biological control in South and East
consumption (producers, herbivores, predators)
Asia. Kyushu University Press, Fukuoka, Japan, 68 pp are called trophic levels, although in reality many
Ooi PAC (1999) Insects in Malaysian agriculture. Tropical organisms do not restrict their feeding to one level.
Press, Kuala Lumpur, Malaysia, 106 pp For instance, some Hymenoptera that are parasi-
Robinson GS, Tuck KR, Shaffer M (1994) Field guide to
smaller moths of Southeast Asia. Malaysian Nature toids as larvae may feed on pollen or nectar as
Society, Kuala Lumpur & Natural History Museum, adults. As a result, the paths along which materials
London, 308 pp move through the organisms in the community
Waterhouse DF (1998) Prospects for the classical biological
can be quite complicated, and collectively they
control of major insect pests and weeds in Southern
China. Entomologia Sinica 5:320341 make up the food web within the community.
Many nutrients, including nitrogen, phospho-
rus, potassium, and other elements, are essential
components of plant and animal tissues. These
Agroecology nutrients have distinct cycles in ecosystems, and
they cycle in food webs along with carbon and other
robert mcsorley materials. Nutrients are released into the soil during
University of Florida, Gainesville, FL, USA decomposition of organic molecules, where they
A Agroecology

are converted into forms that can be taken up by some regions, and many variations exist. Some
plant roots, completing the nutrient cycle. Microar- tropical subsistence intercropping systems are espe-
thropods such as mites and springtails are particu- cially diverse and complicated.
larly important in the decomposition process.

Pest Management
Cropping Systems
The cropping system used has a direct effect on
Crop performance depends on a range of key pest management, which is an important aspect of
resources including nutrients, water, soils, andother agroecology. For example, monoculture may
environmental factors, and many agricultural man- encourage buildup of some pests, such as corn
agement practices are aimed at optimizing and con- rootworms (Diabrotica spp.) or wireworms (Elat-
serving these resources. Various types of cropping eridae), that can be managed by appropriate crop
systems may be selected to address specific goals or rotation. In many cases, the use of intercropping
conservation issues. Conservation tillage and other has resulted in less severe pest outbreaks and
reduced tillage practices are important for soil con- increased diversity of natural enemies compared
servation and reduction of erosion. The crop resi- to monocultured systems.
dues that remain on the soil surface in uncultivated Tritrophic (plant-herbivore-predator) inter-
sites can also aid in conservation of water and actions and the structure of the food web may be
organic matter, and may provide some nutrients affected by changes in the cropping system, such
when they decompose (Fig.28). Monoculture as use of an intercrop, or changes in crop variety or
allows a grower to specialize by growing only one fertility level. The development and use of biologi-
crop, while polyculture permits a grower to diver- cally based pest management tactics such as use of
sify by growing multiple crops on the same land. natural enemies or resistant varieties require a
Multiple cropping on the same site may occur at the detailed knowledge of pest biology and ecology,
same time or over time. In the United States, the including life cycle, population dynamics, and
most common form of multiple cropping is crop interactions with the physical and biological envi-
rotation, in which different crops are grown on the ronment, including potential competitors, preda-
same site in different seasons or years. Cover crops tors, and parasites. Such tactics may be directed
are crops with limited market value that are grown toward preventing pest buildup rather than react-
on the site during seasons that are unfavorable for ing to high pest numbers already present in crisis
growing the main economic crops for the region. In situations. However, the design of a system in
the southeastern United States for example, a winter which pest numbers are less likely to reach crisis
cover crop of rye (Secale cereale) or crimson clover levels, due to the presence of effective natural ene-
(Trifolium incarnatum) may be grown in a field mies for example, requires advanced planning
reserved for cotton or peanut production during based on sound ecological data.
the summer. Cover crops can provide various
advantages, including erosion reduction, increased
supply of nitrogen, competition with weeds, or hay Landscape Ecology
for animals. Green manures, which are usually
legume cover crops, are grown specifically for their An individual agroecosystem does not stand alone,
nitrogen-rich residues and soil fertility benefits. since organisms, materials, and energy move freely
Intercropping, or mixed cropping (Fig.29), refers to in and out of the system. Landscape ecology exam-
the growing of two or more crops at the same time ines the agroecosystem in the context of the
on the same land. The practice is quite common in surrounding region or landscape, an essential
A 83

Agroecology, Figure 28 Residues from a previous rye crop cover the soil surface between plant rows in
this conservation tillage system.

approach in dealing with migrating insects or Many natural ecosystems are highly depen-
regulated pests. The condition of field borders, dent on recycling, since cycles of nutrients, water,
hedgerows, and adjacent fields critically affects and other materials tend to be relatively closed.
pest management within a specific field. Move- In contrast, an agroecosystem is not a closed sys-
ment of pesticides, fertilizers, and other potential tem, because its purpose is to produce harvest for
pollutants from the agroecosystem to natural eco- export to other ecosystems. This process depletes
systems is a major environmental concern. essential resources from the agroecosystem
A Agromyzidae

Agroecology, Figure 29 One of the simplest forms of intercropping is to use one crop as a windbreak,
like the sugarcane planted along with the eggplant crop shown here.

which must be restored if the system is to remain Gliessman SR (1998) Agroecology: ecological processes in
productive. The movement of essential resources sustainable agriculture. Sleeping Bear Press, Chelsea, MI
Jackson LE (ed) (1997) Ecology in agriculture. Academic
into the agroecosystem and the recycling of exist- Press, San Diego, CA
ing resources are therefore critical concerns, to Powers LE, McSorley R (2000) Ecological principles of
anticipate and ensure that supplies of critical agriculture. Delmar Thomson Learning, Albany, NY
resources for agricultural production will be con-
served over time to sustain future agricultural
production. Agromyzidae
Organic Agriculture
Integrated Pest Management (IPM) A family of flies (order Diptera). They commonly
Conservation Biological Control are known as leaf-miner flies.
Flower Strips as Conservation Areas for Pest Flies
Management Leaf-miner Flies (Agromyzidae)
Plant Resistance to Insects
Cultural Control of Insect Pests

References A family of beetles (order Coleoptera). They com-

monly are known as primitive carrion beetles.
Altieri MA (1994) Biodiversity and pest management in Beetles
agroecosystems. Food Products Press, New York, NY
Cavigelli MA, Deming SR, Probyn KL, Harwood RR (eds)
(1998) Michigan field crop ecology. Extension Bulletin
E-2646. Michigan State University, East Lansing, MI A.I.
Coleman DC, Crossley DA Jr (1996) Fundamentals of soil
ecology. Academic Press, San Diego, CA
Collins WW, Qualset CO (eds) (1991) Biodiversity in agro- An abbreviation for active ingredient, the active
ecosystems. CRC Press, Boca Raton, FL component of an insecticide, or the toxicant.
Alarm Pheromones of Insects
A 85

Insecticides heromone also is employed when the respond-

Insecticide Formulation ing individuals are stimulated to show aggression
Insecticide Toxicity towards the attacking agent. This is common in
the social Hymenoptera; for example, the honey-
bee, Apis mellifera (Hymenoptera: Apidae), and
Alarm-Defense System many ant species respond aggressively to their
alarm pheromones.
Defensive behavior that also serves as an alarm As alarm pheromones can benefit the sur-
signaling mechanism. vival of members of the species involved, it is
common for insects that employ alarm phero-
mones to live in congregations for some or all of
their life cycle. In the case of social Hymenoptera,
Air Sacs the colony is genetically related, and in asexually
reproducing aphids, the colony is clonal.
The trachea of insects are sometimes dilated or Although the survival of siblings or clones by
expanded to form pouch-like structures called air alarm pheromone response at the cost of the
sacs. Their occurrence varies among taxa, but their attacked individual appears altruistic, in geneti-
presence lowers specific gravity and enhances air cally related colonies, genes from the individual
exchange, thus enhancing flight. will predominate in the survivors and be passed
Active Ventilation on to their kin.
Abdominal Pumping

Hemipteran Alarm Pheromones

Alarm Pheromone
Alarm Pheromones of Aphids
A pheromone released to trigger alertness, disper- (Hemiptera: Aphididae)
sion or group defense by insects.
Alarm Pheromones of Insects When disturbed or attacked, many aphid species
Chemical Ecology release alarm pheromone from droplets secreted
from tube-like structures called cornicles on their
dorsal posterior. This phenomenon has been stud-
Alarm Pheromones of Insects ied exclusively in the asexual forms and most often
in asexually reproducing wingless females. Aphids
emma napper, John. a. pickett nearby exhibit a variety of behaviors ranging from
Rothamsted, Harpenden, Hertfordshire, UK stopping feeding and moving away, to running or
dropping off the plant and even attacking the
Alarm pheromones are defined as chemical sub- predator. However, not all aphids in a group
stances, produced and released by an organism, respond. The relative risks of predation and costs
that warn or alert another of the same species of of escape, for example, cessation of feeding and
impending danger. This is exemplified by many risk of desiccation, affect the likelihood of any par-
species of aphids (Hemiptera: Aphididae) in ticular response. In studies of the peach-potato
which the pheromone is caused to be released by aphid, Myzus persicae, the pea aphid, Acyrthosi-
attack, for example, by predators, with ensuing phon pisum, and the rose-grain aphid, Metopolo-
dispersal by which individual aphids may avoid phium dirhodum, early stages were found to be less
a subsequent attack. However, the term alarm sensitive to alarm pheromone than later ones.
A Alarm Pheromones of Insects

However, older wingless M. persicae require the with resistance, which could affect mobility or sensi-
greatest stimulation of alarm pheromone before tivity of the nervous system to stimuli.
responding, while winged M. persicae, particularly Aphids that have dropped from a plant may
those not feeding, are extremely sensitive to alarm re-colonize or may move to another host plant fur-
pheromone. The lack of response from the early ther away. The turnip aphid, Lipaphis erysimi, and
stages suggests that the risk of predation to these M. persicae dislodged by alarm pheromone are less
nymphs is lower than the risk involved in ceasing likely to return to the original host plant than when
to feed and dropping from the plant. When young mechanically dislodged. Similar patterns of behav-
M. dirhodum respond to alarm pheromone, they ior are found in A. pisum. Aphids dislodged by a
do so by moving to another part of the plant rather predator or experimentally with synthetic alarm
than by dropping. Winged adults, on the other pheromone spend longer running before the
hand, are more responsive to alarm pheromone, search for a host plant began, whereas aphids dis-
perhaps because they can more readily move off lodged mechanically are more likely to begin to
the host. The sugarcane woolly aphid, Ceratova- search for a host plant immediately.
cuna lanigera, also shows different reactions to Droplets secreted from the cornicles comprise
alarm pheromones at different life stages; it shows two types of material: a volatile, rapidly vaporizing
attack behavior until adult, when the normal aphid fraction which is the alarm pheromone, and a waxy
dispersal response takes over. fraction, consisting mainly of triglycerides, that
Considerable variation is seen between aphid crystallizes on contact with foreign particles outside
species in their sensitivity to alarm pheromones of the aphids body. The waxy component appears
and in both the speed and the form of the response. to function as a sticky or quick-setting irritant to
This variation often can be explained by differ- predators and parasitoids and a releasing substrate
ences in the ecology of the species. Some aphids, for the alarm pheromone component.
particularly those tended by ants, stay on the plant The main component of the alarm pheromone
and respond by walking or waggling their abdo- (Fig.30) of many aphids is the sesquiterpene
mens rather than falling off the plant. These aphids hydrocarbon (E)- -farnesene (1). Other compo-
appear to depend more on the protection afforded nents may also be present as found in the alarm
by their ant attendants than their own defensive pheromone blend of the vetch aphid, Megoura
mechanisms. Aphids that walk away from a source viciae, which contains the monoterpenes (-)- -
of alarm pheromone tend to form new clusters a pinene, (-)- -pinene, (Z,E)- -farnesene and
short distance from the original site, thus ensuring (E,E)- -farnesene, in addition to (E)- -farnesene.
continued ant attendance. There is a high degree of cross-activity of both nat-
Susceptibility to insecticide also has been found ural alarm pheromone and (E)- -farnesene among
to correlate with responses to alarm pheromone. Sus- species within the aphid subfamilies, Aphidinae
ceptible strains produce more pheromone and and Chaitophorinae. This is typical of insect alarm
respond more quickly and in higher numbers than pheromones in general, since such cross-activity
insecticide-resistant strains. In addition, clones col- does not reduce their evolutionary value. However,
lected from around the world showing knockdown the main component of the alarm pheromone of
resistance to pyrethroid insecticides, and esterase- the spotted alfalfa aphid, Therioaphis maculata,
based insecticide resistance, showed lower levels of and the sweet clover aphid, T. riehmi, in the
disturbance to the synthetic alarm pheromone. These Drepanosiphinae, is the cyclic sesquiterpene
aphids may therefore suffer increased predation or (-)-germacrene-A (2).
parasitism in the absence of insecticides, affecting In the turnip aphid, L. erysimi, it has been
the evolutionary fitness of insecticide resistant clones. demonstrated that isothiocyanates, acquired from
This may be due to physiological effects associated chemicals in the host plants, synergize the effect of
Alarm Pheromones of Insects
A 87

the alarm pheromone. These isothiocyanates are metathoracic gland, while in nymphs it is the dorsal
likely to be released from aphid honeydew so that, abdominal glands. These are precursors to the
when there is a high number of other aphids in the metathoracic gland in adults and perform the same
immediate vicinity, the percentage of aphids defense function. The chemical content of these
responding to alarm pheromone increases. secretions is similar throughout the order; for
example, the components of the secretion of the
stinkbug Cosmopepla bimaculata are a complex
Alarm Pheromones of True Bugs mixture of hydrocarbons, aldehydes and esters. The
(Hemiptera, Heteroptera) secretion, which can be ejected from either or both
metathoracic glands in controlled amounts or even
The Pentatomidae is the dominant family of stink- resorbed, displays a defensive function as a preda-
bugs, or shield bugs. The family comprises many tor repellent. In one case, researchers have shown
species that are pests of economic importance, uncommon dedication in describing the repellency
especially in warmer climates. These insects secrete by squeezing adults in their mouths and chewing
a complex mixture of chemicals when strongly nymphs. The effects were a burning sensation and
molested. The energetic cost of the defense numbness of the tongue for up to two hours. In
response, especially production of defense chem- addition to repelling predators, the secretions
icals, is significant and considerable provocation possess alarm pheromone activity and cause adults
usually is required to cause release. In adults, to drop off plants. In the field, C. bimaculata are
the source of the defense compounds is the found highly clumped and the occurrence of large

Alarm Pheromones of Insects , Figure 30 Defense secretion of Cosmopepla bimaculata showing a typical
range of compounds produced by stinkbugs. Chemical structures of alarm pheromones are referenced
in the text by bold numbers.
A Alarm Pheromones of Insects

Alarm Pheromones of Insects, Figure 30 (Continued)

numbers living together gives an evolutionary spread the oily secretion so that the aldehydic
advantage to possessing an alarm pheromone. components can exert full irritant effect. (E)-2-
Six-carbon-long aldehydes, in particular (E)- Hexenal has been reported to have the added dual
2-hexenal, are common components of defensive functions of both alarm and aggregation phero-
secretions and are found in many families of het- mone, depending on the stimulus concentration,
eropterous insects, including the Pentatomidae, as well as use as a defense chemical. In the case of
Coreidae, Pyrrhocoridae, Cimicidae, Cynidae and the bed bug, Cimex lectularius, and Eurydema
Alydidae. it is thought that the general irritant rugosa, low concentration of (E)-2-hexenal acts as
properties of aldehydes provide a repellent effect an aggregation pheromone while high concentra-
to predators with hydrocarbons such as n-tride- tion produces an alarm response. Alternatively, it
cane, another ubiquitous component, acting to has been reported that n-tridecane, the other
Alarm Pheromones of Insects
A 89

The pentatomid bug Erthesina fullo is a major

pest of pine and hardwood trees. Both sexes pro-
duce a secretion from the metathoracic gland
causing conspecific adults to drop from plants,
fly or move away. The secretion comprises nine
identified compounds, esters and aldehydes
(about 35%) including (E)-2-hexenal and (E)-4-
oxo-2-hexenal (3) and long chain alkanes, including
50% n-tridecane. Likewise, adult and nymph
secretions from Dysdercus cingulatus (Het-
eroptera: Pyrrhocoridae) revealed 55 identified
compounds, although the major components are
again aldehydes and n-tridecane, features com-
mon with several more species of pentatomids
from the genus Chlorochroa and Piezodorus
The leaf-footed bug, Leptoglossus zonatus
(Heteroptera: Coreidae), is an economically
important pest of Brazilian corn. An extract
obtained from the metathoracic gland by immer-
sion in hexane showed that the major com-
pounds were all of six-carbon length: hexanal,
hexanol, hexyl acetate, hexanoic acid, and (E)-4-
oxo-2-hexenal (3). (E)-2-Hexenal was found in
the nymph extracts but not in the adult, an
example of the general rule that exocrine chem-
istry of heteropterous nymphs is distinct from
that of the adult. In this case, different life-stages
possess different alarm pheromone systems.
When tested individually, all components pro-
duced varying degrees of alarm response in
Alarm Pheromones of Insects, Figure 30 (Continued) adults and nymphs and even mating insects
would stop and disperse, over-riding the sex
biquitous component of defense secretions, is a
u pheromone response. These compounds are not
bifunctional pheromone for the southern green species specific and are, for instance, found in
stinkbug, Nezara viridula (Heteroptera: Pentato- L.oppositus and L. clypealis, a situation that mir-
midae), which causes dispersal at high concentra- rors the cross-activity of (E)--farnesene in
tion (one individual equivalent) and aggregation many aphid species (see above), providing more
at low concentrations. The multifunctional aspect evidence that this non-specific activity does not
of these compounds has important repercussions reduce alarm pheromone value in evolutionary
for their practical use as dispersal agents for pest terms.
species in the field, as the low concentration Adults of the bean bug, Riptortus clavatus
response of aggregation may dominate once the (Heteroptera: Alydidae), a pest of Japanese soy-
applied high concentration of compounds has bean, secrete (E)-2-hexenyl acetate in its defensive
diminished. response. This causes an alarm response in adults
A Alarm Pheromones of Insects

and nymphs. Interestingly, adults also produce Alarm Pheromones of Social

(E)-2-hexenal, and although some response was Insects (Hymenoptera)
found when tested at high concentration, there
was no response at physiological concentration, The Honeybee (Hymenoptera: Apidae)
suggesting that this compound is not an alarm
pheromone. When the honeybee (Apis mellifera) is attacked,
The examples shown above demonstrate alarm pheromones released serve to muster help
that alarm pheromones of heteropteran families and to direct the attack. Specialized guard bees
are based on a chemical selection general to present at the nest entrance carry out attacks.
insects of a wide taxonomy and show little spe- Although these guards are relatively few compared
cies specificity. The alarm behavior caused by to the colony population, release of alarm phero-
high concentrations of n-tridecane or (E)-2- mone can result in synchronized attacks by more
hexenal is rationalized easily but the aggregation than 100 workers against an intruder. Guard bees
due to low concentrations is more difficult to initiate attacks by raising their abdomens, protrud-
explain. Perhaps these substances are constantly ing their stings and releasing alarm pheromone
emitted in very small quantities due to their vol- from the sting chamber. The workers then alert the
atility and act so as to direct individuals to a rest of the hive by wing beating, aiding dispersal of
region where conspecifics can be found and the pheromone, and by running into the hive. After
therefore where food most probably is located, a few seconds, many excited bees may rush out of
as well as the defensive advantage in being part the hive entrance and search, or stop and assume a
of a large group. characteristic tense and aggressive posture with a
Triatomine bugs (Heteroptera: Reduviidae) slightly raised body, wings extended, mandibles
are blood-sucking insects that live throughout agape and antennae waving. In this highly activated
the Americas and cause public health problems state, they will fly to attack at the slightest further
by transmitting the protozoa Trypanosoma cruzi, provocation. These two stages of alarm response
the causative agent of Chagas disease, to humans. are called alerting and activation and are charac-
Secretions from Brindleys gland (a simple sac, teristic of alarm pheromones.
metathoracic in origin) of several species all Alerted workers need to search for and dis-
revealed isobutyric acid as the major component. cover the enemy to prevent any further threat. To
Subsequently, other short-chain and branched- do this, they rely on other cues to direct the attack
chain fatty acids have been identified, and such as odor, jerky movement and hairy body
together with isobutyric acid, they act as a pow- covering. Once the threat is located, it is stung,
erful defensive secretion. Pure isobutyric acid injecting a dose of venom. However, the shaft of a
vapor, however, also caused an alarm response in sting is barbed and a bee is unable to withdraw it
Rhodnius prolixus while another report revealed from the skin of vertebrates, so the sting, together
that low concentrations of isobutyric acid with associated motor apparatus and glands, are
attracted R. prolixus adults. This defense com- severed from the bee as it attempts to fly away
pound therefore shows the same multifunctional and are left attached to the enemy. The severed
alarm and aggregation properties as described sting apparatus continues to pump venom into
for components of the stinkbug secretions (see the victim and alarm pheromone is dispersed
above). Triatomine bugs are inactive and hide from the exposed under-surface of the sting shaft
during the day, congregating in protective sites. membrane to mark an enemy and make it a more
This aspect of group living can help explain the obvious target.
evolutionary advantage in possessing aggrega- The main alarm pheromone component of
tion and alarm responses. the sting gland was identified in 1962 as isopentyl
Alarm Pheromones of Insects
A 91

acetate (4). Although a number of other com- threat is closer to the nest and may be specialized
pounds are known to be present, isopentyl acetate to deal with vertebrate predators. Also, workers
and (Z)-11-eicosen-1-ol (5) account fully for the of the Texas leaf-cutting ant, Atta texana (Formi-
activity of the sting pheromone. The roles of the cidae: Myrmicinae), have a lower threshold for
two compounds in the pheromone appear to dif- alarm pheromone response than the queen and
fer, with (Z)-11-eicosen-1-ol responsible for pro- males. Other factors governing the type and
longing the activity of isopentyl acetate. Other intensity of alarm response are the age and size of
compounds such as 1-hexanol and 1-butanol ant colony. When alarm pheromone is present in
increase the number of bees responding. sufficient concentration to excite the workers,
Stinging bees often grip an enemy with their other stimuli are needed to direct an attack.
mandibles and deposit an alarm substance. At the Workers often will touch everything they encoun-
hive entrance, more bees examine mandibular ter and the full-scale alarm response may rely on
gland extracts of worker honeybees applied to fil- additional cues, such as the presence of an alien
ter paper than examined unscented filter paper. object. Alarm pheromones also may function
2-Heptanone (6) has been identified from the with acoustic alarm signals. Ant species in the
mandibular gland secretion, and when filter papers sub-family Dolichoderinae produce vibration
or small corks carrying 2-heptanone were placed signals using their mandibles to scratch the
at the hive entrance, the guard bees were alerted ground or the abdomen to hit the ground, increas-
and attacked them. As the mandibles are used for ing alarm behavior in other workers. Vibrations
grasping an intruder, it seems likely that the main are produced also by leaf-cutting ants which act
function of 2-heptanone is to label the intruder to as warning signals. Alarm pheromones also are
be attacked. used by ants to attract attention if they are
Under certain circumstances, honeybee alarm trapped, and may be released by reproductive
pheromones are repellent. The presence of alarm ants just before mating flights to ensure that
pheromone deters honeybees from foraging at aggressive workers protect them from potential
dishes of sugar syrup and from exposing their predators.
Nasonov glands and fanning which normally The context in which a worker encounters an
attracts other bees. Furthermore, a high concen- alarm pheromone also influences the response.
tration of alarm pheromone repelled foraging Workers of the grass-cutting ant Atta capiguara
beesfrom crops including oilseed rape, normally (Formicidae: Myrmicinae) are less likely to show
highly attractive to bees, in an area that had many alarm behavior if already engaged in a task. Forag-
honeybee colonies. ers carrying leaves do not respond to alarm phero-
mone, whereas minor workers and foragers that
are not carrying leaves do respond.
Alarm Pheromones of Ants Ant alarm pheromones may be produced
(Hymenoptera: Formicidae) from one or several sources. The army ants or
Eciton ants (Formicidae: Ecitoninae) and the rare
The Formicidae is a huge family comprising Leptanilla sp. (Formicidae: Leptanillinae) of
thousands of ant species, all of which are social Indonesia have large mandibular glands, which
insects, living in colonies that vary hugely in size. are believed to be the sole source of the alarm
Members of an ant colony may be differentiated pheromone. However, other species rely on a
into castes that specialize in carrying out particu- combination of secretions from several glands.
lar tasks and vary in their response to alarm Formica and Myrmica species use products from
pheromone. Soldiers show a more aggressive the poison and Dufours gland (both opening
response, are more likely to respond when the near the base of the sting), as well as the
A Alarm Pheromones of Insects

andibular gland. Ponerine ants (Formicidae:

m believed to be patrollers as they have a stronger
Ponerinae) use secretions from the pygidial gland response to alarm pheromone than foragers and
as alarm components, whereas the poison gland soldiers. Minors of other Atta species are also
is the most important gland of several other spe- more efficient at recognizing intruding ants than
cies, including the harvester ant, Messor barbarus other castes.
(Formicidae: Myrmicinae). As is the case with honeybees described above,
Ants are able to detect and respond to spe- the complete alarm response can be described by a
cific isomers of their alarm pheromone. Myrmica number of behaviors. These behaviors and their
rubra and M. scabrinodis use 3-octanol (7) as the elicitors have been dissected in an elegant piece of
alarm pheromone produced from mandibular research on the African weaver ant, Oecophylla
glands. Experiments carried out using the two longinoda. The major workers produce a secretion
optical isomers (8a, b) of this compound showed from the mandibular gland comprising four active
that M. rubra workers only responded to one of components: hexanal, hexanol, (E)-2-butyl-2-
the isomers, (R)-3-octanol (8a), while M. scabrin- octenal (10), a dimer of hexanal produced chemi-
odis workers reacted more strongly to the natural cally by self-condensation, and 3-undecanone
9:1 mixture of R and S isomers. This work sug- (11). Hexanal, a highly volatile component with an
gests that there may not only be specific chemi- active space of 510 cm (the area around an emis-
cals but also species-specific mixtures of isomers. sion where the concentration is at or above that
Many other Myrmica species have 3-octanol and required for a behavioral response), causes the
3-octanone as alarm pheromones but may have ants to be alerted, making quick runs in random
species-specific ratios of the two, which allow ant and changing direction with mandibles open and
species to only show a full alarm reaction to their antennae waving. Hexanol attracts directly to the
specific alarm pheromone blend. source at a range of 15 cm; it is repellent at very
Myrmica species of grass-cutting ants share close range and also causes further excitement. As
the main component of alarm pheromone, the hexanol disperses, 3-undecanone is attractive
4-methyl-3-heptanone (9), but they have species- over this close range and, along with (E)-2-butyl-
specific modifying components. The response to 2-octenal, acts as a marker for attack and biting
4-methyl-3- heptanone was compared to that elic- to hold the source occurs. This process is called
ited by the bodies of workers that had their heads alocal attack. In addition, O. longinoda also has a
crushed to release the natural alarm pheromone. mass attack alarm response. The poison gland of
4-Methyl-3-heptanone and bodies caused the same the major and minor workers contains venom that
level of attraction but the full range of alarm beha is ejected by raising the gasters above vertical when
vior was seen only with the bodies. In contrast, an attacked object is held in the jaws. The venom
workers of the giant tropical ant, Paraponara contains a blend of straight chain hydrocarbons
clavata (Formicidae: Paraponera), produce two and formic acid. Formic acid initiates approach
components, 4-methyl-3-heptanone and 4-methyl- and attack while n-undecane causes mandible
3-heptanol. opening, gaster raising and also short-range
Atta capiguara is a grass-cutting ant species approach to the source. The combination of these
that lives in colonies with hundreds of thousands behaviors allows location and initial attack of still
of workers. Workers are polymorphic, varying in and moving objects followed by recruitment of
size from small minors and medias to the larger workers to continue attack.
foragers and soldiers. Minors and medias do most The properties that make alarm pheromone
of the nest tasks whereas foragers collect the grass; cues for conspecifics also enable them to act as cues
however, minors often are found on foraging trails for parasites and predators of ants to find their
despite the fact they do not carry grass. They are prey. Apocephalus paraponerae is a parasitic fly that
Alarm Pheromones of Insects
A 93

attacks the ant Paraponera clavata. Females and Cockroach Alarm Pheromones
males of A. paraponerae are attracted to injured, (Blattodea: Blattidae)
fighting or freshly killed workers. After finding a
worker, the female lays a few eggs, which will hatch Defensive secretions are well known in cockroaches.
and then feed on the victim for 37 days. Both male They are produced from ventral inter-segmental
and female A. paraponerae also feed on the wounds glands and comprise an organic and an aqueous
of the injured workers and gather near their vic- phase. In the case of the Florida woods cockroach,
tims to mate. The heads of P. clavata workers Eurycotis floridana (Dictyoptera: Blattidae), 90% of
contain two chemicals, 4-methyl-3-heptanone and the organic phase (which comprises 85% of the total
4-methyl-3-heptanol, which are particularly attrac- secretion) is (E)-2-hexenal, a compound found in
tive to A. paraponerae. These compounds are com- many heteropteran bugs and discussed above. The
mon alarm pheromone components of ants and rest of the organic secretion comprises approxi-
are released when the workers are stressed. How- mately 40 other components, including mainly alde-
ever, it has been suggested that because these para- hydes, alcohols and carboxylic acids, while the
sites use alarm pheromone for finding their host, aqueous phase contains gluconic acid, glucose and
P. clavata may be under pressure to reduce the gluconolactone. The secretion acts as a conspecific
amount of alarm pheromone released and that this alarm pheromone in these gregarious insects with
ant may even have lost alarm behavior response as nymphs responding at lower concentrations than
a result of this pressure. Similarly, the zodariid spi- adults do. Ethanolic extracts of the American cock-
der, Habronestes bradleyi (Zodariidae), a predator roach, Periplaneta americana, also repel conspecifics
of the meat ant, Iridomyrmex purpureus, detects from aggregations in daytime shelters. However,
the alarm pheromone, in this case 6-methyl-5- there was no evidence that this repellent is released
hepten-2-one (12), given out by fighting workers by living insects as an alarm pheromone but is
and uses it to locate its prey. instead endogenously produced from dead insects
and is effective against other cockroaches with
diverse phylogenetic relationships. The effect, there-
Alarm Pheromones of Thrips fore, is not pheromonal, as the authors explain the
(Thysanoptera: Thripidae) activity in terms of unsaturated fatty acids (oleic,
linoleic and linolenic acids) which emerge as signals
Thrips are small, economically important pest of death and injury among organisms from a wide
insects, often known as thunderflies. The defensive phylogenetic background. Both of the reports
behavior of thrips includes raising and lowering described above provide evidence that the use of
the abdomen and secretion of a droplet of anal alarm pheromones to increase dispersal for pest
fluid highly repellent to predatory ants. Western management purposes will be of limited value. Due
flower thrips, Frankliniella occidentalis (Thys- to the aggregation effect at low concentration, treated
anoptera: Thripidae), are not social but tend to be areas could become attractive. Also, if low concen-
found in clumped distributions. Adults and nymphs trations were used as attractants in a lure and kill
of western flower thrips produce an anal droplet approach, dead insects would repel others before
containing decyl acetate and dodecyl acetate (13) they become ensnared.
in a molar ratio of 1.5:1. Each component, at levels
of 1 ng, produces the alarm response of walking Alarm Pheromones of Beetles
away from the source or dropping from leaves. The (Coleoptera)
response, however, is only over short distances and
limits the potential for pheromone use in pest Despite the vast numbers of species in the order
management. Coleoptera, inhabiting a wide range of ecological
A Alarm Pheromones of Insects

niches, little is known of the existence of behavioral Conclusions

responses to alarm pheromones they may possess.
Species of beetles that are group-living are most It can be seen that alarm pheromones are used
likely to demonstrate alarm responses. Gyrinid bee- widely by a broad taxonomic diversity of insects
tles (Coleoptera: Gyrinidae), known as whirligig and elicit equally varied behavioral responses,
beetles, live in open habitats on fresh water surfaces including escape or aggressive behavior. Alarm
and typically aggregate in groups containing hun- pheromones are generally low molecular weight,
dreds of individuals, dispersing in the evening to organic compounds and so are volatile, dispersing
forage. Although easy to detect by fish, they are sel- quickly, and do not persist in the environment. In
dom preyed upon due to a repellent secretion addition, the chemical nature of the alarm phero-
released as a last resort to physical attack. The secre- mone often is unstable, increasing the lack of per-
tion also acts as an alarm pheromone over short sistence. This allows conspecifics to be alerted very
distances, increasing locomotory activity and defen- quickly over a fairly large area and yet not cause
sive movement, such as diving and active underwa- false alarm after the danger has passed. Alarm pher-
ter swimming. Although alarm dispersal after attack omones are often produced in glands responsible
can occur, the aggregations of beetles themselves for biosynthesis, storage or release of defense secre-
indicate to experienced predators to avoid the group tions. This association between alarm pheromones
and confer an aposematic effect acting at the group and defense glands, including those near the sting
level rather than the individual level. or mandibles, has led to the hypothesis that alarm
pheromones have evolved from chemicals that
originally had a defensive role, or are themselves
Lacewing Alarm Pheromones defense compounds that have taken on an addi-
(Neuroptera: Chrysopidae) tional alarm pheromonal role. The fact that known
defense components have additional multifunc-
The green lacewing, Chrysoperla carnea (Neu- tional pheromonal roles of alarm (high concentra-
roptera: Chrysopidae), is an important predator of tion) and aggregation (low concentration) also
pest aphids and, as such, is a beneficial insect. It dis- points to the possibility that these pheromonal roles
charges a malodorous secretion from glands at the have evolved from compounds originally used for
anterior of the prothorax. The major component of defense. Of particular interest is the common lack
this secretion has been identified by gas chroma- of species specificity found in alarm pheromones,
tography, mass spectroscopy and chemical synthe- which is in contrast to that of other pheromones.
sis as (Z)-4-tridecene (14) and gas chromatography Sex pheromones, for example, are so specific that
coupled electroantennagrams revealed that it is they can be the sole identifiable trait in defining
detected by the lacewing antennae. Predatory ants morphologically identical populations, such as
displayed avoidance behavior in response to it, sug- within the species complex of the sandfly, Lutzo-
gesting a defensive function, and in laboratory myia longipalpis (Diptera: Psychodidae). However,
experiments, adult lacewings avoid entering areas the alarm pheromone of different aphid species is
where it is present. In the field, it acts as an antago- (E)--farnesene and different species of Atta grass-
nist to trap catches using known attractants and, as cutting ant use 4-methyl-3-heptanone. In addition,
such, could be described as an alarm pheromone. production of (E)-2-hexenal and n-tridecane is
Another species of lacewing, Peyerimhoffina ubiquitous as multifunctional pheromone compo-
gracilis, also produces the identical compound. As nents in terrestrial true bugs, and it is possible that
lacewings are not known to be gregarious, the exact (E)-4-tridecene may reveal itself tobe common in
ecological purpose of this compound is being green lacewings. Discrimination between these
investigated. behavioral signal compounds, therefore, is not
Alderflies and Dobsonflies (Megaloptera)
A 95

essential to their function as alarm pheromones, robably aid the dorsal diaphragm in providing sup-
and there may even be evolutionary benefits in port for the heart, the part of the dorsal vessel in the
being able to respond to alarm pheromones of abdomen. The muscle fibers fan out from a small
related species of insects. point of origin on the lateral wall of the dorsum to a
broad insertion on the heart in many insects, pre-
senting the typical delta appearance. In some insects,
however, the delta shape is not so evident. Some
alary muscle fibers pass beneath the heart and extend
laterally from side to side, and thus help to support
Aldrich JR, Blum MS, Lloyd HA, Fales HM (1978) Pentato-
mid natural products. Chemistry and morphology of the heart. In places, the fibers may also run parallel
the III-IV dorsal abdominal glands of adults. J Chem to the long axis of the heart for a short distance. The
Ecol 4:161172 pairs of alary muscles tend to agree with the number
Bowers WS, Nault LR, Webb RE, Dutky SR (1972) Aphid
of pairs of ostia, the (usually) lateral openings in the
alarm pheromone: isolation, identification, synthesis.
Science 177:11211122 dorsal vessel that allow hemolymph to flow into the
Bradshaw JWS, Baker R, Howse PE (1979) Multicomponent heart in the abdomen. Alary muscles generally do
alarm pheromones in the mandibular glands of major not occur in the thorax, but in some insects, a few
workers of the African weaver ant. Physiol Entomol
ostia open outward in the thorax, allowing hemo-
Dawson GW, Griffiths DC, Pickett JA, Woodcock CM (1983) lymph to flow outward.
Decreased response to alarm pheromone by insect- In addition to support, the alary muscle may
resistant aphids. Naturwissenschaften 70:254255 assist in the expansion (diastole) of the heart after
Free JB, Pickett JA, Ferguson AW, Simpkins JR, Smith MC
(1985) Repelling foraging honeybees with alarm phero- each contractile wave passes a given point, and
mones. J Agric Sci 105:255260 thus aid in pulling hemolymph into the incurrent
Hlldobler B, Wilson EO (1990) The ants, 1st ed. The Belknap ostia. They are not necessary for diastole, however,
Press of Harvard University Press, Cambridge, MA
as evidenced by severing them with little or no
Nault LR, Phelan PL (1984) Alarm pheromones and sociality
in pre-social insects. In: Chemical ecology of insects. apparent effect on the heart beat.
Chapman and Hall, London, UK, pp 237256
Pickett JA, Griffiths DC (1980) Composition of aphid alarm
pheromones. J Chem Ecol 6:349360
Alate (pl., Alatae or Alates)

The winged forms of insects, particularly aphids.

Alarm-Recruitment System

Recruitment of nest members to a particular loca- Alderflies

tion to aid in colony defense.
Sociality of Insects Members of the family Sialidae (order Megaloptera).
Alderflies and Dobsonflies

Alary Muscles
Alderflies and Dobsonflies
james l. nation (Megaloptera)
University of Florida, Gainesville, FL, USA
lionel stange
The alary muscles, so named because of their general Florida Department of Consumer and Agricul-
wing or delta shape in many insects, lie immediately tural Services, Division of Plant Industry, Gainesville,
on top of the dorsal diaphragm. The muscles FL, USA
A Alderflies and Dobsonflies (Megaloptera)

The order Megaloptera comprises about 190 spe-

cies in 60 genera in two families. All the larvae are
aquatic. The larvae, especially of Corydalinae, are
among the most primitive of the Holometabola.
The metamorphosis from larva to adult is rela-
tively simple.
The family Sialidae, commonly called alder-
flies, is a small group of about 70 species in about
eight genera. They are worldwide. Most of the
adults have a similar appearance and are usually
dark brown to black in coloration. They lack ocelli
and have the fourth tarsomere bilobed. They are
an ancient group known from the Permian, about
200 million years ago, and evidently have not
evolved much since then. In fact, the wing vena-
tion has many features in common with the Pro-
toperlaria, a fossil order considered by some as
ancestral to the Plecoptera. The adult life span is
probably short since the reduced mouthparts do
not seem adapted for extensive feeding. The eggs
are laid in rows, forming large masses situated on Alderflies and Dobsonflies (Megaloptera),
branches, bridges, and other objects overhanging Figure31 Adult dobsonfly (Corydalidae).
the water. The larvae hatch and fall into the water
where they are predacious on other aquatic The Corydalinae with about 60 species, is dis-
insects, especially caddisflies. There are as many tributed in the New World (three genera), South
as 10 larval instars which may last up to two or Africa (one genus) and Asia (five genera). This
more years until pupation. The larva crawls out of subfamily does not have pectinate antennae and
the water and digs into the bank to form an the head is usually quadrate, often with a postocu-
earthen cell several feet from theedge of the water. lar spine. Often there are more than four crossveins
The genera are restricted geographically. The between the radius and radial sector. The male ter-
genus Sialis Latreille is Holarctic, Protosialis minalia are distinctive with a well developed ninth
Van der Weele is South American, Austrosialis gonostylus. Many of the species are very large.
Tillyard and Stenostialis are Australian, Haplosia- Corydalis Latreille is the largest genus in the New
lis Navas is from Madagascar, Leptosialis is African, World with about 30 species. The males of this
Indosialis Lestage is Oriental and Nipponsialis genus often have the mandibles greatly extended
Kuwayama is Japanese. The larva has seven lateral which is similar to Acanthacorydalis Van der Weele
processes and the abdomen terminates in an elon- from Asia. Platyneuromus christil (Navs) from
gate process. The only world compilation is by Central America, has a tremendously expanded
Van der Weele (1910) but is greatly out of date. postocular flange. The larva has eight lateral pro-
The family Corydalidae, or dobsonflies, is cesses and the abdomen ends in a pair of claw-like
characterized by having three ocelli and the anal structures. In America, the larvae are called hell-
region of the hindwing is very wide, folded fanlike grammites and are used for fishing. In one case, a
at rest (Fig.31). The fourth tarsomere is not modi- larva was found inside a fish stomach many hours
fied. There are several hundred species in about 20 after ingestion. Glorioso (1982) has provided a
genera and two subfamilies. good reference.
Alexander, Charles Paul
A 97

The Chauliodini was reviewed by Kimmins References

(1954) who recognized 12 genera. The genera are
restricted in distribution similar to the Corydalinae Beier M (1973) The early naturalists and anatomists during
and are found in the Cape region of South Africa the renaissance and seventeenth century. In: Smith RF,
(two genera), North America (four genera), Chile Mittler TE, Smith CN (eds) History of entomology.
Annual Reviews, Inc., Palo Alto, CA, pp 8194
(three genera), Australia (one genus), Madagascar Nordenskild E (1935) The history of biology: a survey.
(one genus), and Asia (four genera). Males and Tudor, New York, NY, 629 pp
rarely females have pectinate antennae in some

A family of bugs (order Hemiptera). They some-
Glorioso MJ (1981) Systematics of the dobsonfly subfamily times are called broad-headed bugs.
Corydalinae (Megaloptera: Corydalidae). Sys Entomol Bugs
Kimmins DE (1954) A new genus and some new species of
the Chauliodini (Megaloptera), with notes on certain
previously described species. Bull Br Mus Nat Hist Aleyrodidae
Entomol 3:417444
Van der Weele (1910) Megaloptera. Collections Zoologiques A family of insects in the order Hemiptera. They
du Baron Edm. de Selys Longchamps, fasc 5, 93 pp
sometimes are called whiteflies.
Aldrovandi, Ulisse (Ulysse,
Alexander, Charles Paul
Ulisse Aldrovandi was born in Bologna, Italy,
in 1522. He studied law in Bologna, and then Charles Alexander was born in New York state on
philosophy and medicine in Padua and Rome, earn- September 25, 1889. He entered Cornell Univer-
ing a doctorate in medicine in 1552. In 1560, he was sity in 1909, receiving B.Sc. and Ph.D. degrees in
appointed professor in Bologna, a position that he 1913 and 1918, respectively. He was employed as
held for 40 years. He lectured mainly on pharma- systematic entomologist in the Snow Entomologi-
cology, but he collected natural history objects and cal Museum at the University of Kansas in 1917
employed artists to draw them. He published four 1919 and then by Illinois Natural History Survey
large volumes during his lifetime, but his friends in 19191922. Next he moved to Massachusetts
and pupils used his voluminous manuscripts to Agricultural College and was placed in charge of
publish 10 more volumes after his death. His (1602) teaching entomology. He served as chairman of
De animalibus insectis libri VII was the first book the Department of Entomology and Zoology for
to be published on insects, although the insects 10 years, for the last three of which he was dean of
included various other kinds of invertebrates. A the School of Science (of what had by then become
chapter was devoted to the structure of the insect the University of Massachusetts). He was presi-
body. Insect reproduction and metamorphosis were dent of the Entomological Society of America in
described; respiration and the senses of touch, taste, 19411943 (two terms). His almost exclusive sub-
and smell are discussed, and the life of honey bees is ject of research was the family Tipulidae (crane
described. He died in 1605. flies) about which he published over 1,000 papers
A Alfalfa Leafcutting Bee, Megachile Rotundata (Hymenoptera: Megachilidae)

and described over 10,000 species, an enormous commercial scale. M. rotundata is of eastern Medi-
production. After retirement from teaching, he terranean origin and was first found in North
moved his insect collection to his house and con- America in the 1940s near seaports. It probably
tinued working on it until the death of his wife, gained entry as diapausing pre-pupae within tun-
Mabel, in 1979. Two years later he transferred his nels in the wood used to make shipping crates or
collection to the National Museum of Natural pallets.
History in Washington, DC. He died at home in
Massachusetts on December 12, 1981.
Life History

Reference Once a suitable tunnel has been found, the female

uses her mandibles to neatly cut oblong pieces of
Byers GW (1982) In memoriam Charles P. Alexander 1889 leaves or flower petals which she uses to build
1981. Journal of the Kansas Entomological Society cells end to end in the tunnel, starting at the far
end and finishing near the entrance (Fig.32).
About 15 leaf pieces are arranged in overlapping
layers and cemented together to form a thimble-
Alfalfa Leafcutting Bee, Megachile shaped cell with a concave bottom. The cell is then
rotundata (Hymenoptera: provisioned with nectar and pollen. During this
Megachilidae) process, the female enters the tunnel head first,
regurgitates the nectar, then turns around to
mark s. goettel remove the pollen from the scopa (the pollen-
Agriculture and Agri-Food Canada, Lethbridge, collecting hairs on the underside of her abdomen)
AB, Canada and tamps the pollen into the nectar with the
tipof her abdomen. The provisions for each cell
The alfalfa leafcutting bee, Megachile rotundata consist of about two-thirds nectar and one-third
Fabricius (Hymenoptera: Megachilidae), has been pollen, requiring 1525 provisioning trips. It is
successfully semi-domesticated within the last 50 while collecting the nectar and pollen that the
years to pollinate alfalfa for seed production in bees pollinate the flowers that they visit. When
North America. Honey bees are inefficient polli- the cell has been adequately provisioned, the
nators of alfalfa and, although bumbles bees and female lays a single egg directly on the surface of
some other wild bees are efficient pollinators, they the provisions and then caps the cell with several
have proved difficult to manage. The use of the circular leaf pieces. She then proceeds to con-
alfalfa leafcutting bee has succeeded in greatly struct the next cell, repeating this process until
increasing the seed yield of alfalfa. In western the tunnel is filled. She then plugs the end of the
Canada, the average alfalfa seed yield using this tunnel with 1015 leaf pieces cemented together
bee exceeds 300 kg/ha, whereas without it is usu- to form a plug. Females continue filling tunnels
ally less than 50 kg/ha. The genus Megachile con- with cells until pollen and nectar sources are no
tains many species that nest in tunnels in dead longer available. Upon hatching, the larva imme-
trees or fallen logs. Most are solitary, but M. rotun- diately begins feeding on the provisions within its
data is gregarious and, although each female cell, undergoing four instars before reaching
constructs and provisions her own tunnel, she maturity. It then deposits a ring of fecal pellets
willtolerate close neighbors. This behavioral char- within the cell and spins a tough silken cocoon
acteristic is one of the main reasons why this within which it overwinters as a diapausing
species has been amenable to management on a pre-pupa. During the feeding period, the waste
Alfalfa Leafcutting Bee, Megachile Rotundata (Hymenoptera: Megachilidae)
A 99

Alfalfa Leafcutting Bee, Megachile Rotundata (Hymenoptera: Megachilidae), Figure 32 Alfalfa leafcutting
bee, Megachile rotundata. (1) Adult on alfalfa flower. Flowers are pollinated while the bees visit the flowers
to collect nectar and pollen for provisioning their cells. (2) A single egg is deposited on the surface of pollen/
nectar provisions within a cell which the female constructs within tunnels using oblong pieces of leaves or
flower petals. (3) The egg placed into the cell hatches within 2 to 3 days and the larva immediately begins to
consume the provisions. The larva pupates after undergoing 4 instars. (a) single egg, (b) 3rd instar, and (c) 4th
instar larvae within the cell. Cell caps have been removed. (4) X-ray of leafcutting bee cells used to determine
quality of c ommercial bees. (a) empty cell, (b) chalkbrood cadavers, (c) Pteromalus venustus parasitoid cocoons.
(5) Nesting boards separated to show arrangement of bee cells constructed within the tunnels. In the fall, the
boards are removed from the field and the cells are stripped from the boards using specialized automated
equipment. (6) Chalkbrood cadaver within the cell. Note the ring of frass deposited on the outside edge of the
cell. Normally, the larva would spin a tough silken cocoon within which it overwinters as a diapausing pre-pupa.
Larvae infected with chalkbrood usually succumb just after defecating and just prior to cocoon spinning.
A Alfalfa Leafcutting Bee, Megachile Rotundata (Hymenoptera: Megachilidae)

products of digestion are accumulated internally R.H. and 30C. A few days before the bees are due to
until the larva defecates just before forming the emerge, the trays are moved to especially designed
cocoon. In the spring, the pre-pupa pupates. After shelters spaced throughout the alfalfa seed fields. By
a pupal period of 34 weeks, the adult emerges selecting the date when incubation begins, and if
and chews its way out of the cocoon. Mating takes necessary manipulating the incubation tempera-
place soon after emergence of the adults. Females ture, the emergence of the bees can be adjusted to
store enough sperm from a single mating to fertil- coincide with the start of alfalfa bloom.
ize all of their eggs. Soon after mating, the females An advantage of the loose cell system of
seek out suitable sites in which to excavate tun- management is that it facilitates the control of
nels or select suitable preexisting ones, either nat- parasitoids, predators and disease, and assess-
ural or man-made. ment of the quality of the progeny. Leafcutting
beekeepers routinely send samples of cells to
specialized leafcutting bee cocoon testing cen-
Domestication ters, where they are x-rayed and incubated to
provide estimates of numbers of intact cells,
The gregarious nature of M. rotundata and its incidence of parasites and pathogens, and sex
willingness to accept artificial domiciles has per- ratio. These data are used to determine stocking
mitted the commercial scale management of this rates and to set a price if bees are to be marketed.
species for crop pollination. Initially, observant The proportion of females, which is usually only
alfalfa seed producers in the northwestern U.S.A. about a third, is of particular interest because
noticed that this species, which had undergone a they are the primary pollinators.
population increase following natural establish-
ment, would nest in man-made structures such as
shingled roofs and they started to provide artificial Natural Enemies
tunnels by drilling holes in logs positioned around
the edges of the seed fields. The next step was to About 20 species of insects are known to parasitize
provide nests consisting of wooden blocks drilled or prey on the immature stages of the alfalfa leaf-
with closely spaced tunnels. Although reasonably cutting bee. The most important of these are sev-
successful on a small scale, this method was not eral species of chalcid wasps, including Pteromalus
suitable for the management of the large numbers venustus Walker, Monodontomerus obscurus West-
of bees (50,00075,000/ha) required for commer- wood, Melittobia chalbii Ashmead and Diachys
cial alfalfa seed production. Consequently a loose- confusus (Girault). The most widespread and dam-
cell system was developed. This system uses 10 mm aging is P. venustus, which probably arrived in
thick boards of wood or polystyrene which are North America with its host. The female parasit-
grooved on both sides and stacked together to form oid pierces the host cocoon with her ovipositor,
hives of closely packed tunnels about 7 mm in stings the larva or pupa to paralyze it, and then
diameter and 150 mm in length. At the end of the lays some eggs on its surface. The parasitoid larvae
season, the boards are separated and the cells then feed upon the bee larva eventually killing it.
removed using specialized automated equipment. Normally 1520 adult P. venustus emerge from
After being stripped from the boards the cells are each host cocoon.
tumbled and screened to remove loose leaf pieces, Two other enemies, which are more of bio-
molds and some parasites and predators. The clean logical interest than economic significance, are
cells are then placed in containers for overwinter- several species of cuckoo bees, Coelioxys (Mega-
ing storage at about 50% R.H. and 5C. In the spring, chilidae) and the brown blister beetle, Nemogna-
the cells are placed in trays for incubation at 70% tha lutea LeConte. Cuckoo bees are very similar to
Alfalfa (Lucerne) Pests and their Management
A 101

leafcutting bees, but lack the structures required way through infected cadavers in order to exit
for collecting pollen. The female cuckoo bee lays their nesting tunnels. Chalkbrood can be ade-
her egg in the partially provisioned cell of the leaf- quately managed through strict hygiene and
cutting bee while the rightful owner is out forag- decontamination of the bee cells, nest materials
ing. When partly grown, the cuckoo bee larva kills and shelters. Initially, decontamination was per-
the leafcutting bee larva and usurps the provi- formed by dipping in household bleach. However,
sions. Brown blister beetle females lay their eggs fumigation with paraformaldehyde has become
on flowers and the first instar larvae (triungulins) the method of choice, and is highly effective for
attach themselves to any bee that visits the flower. the control of both A. aggregata and foliar molds,
When the bee returns to its nest, the triungulin which can sometimes pose a health risk to the
detaches and begins feeding on the cell contents, beekeeper.
destroying 2 or 3 cells before reaching maturity. Bees
Several stored-product insects including the
driedfruit moth, Vitula edmandsae serratilinnella
Ragonot, and stored-product beetles such as the
sawtoothed grain beetle, Oryzaephilus surinamen-
sis (Linnaeus), the red flour beetle, Tribolium cas-
Goerzen DW, Watts TC (1991) Efficacy of the fumigant
taneum (Herbst) and the confused flour beetle, paraformaldehyde for control of microflora associated
Tribolium confusum (Jacquelin du Val) can cause with the alfalfa leafcutting bee, Megachile rotundata
serious damage during overwintering storage, (Fabricius) (Hymenoptera: Megachilidae). BeeScience
especially if sanitation practices are lax. Goettel MS, Richards KW, Goerzen DW (1993) Decontami-
Most of the parasitoids and predators can be nation of Ascosphaera aggregata spores from alfalfa
largely controlled by proper construction of hives leafcutting bee (Megachile rotundata) nesting materials
and nesting materials, physical removal during by fumigation with paraformaldehyde. BeeScience
the loose-cell processing and strict hygiene during Hill BD, Richards KW, Schaaljie GB (1984) Use of dichlorvos
storage. However, successful control of the major resin strips to reduce parasitism of alfalfa leafcutter
pest, the chalcid P. venustus, often requires care- bee (Hymenoptera: Megachilidae) cocoons during
incubation. J Econ Entomol 77:13071312
fully controlled fumigation using dichlorvos (2,
Richards KW (1984) Alfalfa leafcutter bee management in
2-dichloro-vinyl dimethyl phosphate) resin strips. Western Canada. Publication #1495E. Agriculture
The only disease causing significant losses to Canada, Ottawa, Canada, 53 pp
the leafcutting bee industry is chalkbrood, caused Richards KW (1987) Alfalfa leafcutter bee management in
Canada. Bee World 68:168178
by the fungus Ascosphaera aggregata Skou. The Vandenberg, JD, Stephen WP (1982) Etiology and symptoma-
disease was first reported in leafcutting bees in tology of chalkbrood in the alfalfa leafcutting bee,
1973, and remains most severe in the western U.S. Megachile rotundata. J Invertebr Pathol 39:133137
states, where losses of more than 65% of bees are
not uncommon. Bee larvae become infected after
consuming pollen provisions contaminated with Alfalfa (Lucerne) Pests and their
the fungal spores which germinate within the Management
midgut and penetrate into the hemocoel. Larvae
soon die and turn a chalk white color as the myce- john l. capinera
lium fills the body. Sporogenesis occurs beneath University of Florida, Gainesville, FL, USA
the host cuticle resulting in the formation of asco-
spores which are bound in spore balls within Alfalfa (lucerne), Medicago sativae, is one of the
ascomata. At this stage, the cadaver turns black. most important legumes used in agriculture. It is
Spores are spread by adults that must chew their the principal roughage for ruminants, as well as
A Alfalfa (Lucerne) Pests and their Management

being an important source of protein in animal climates it is irrigated. It is tolerant of cold, grow-
diets. It is surpassed only by grass, corn, and soy- ing well in cool and cold climates. It does not tol-
bean as an animal feed, and is especially important erate hot, humid climates, however.
to the dairy industry. The USA is the worlds larg- Alfalfa often is cut and dried before it is baled
est producer of alfalfa, but it also is an important and stored. To speed up the process of drying, alfalfa
crop in Australia, Europe, Argentina, China, South is commonly flailed or passed through a set of roll-
Africa, and the Middle East. ers to break or crush the stems, facilitating the dry-
There are other uses for alfalfa, though they are ing process. The crushing process is called crimping
minor. Alfalfa sprouts are a salad ingredient, alfalfa and sometimes can cause problems for horses
shoots are sometimes consumed as a leafy vegeta- because blister beetles (Coleoptera: Meloidae) are
ble, and dehydrated alfalfa is sometimes formulated incorporated into the hay (see below, blister beetles).
as a tablet to be consumed as a dietary supplement. Dried alfalfa is tied into bales of various sizes,
Alfalfa is a cross-pollinated species. It relies on including large cylindrical bales, and stored under
insects, often domesticated leafcutting bees, honey shelter, or packaged in plastic, to avoid moisture. If
bees, alkali bees, and various wild bees, for pollina- the alfalfa is to be fed to cattle, however, it is not
tion. Wind pollination does not occur because the dried, and instead it is finely chopped and stored in
blossom is structured in a way that physical trip- trenches, silos, or bags where it can ferment and
ping to expose the stigma to the anthers is required. maintain high nutrient levels. Cattle are not very
Bees manipulate the blossom when foraging for susceptible to poisoning by blister beetles.
nectar and pollen and thereby trip the blossom, an Alfalfa has undergone considerable breeding
action that results in the bee being struck in the to produce strains that have not only suitable agro-
head. An interesting aspect of pollination is that nomic conditions, but also are disease and pest
some bees learn to avoid the tripping process to resistant. Nevertheless, insects can damage alfalfa
avoid being struck, thereby robbing the flower with- nearly everywhere it is grown. Some of the impor-
out pollination occurring. Older honey bees are tant pests are listed in the table, and the most
good at avoiding tripping, but nave young honey important are discussed below.
bees trip the blossom and provide pollination.
Alfalfa is normally harvested before, or at, the
initiation of flowering, which maximizes protein Alfalfa Weevil, Hypera postica
content of the harvested hay. Because pollinators (Gyllenhal) (Coleoptera:
are often present in alfalfa fields during the bloom Curculionidae)
period, care must be taken when using insecticides
for pest suppression to avoid products that are In many regions, this is the most important pest of
highly toxic to pollinators, at least if seed produc- alfalfa. It is found in Europe, the Middle East, Cen-
tion is a concern. However, most alfalfa is grown tral Asia, and North America. Alfalfa weevils over-
only for forage, and without regard for seed pro- winter as adults in the soil of weedy, brushy areas
duction. Thus, insecticide use may include the near alfalfa fields. They disperse to alfalfa in the
bloom period, though if pollinator populations spring and oviposit within the stems. The eggs are
are reduced, other crops that require pollination oval and yellow. Early instars developing from
may be inadvertently affected. these eggs are slate colored, but develop a bright
Alfalfa is unusual as a field crop in that it is a green color and a white stripe down the middle of
short-lived perennial, living 312 years. It may be the back as they mature. Larvae have a black head
harvested from once to 12 times per year, depend- capsule. They display four instars and will grow to
ing on climate and growing conditions. It has deep about 810 mm in length. After feeding for 34
roots, and is resistant to drought, though in arid weeks, larvae spin loosely constructed cocoons on
Alfalfa (Lucerne) Pests and their Management
A 103

Alfalfa (Lucerne) Pests and their Management, Table 5 Some pests of alfalfa (lucerne), and locations
where they are considered to be damaging
Feeding behavior Primary taxon Common name Scientific name Location
Above-ground, Coleoptera Sitona weevil Sitona discoides Australia
Small lucerne Atrichonotus Australia
weevil taeniatulus
Vegetable weevil Listroderes Australia
Alfalfa weevil Hypera postica Europe, Asia,
Clover leaf weevil Hypera punctata Europe, Asia,
Clover head weevil Hypera meles Europe, N. America
Blister beetles Epicauta spp. N. America
Flea beetles Epitrix, Systena, N. America
Disonycha spp.
Orthoptera Grasshoppers Melanoplus spp. N. America
Wingless Phaulacridium Australia
grasshopper spp.
Lepidoptera Armyworm Mythimna spp., Australia
Armyworm Persectania spp. Australia
Armyworm Pseudaletia N. America
Variegated Peridroma saucia Europe, Asia, Africa,
cutworm N.America
Army cutworm Euxoa auxiliaris N. America
Granulate cutworm Agrotis N. America, S.America
Black cutworm Agrotis ipsilon N. America, Europe,
Beet armyworm Spodoptera Asia, N. America
Fall armyworm Spodoptera N. America, S.America
Budworm Helicoverpa Australia
Corn earworm Helicoverpa zea N. America, S.America
Alfalfa looper Autographa N. America
A Alfalfa (Lucerne) Pests and their Management

Alfalfa (Lucerne) Pests and their Management, Table 5 Some pests of alfalfa (lucerne), and locations
where they are considered to be damaging (Continued)
Feeding behavior Primary taxon Common name Scientific name Location
Lucerne leafroller Merophyas Australia
Alfalfa caterpillar Colias eurytheme N. America
Webworms Loxostege spp. N. America, Europe,
Collembola Lucerne flea Sminthurus Australia, Europe,
viridis Africa
Diptera Alfalfa blotch Agromyza Europe, N. America
leafminer frontella
Above-ground, Acari Redlegged earth Halotydeus Australia
sucking mite destructor
Clover mite Bryobia spp. Australia
Twospotted spider Tetranychus No. America
mite urticae
Collembola Lucerne flea Sminthurus Europe, N. Africa,
viridis Australia
Hemiptera Pea aphid Acythosiphum Europe, Asia, Australia,
pisum N. & S. America
Blue alfalfa aphid Acythosiphum Mediterranean,
kondoi Australia, N. & S.
Spotted alfalfa Therioaphis Mediterranean,
aphid maculata Australia, N. America,
Potato leafhopper Empoasca fabae N. America
Lucerne leafhopper Austroasca Australia
3-cornered alfalfa Spissistilus N. America
hopper festinus
Meadow spittlebug Philaneus N. America
Tarnished plant Lygus spp. Europe, N. America
Alfalfa plant bug Adelphocoris Europe, N. America
Thysanoptera Flower thrips Frankliniella spp. Europe, Asia,
Below-ground Coleoptera Clover root Sitona hispidula Europe, N. & C.
curculio America
Alfalfa (Lucerne) Pests and their Management
A 105

Alfalfa (Lucerne) Pests and their Management, Table 5 Some pests of alfalfa (lucerne), and locations
where they are considered to be damaging (Continued)
Feeding behavior Primary taxon Common name Scientific name Location
Alfalfa snout beetle Otiorhynchus N. America
African black Heteronychus Africa, Australia
beetle arator
Whitefringed Naupactus Australia, S. America
beetle leucoloma
Small lucerne Atrichonotus Australia
weevil taeniatulus

plants or in litter on the soil, pupate, and emerge as aethiopoides Loan and M. colesi Drea (Hymenoptera:
adults in 12 weeks. Adults are 56 mm long, have Braconidae) Oomyzus incertus (Ratzenberg)
a long snout, and have a dark stripe down the back. (Hymenoptera: Eulophidae); Dibrachoides dynas-
They are light brown at emergence and darken in tes (Forester) and Peridesmia discus (Walker)
several days. The number of generations varies (Hymenoptera: Pteromalidae); and Anaphes luna
according to climate, but eventually they leave (Girault) (Hymenoptera: Mymaridae). A fungal
fields for grassy, brushy, weedy areas where they pathogen, Zoophthora phytonomi Arthur (Phyco-
become inactive until the onset of winter. mycetes: Entomophthoraceae), attacks weevil lar-
Damage is caused by the larval stage which vae and can control populations in several days,
feeds on leaves; damage ranges from pinholes to though it is most effective under moist conditions.
skeletonization of leaves. Adults generally cause These biological control agents are extremely effec-
minor damage. Peak damage is usually just prior tive control measures in all but major outbreak
to the first cutting or after the first cutting, as both periods. However, when fields show damage on
larvae and adults feed on new growth; this can 3540% of plant tips more than 710 days prior to
seriously affect regrowth of the stand. Also, cool, harvest, chemical suppression is often initiated.
cloudy weather exacerbates damage done by the Early harvest (first crop) is very effective in
alfalfa weevil. Cool and cloudy weather conditions killing larvae, and is preferred to chemical control if
slow the regrowth rate of alfalfa, and also increase the planned harvest is less than 710 days away. If
the daily feeding period of the weevil because both harvesting is used to control alfalfa weevil, the stub-
larvae and adults tend to hide under crop residue ble and debris should be examined closely for adults
during bright sunlight and will not actively feed and larvae, and stems should be examined for feed-
during such periods. ing signs. It may be necessary to spray stubble,
Weevil larvae can be found early in the Spring. though in many areas producers can avoid insecti-
It is important to scout for live larvae and injured cide use consistently through timing of harvest.
terminals on the first crop, but also subsequent
crops. Sweep net sampling can be used to detect
weevil presence. Several species of wasps can be Root Weevils, Sitona spp.,
effective in maintaining weevil populations below Atrichonotus taeniatulus (Berg),
economic threshold levels. Among the effective Others (Coleoptera: Curculionidae)
parasitoids are Bathyplectes curculionis (Thomson),
B. anurus (Thomson) and B. stenostigma (Thomson) Root weevils such as clover root curculio, Sitona his-
(Hymenoptera: Ichneumonidae); Microctonus pidula (Fabricius), in North America; and sitona
A Alfalfa (Lucerne) Pests and their Management

weevil, Sitona discoideus Gyllenhal, small lucerne c ontain crushed blister beetles that may prove toxic
weevil, Atrichonotus taeniatulus (Berg), in Australia, to horses that ingest the hay. The most abundant
and whitefringed beetle, Naupactus leucoloma Bohe- blister beetle in alfalfa fields generally is the black
man in Australia and South America, can be signifi- blister beetle, E. pensylvanica (De Geer). However,
cant pests of alfalfa. Although the adults commonly the species that is most toxic is E. vittata (Fabricius).
feed on the foliage, the principal damage is due to Most blister beetles are recognized by the
larval feeding on the roots of the alfalfa plant. shape of their body. They are narrow, cylindrical,
Eggs are laid in fall or spring, on the soil sur- and soft. The region between the head and wings is
face or lower parts of plants. Eggs hatch in the distinctly narrower than the wings, and is usually
winter or spring. White, legless larvae move into narrower than the head. Most species have one
the soil and feed on roots until they pupate. Pupae generation per year, although some have two.
are found just below the soil surface. Adults emerge Blister beetles overwinter as larvae. The adults
in the summer months and live up to a year. The begin to emerge in the Spring and adults deposit
adults are brown or black, blunt-snouted weevils their eggs where grasshopper egg pods may occur,
up to about 10 mm long. There is one generation as larvae feed on the grasshopper eggs.
per year. Adults migrate by crawling, and thus If grasshoppers are not abundant, then blister
infest new areas rather slowly. beetles are unlikely to be abundant. When both
The adults feed on alfalfa leaf margins, leaving are numerous, it is advisable to harvest alfalfa
crescent-shaped notches, and chew on stems and early, before bloom, as this is the only time that
leaf buds of seedlings, but this tends to cause beetles are attracted to the crop. There is some
minor loss. Most damage is caused by the larvae. yield loss associated with this approach, of course,
First larval instars feed on root nodules and lateral and an alternative it to treat the crop with insecti-
roots; later instars feed on the taproot. Feeding on cides. If insecticide is used, alfalfa should be har-
the taproot can girdle the plant, resulting in plant vested as soon as possible after the pre-harvest
death. Such damage also weakens the overall vigor interval expires, to get hay out of the field before it
of a stand, perhaps contributing to winter-kill and is re-infested. A principal problem with blister
increased susceptibility to disease. beetle management is that the beetles tend to
It is difficult to control larvae because they are aggregate. Thus, there may be relatively few bee-
in the soil and largely protected from insecticide. tles in field, but a large number in one location,
Suppression aimed at adults usually requires mul- and these may be crushed together and concen-
tiple applications. It is inadvisable to plant alfalfa trated into one or a few bales of hay. Thus, they are
into a field which has previous ly been infested, to hard to detect by standard sampling methods.
plant into fields previously supporting legume When alfalfa hay is purchased for horses, it is
crops, or to seed alfalfa next to established stands. advisable to acquire early-crop hay, or hay from
areas free of high grasshopper populations. Alter-
natively, inspection of the hay as it is fed to horses
Blister Beetles (Epicauta spp.) can reveal the presence or absence of beetles.
(Coleoptera: Meloidae)

There are several species of North American blister Potato Leafhopper, Empoasca
beetles that can be of concern in alfalfa. They are a fabae (Harris) (Hemiptera:
problem not because of their food habits (they tend Cicadellidae)
to feed mostly on blossoms) but because they con-
tain the toxin cantharidin within their bodies. When Potato leafhopper is indigenous to eastern
alfalfa is harvested, if the hay is crimped it may North America. Adults are about 3.5 mm long,
Alfalfa (Lucerne) Pests and their Management
A 107

wedge-shaped, winged, and green. Nymphs are Although the potato leafhopper has natural
similar in appearance, but are smaller, yellowish- enemies, they often get left behind when the adults
green to fluorescent green, and wingless. Each disperse. Thus, a combination of crop monitoring
Spring, potato leafhoppers migrate north from and insecticide suppression is often the principal
southern states where they overwinter. Timing of management strategy. Chemical control of potato
the first and subsequent arrivals in the north is leafhopper is effective, but should not be used if
heavily dependent on weather patterns. Adults lay harvest is within seven days of harvest. Cutting
eggs in stems and leaf veins; eggs hatch in 69 days will kill a large percentage of nymphs, and will
in mid-summer. Each generation takes approxi- force adults out of the field. Cutting is the control
mately 3035 days to mature, resulting in several of choice if thresholds are reached within seven
generations. days of harvest. Additionally, early harvest may be
Adults and nymphs both feed on alfalfa with an alternative to insecticides when thresholds are
piercing-sucking mouthparts, sucking plant sap reached late in the year.
and injecting a toxin into the plant. Damage is
called hopperburn, and is a yellow wedge-shape
area beginning at the tips of leaves. The leaves may Aphids (Hemiptera: Aphididae)
eventually turn entirely yellow or reddish. Plants
may become stunted. Leafhoppers cause yield loss, Several aphids are pests of alfalfa, including pea
reduced nutritional quality of alfalfa, and reduced aphid, Acyrthosiphon pisum (Harris); blue alfalfa
plant vigor that results in increased winter-kill and aphid or bluegreen aphid, Acyrthosiphon kondoi
slower regrowth of the crop the next spring. Shinji, cowpea aphid, Aphis craccivora Koch; green
In some regions of the USA, the potato leafhop- peach aphid, Myzus persicae Sulzer; and spotted
per is the worst insect pest of alfalfa, and can cause alfalfa aphid, Therioaphis maculata Buckton.
losses of 80% or more if not controlled. Leafhoppers All these aphids are small, measuring 3 mm
are not generally a problem in the first crop in an or less. Their color varies, depending on species.
established stand, but as the population increases, all They may or may not be winged. In most climates,
subsequent crops will need to be monitored for in early Spring nymphs hatch from eggs that were
infestation. The characteristic hopperburn will not laid in the fall; these aphids are all female. Females
appear until some yield and quality loss has occurred, can reproduce without mating when conditions
so it is important to scout for leafhoppers weekly are favorable, and they do so in Spring and Sum-
onthe second and subsequent crops. Scouting may mer. In the Summer, the entire life cycle takes
be concluded 710 days prior to harvest. only a few days. Males appear in late Summer,
Potato leafhopper economic thresholds are and mate with females to produce eggs capable of
based on plant height. Scouting is accomplished overwintering.
by sweep net sampling. As an example, following Aphids use piercing-sucking mouthparts to
are treatment thresholds recommended for remove plant sap, and prefer to feed on young
Minnesota, USA. growth. Aphid feeding can result in stunted or
wilted plants. The plants may also turn yellow.
Average plant height # adult leafhoppers/ Aphids commonly attain high densities in
sweep alfalfa, but in most years natural enemies keep
< 3 inches 0.3 aphid populations at levels that are not economi-
37 inches 0.5
cally important. Many natural enemies of pea
aphids exist, including green lacewing larvae
812 inches 1.0
(Neuroptera: Chrysopidae), damsel bugs
> 12 inches 2.0
(Hemiptera: Nabidae), and parasitic wasps
A Alfalfa (Lucerne) Pests and their Management

(Hymenoptera, various families), lady beetles Most egg laying occurs in late summer and fall in
(Coleoptera: Coccinellidae), and disease (fungi). production areas; most species prefer uncultivated,
grassy or weedy areas, and lay eggs 13 cm below
the soil surface.
Plant Bugs (Hemiptera: Miridae) Grasshoppers are generally considered a
minor pest except during periods of great abun-
Several species of plant bugs affect alfalfa, but the dance, and then they can do great damage. An
most common are tarnished plant bug, Lygus line- exception is Australia, where wingless grasshop-
olaris (Palisot de Beauvois), and alfalfa plant bug, per has become an increasingly severe pest of
Adelphocoris lineolatus (Goeze). Adult tarnished alfalfa. Damage has increased in Australia due
plant bugs are brown, winged, and 46 mm long; to widespread cultivation of alfalfa, which is
nymphs are green, wingless, and the third and more suitable than grasses for nymphal growth
subsequent instars have black spots. Adult alfalfa and survival. Grasshopper nymphs and adults
plant bugs are light green, winged, and 7.510 mm damage alfalfa by chewing on leaves from the
long; nymphs are green, wingless, and have red margin inward in an irregular pattern. Attacks
eyes. Tarnished plant bugs overwinter as adults; are often on new growth, but will occur on any
alfalfa plant bugs overwinter as eggs in plant tis- stage. The margins of fields are most likely to be
sue. During the growing season, the entire life damaged.
cycle takes 2050 days, depending on temperature. In North America, grasshopper infestations
There are two to five generations per year. are more severe in warm and dry years. Warm,
Plant bugs suck sap from plants and inject dry weather immediately following egg hatch
toxic saliva into the plant. They cause leaves to favors survival of nymphs, because nymphal
crinkle, plants to be stunted, and flower buds to growth rates and survival are lower in cool, wet
abort. They are abundant in all but the earliest weather. Long, warm autumns prolong the egg-
portions of the season. Although traditionally laying season, and result in larger populations in
considered mostly a seed pest, plant bugs also the next growing season. It can take 35 years for
contribute to forage yield reductions. If bugs are populations to build to economically important
abundant more than seven days prior harvest, levels. In Australia, drought also is implicated,
chemical control may be warranted. but mermithid nematodes are a critical element
in grasshopper biology. Absence of rainfall, and
clearing of drier, higher elevation pasture impedes
Grasshoppers (Melanoplus spp. the ability of the nematodes to parasitize the
and Phaulacridium spp.) grasshoppers.
(Orthoptera: Acrididae) Grasshoppers are naturally suppressed by
numerous natural enemies, but when weather con-
Everywhere alfalfa is grown, grasshoppers and ditions favor the grasshoppers their populations
locusts will feed on the crop. However, they are increase quickly. The natural enemy population
only casually associated with alfalfa, attacking the increases as their food supply becomes more avail-
crop only when abundant. None feed preferentially able, but the lag in natural enemy abundance can
on alfalfa. In North America, the principal pests are result in crop damage by the grasshoppers. Weedy
Melanoplus spp., and in Australia Phaulacridium fence rows, irrigation ditches, and fallow fields are
spp. is the major grasshopper pest. These economi- important sources of grasshoppers. Weed popula-
cally important grasshoppers overwinter as eggs. tions should be managed, which may require tillage
Populations disperse into cultivated fields or pas- or burning to make these habitats less productive
tures as their populations build through the season. for grasshoppers.
Alfalfa (Lucerne) Pests and their Management
A 109

Cutworms, Armyworms and ability and its minute size, not because it is a flea
Budworms (Lepidoptera: or related to fleas. This species has a patchy dis-
Noctuidae) tribution in Europe and North Africa, and has
been accidentally introduced to Australia, where
The caterpillars of several moths can become it is most injurious. It also affects lupine flowers,
abundant enough to cause significant loss to lentils, beans, and field peas. Immature lucerne
alfalfa.Among these are the armyworms Mythimna fleas consume small patches of foliage, whereas
spp., Persectania spp., and Pseudaletia unipuncta adults consume the entire leaf except for the
Haworth; variegated cutworm, Peridroma saucia veins. Early season spraying of insecticide is the
(Hbner); army cutworm, Euxoa auxiliaris most common recommendation to curb their
(Grote); granulate cutworm, Agrotis subterranea damage.
(Fabricius); beet armyworm, Spodoptera exigua
(Hbner); budworm, Helicoverpa punctigera
(Wallengren), and many others. The important Mites (Acari)
species vary among regions, though they are simi-
lar ecologically. Mites generally are not major pests of alfalfa, but
The larvae of cutworms, armyworms, and under arid conditions or along the margins of
budworms range in color from greenish-yellow to fields they can be quite damaging. The most impor-
brownish-black. Larvae are 25 cm long at matu- tant are clover mites, Bryobia spp. (Acari: Tetrany-
rity. The wings of the adults vary from tan to dark chidae), and redlegged earth mite, Halotydeus
brown with mottling or stripes. Pupae are 13 cm destructor Tucker (Acari: Penthaleidae), in Austra-
long and are reddish-brown to black in color. lia, and twospotted spider mite, Tetranychus urticae
There are one to six generations per year. Larvae Koch (Acari: Tetranychidae) in North America.
overwinter in the larval or pupal stage, depending They rupture the cells of leaf tissue, imparting a sil-
on species. ver or yellow appearance, and reducing yield.
Larvae feed on stems and leaves of plants, and
can limit regrowth after harvest. Larvae will also
cut the stems of seedlings. Their occurrence as Pest Management in Alfalfa
economic pests is sporadic. Although these insects
have many natural enemies, when they are abun- Alfalfa is an excellent crop for the practice of mod-
dant insecticides are the preferred approach to ern pest management tactics because (i) it is quite
population reduction. tolerant of damage; cosmetic injury is not impor-
tant; (ii) it is a perennial crop, providing harbor-
age throughout the year for an immense assemblage
Lucerne Flea, Sminthurus viridis of insects, including predators and parasitoids;
(Collembola) (iii) it is an important crop, so extensive research
on the pests have been conducted; (iv) it is amena-
Sminthurus viridis, the lucerne flea or clover ble to various cultural manipulations, and pro-
springtail, is an insect relative (hexapod) belong- duces multiple crops over a large portion of the
ing to the order Collembola (the springtails). It year; and (v) it is a favorite crop for rotations, so
is bright green with a roughly spherical body there is ample opportunity to integrate its culture
and may swarms in large numbers on living with the culture of other crops.
plants, including alfalfa or lucerne, thus the first The principal tactics used for alfalfa produc-
part of the common name. The second part of tion include scouting and use of an economic
the common name was given for its jumping threshold for decision making, natural and classical
A Alfalfa Weevil, Hypera Postica (Gyllenhal) (Coleoptera: Curculionidae)

biological control, cultural control, and chemical populations. Strip cropping is commonly recom-
control. The economic threshold varies among mended because the uncut areas retain popula-
insect species, geographic locations, crop manage- tions of natural enemies, allowing the beneficial
ment practices and economic conditions, but most insect to move into newly harvested alfalfa as it
locations have established such benchmarks for regrows and becomes infested with pests. Farmers
initiating chemical control. A large number of rarely embrace this approach, however, opting for
insecticides are registered for this crop, so growers operational efficiency over economic pest control.
have ample opportunity to select products accord-
ing to their need and budget. A modest level of
host plant resistance apparently exists in alfalfa, References
and although resistance is effective mostly against
aphids, there is also some success with alfalfa wee- University of California (1981) Integrated pest management
vil and leafhoppers. for alfalfa hay. Publication #3312, 98 pages
A large number of beneficial arthropods have Summers CG (1998) Integrated pest management in forage
alfalfa. Integr Pest Manage Rev 3:127154
been moved around the world in an effort to attain
biological suppression of invading alfalfa-feeding
insects. In some cases this has met with success. For
example, alfalfa blotch leafminer, Agromyza fron- Alfalfa Weevil, Hypera postica
tella (Diptera: Agromyzidae), was considered a seri- (Gyllenhal) (Coleoptera:
ous pest when it first invaded the eastern USA, but Curculionidae)
following release of wasp parasitoids it fell to minor
pest status. Similarly, the status of spotted alfalfa An important defoliator of alfalfa (lucerne).
aphid, pea aphid and blue alfalfa aphid was affected Alfalfa (Lucerne) Pests and their Management
by importation of beneficial insects. A native ento-
mopathogenic fungus, Zoophthora phytonomi, has
adapted to the invasive alfalfa weevil and sometimes Alga (pl., Algae)
provides good suppression. Pea aphid is affected by
the fungus Erynia neoaphidis under favorable An aquatic non vascular plant, often very small in
weather conditions. Generalist predators such as size. Algae can reach pest status when weather and
lacewings, lady bird beetles, nabids, soft-winged nutrient levels favor its growth, and pesticides may
flower beetles, big-eyed bugs, and minute pirate be needed to suppress it.
bugs are often active in alfalfa, and provide good
suppression of aphids, thrips, and also consume Alien
eggs and young larvae of caterpillars.
Cultural manipulations are the most impor- An organism that is native elsewhere. These are
tant tactics for management of alfalfa pests. In also referred to as exotic or foreign.
particular, early harvesting can provide acceptable Invasive Species
or even nearly complete control of alfalfa weevil,
alfalfa blotch leafminer, several caterpillars, aphids,
and leafhoppers because when the crop is cut the Alienicolae
insects are exposed to lethal levels of heat and dry-
ness, or the environment becomes so unsuitable In heteroecious aphids, viviparous parthenoge-
that the insects move elsewhere. Crop rotation is netic females developing on herbaceous (second-
most important for root feeding pests, many of ary) host plants.
which take several years to develop damaging Aphids
Alimentary Canal and Digestion
A 111

Alimentary Canal and Digestion

james l. nation
University of Florida, Gainesville, FL, USA

Insects feed upon many different kinds of food,

including paper, wood, plant phloem and xylem
sap, plant leaves, roots and stems, animal tissues,
hair, wool, and vertebrate blood. The alimentary
canal (often simply called the gut in much of the
literature) evolved to accommodate such diverse
foods in a variety of morphological and physio-
logical ways. Thus, there is no typical insect ali-
mentary canal just as there is no typical insect.
Nevertheless, there are similarities in the structure
of the alimentary canal in all insects andnearly all
must digest some of the same complex molecules,
such as proteins, lipids, and carbohydrates.
In every insect alimentary canal three regions
can be identified morphologically and physiolog-
Alimentary Canal and Digestion, Figure 33 A
ically: the foregut or stomadeum, the midgut or
generalized drawing of the alimentary canal
mesenteron, and the hindgut or proctodeum
in a cockroach to show the major divisions of
(Fig.33). One or more of these regions may be
the c anal. Many variations occur in the overall
greatly reduced in size, or expanded in size,
structure of the alimentary canal in insects, and
depending upon the feeding behavior of the
this is not i ntended to suggest that the cockroach
insect. A cuticular layer, the intima, attached to
alimentary canal is typical of insects.
the epithelial cells, lines the fore- and hindgut
regions. The old intima is partially digested and with the food, lubricate the food, and may begin
the residue sloughed off into the gut and excreted some carbohydrate digestion. In some insects the
at each molt, and a new intima is secreted. The crop is not a noticeably modified part of the
midgut does not have anattached cuticular lin- foregut, but often the crop comes off the foregut as
ing, but may have a non-attached peritrophic a diverticulum. In other insects it is an enlarged
membrane that separates the food enclosed portion of the foregut. In opportunistic and pos-
within from the delicate surface of the midgut sibly irregular feeders such as praying mantids, the
cells. If a peritrophic membrane is present, it is crop composes more than half of the alimentary
often secreted several times each day. canal, apparently an evolutionary development to
store a large amount of food when available and
tide the mantid over periods when prey is scarce.
The Foregut Some insects (for example, many orthopterans)
regurgitate enzymes from the midgut into the
The buccal cavity (mouth), pharynx, esophagus, crop, and these enzymes, along with salivary secre-
crop, proventriculus and attached salivary glands tions, digest food in the crop. The digested food
comprise the foregut. Secretions from the salivary components still enter the midgut to be absorbed,
glands attached near the mouth are swallowed and there is no evidence that the crop ever secretes
A Alimentary Canal and Digestion

enzymes itself. The cuticular intima creates a bar- in nidi (nests). The regenerative cells grow into
rier even against the absorption of water from the mature epithelial cells to replace cells worn out or
crop. The proventriculus controls the entry of food those that disintegrate to release digestive
into the midgut in liquid feeders, but in many enzymes. Midgut cells may be completely replaced
insects it is modified into a grinding apparatus every few days in insects that live longer lives.
with hard, sclerotized ridges and spines, and heavy Gastric caeca, small finger or sac-like diverticula
musculature for breaking and tearing the food from the midgut, often arise at or near the origin
into smaller particles. of the midgut, but may be located at various points
along the midgut. The caeca appear to secrete
digestive enzymes and may be important in
The Midgut absorption of digested products.
The midgut does not have an attached cuticu-
The midgut in most, but not all, insects is the main lar lining on the surface of the cells, but midgut
site for digestion, absorption, and secretion of cells in the majority of insects secrete a thin mem-
digestive enzymes. The epithelium is a single layer brane composed of chitin and protein, the peri-
of cells, but several types of cells occur in some trophic membrane, that surrounds the food and
insects. The most common cells are tall, relatively shields the delicate microvilli of the midgut cells
narrow ones called columnar or primary cells by from contact with potentially rough and abrasive
various authors. They have extensive microscopic food particles. Although the peritrophic mem-
microvilli on the apical or lumen surface (Fig.34) brane is thin, varying from 0.13 m to about 0.4
and extensive invaginations of the basal cell mem- m thick, it also is thought to make it more diffi-
brane, features that greatly increase the surface cult for viruses, fungi, bacteria, and protozoans to
area on both sides of the cell over which secretion get to the surface of the midgut cells where they
and absorption occur. The columnar cells are the might be able to enter the cells and create an infec-
primary cells that secrete digestive enzymes and tion. Some insects produce several peritrophic
absorb digested products. In all insects that feed membranes per day, each encasing the one before
as adults and live for days or weeks, there are small it, perhaps increasing protection from random
regenerative cells distributed at the base of the breaks or punctures by larger food particles, and
columnar cells, or sometimes clustered together thus affording more protection for the midgut

Alimentary Canal and Digestion, Figure 34 A brush border of microvilli on the lumen surface of midgut
cells in a mole cricket.
Alimentary Canal and Digestion
A 113

cells from possible fungi, parasites, viruses, and the epithelium cells and the gut lumen and seem to
bacteria ingested with the food. A peritrophic play a major role in amino acid absorption. The
membrane occurs in living representatives of some high midgut pH may provide plant feeding insects
of the earliest insects to evolve, and it is believed to some protection against tannins that are common
have evolved very early in a generalist scavenger in the food plants of phytophagous insects. Tan-
feeder in which protection of midgut microvillar nins can complex with an insects own enzymes
surfaces from food particles, sand, or other hard and proteins in the food, and may result in reduced
substances coincidentally ingested was likely to be digestion and absorption. Many details and the
important. A peritrophic membrane is present in precise metabolic components in the cell mem-
many insects that do not feed upon rough or solid brane that support and enable these secretory and
food, such as some blood feeders (but not all blood absorptive mechanisms remain to be elucidated,
feeders), and in adult lepidopterans that take but what is already known emphasizes the com-
flower and plant nectars. Although the peritrophic plexity of insect digestive functions.
membrane is not present in all groups of insects,
like other gut features, it has been conserved over
long evolutionary time, lending support to views The Hindgut
that it has multiple functions, especially protec-
tion from disease invaders and may even have The hindgut is not only a posterior extension of
properties that could bind toxicants and limit their the alimentary canal, but it also plays a major role
access to cells. in excretion through secretion of some substance
Absorption of digested food substances has into the lumen, and reabsorption of useful sub-
not been studied in most insects, but one mecha- stances such as ions, water and some nutrients
nism has been partially elucidated for absorption from the Malpighian tubule effluent. The Mal-
of amino acids derived from protein digestion in pighian tubules typically arise at the origin of the
larvae of Lepidoptera. Interspersed among the tall hindgut (but exceptions do occur) and pass rela-
columnar cells lining the midgut in larvae of Lepi- tively large volumes of an ultrafiltrate of hemo-
doptera (and some other groups of insects as well) lymph components minus proteins into the
are cells shaped much like a goblet and called, beginning of the hindgut. The cuticular lining on
appropriately enough, goblet cells. The apical cell hindgut cells is thinner and has larger pores than
membrane of the goblet cavity has metabolic the lining in the foregut, permitting reabsorption
machinery that uses energy derived from splitting of water, some ions, and useful metabolites that
ATP to push or pump protons (H+) into the goblet are returned to the hemolymph. In most terres-
cavity. A different set of machinery in the goblet cell trial insects, water conservation is vital to life, and
membrane, an antiporter mechanism, reabsorbs the hindgut must conserve the water that the Mal-
the protons and simultaneously secretes potassium pighian tubules flush into the hindgut. Waste
ions into the goblet cavity. The net result of the products such as undigested food material (cel-
secretion of potassium ions is that a strongly alka- lulose, for example, which most plant-feeding
line midgut (a midgut pH as high as pH 8 to about insects cannot digest and use), uric acid, and other
11) is produced, and a high voltage (up to 240 mV allelochemicals picked up from the food are con-
in some reports) is created between the gut lumen centrated in the rectum and eventually excreted.
(positive) and the interior of cells lining the gut. The hindgut secretes some molecules into the
The voltage created by the pump enables an absorp- lumen for excretion. Experimental evidence indi-
tive mechanism in membranes of columnar cells to cates that secretion and selective reabsorption
reabsorb K+ and amino acids from protein diges- helps regulate pH of the hemolymph in some
tion. Thus, potassium ions are recycled between insects. Specialized cells, the rectal papillae and
A Alimentary Canal and Digestion

rectal pad cells, in the rectum of many insects of the nutrient value of the food source. Some
have characteristic ultrastructure and physiologi- insects reflux enzyme secretions and partially
cal mechanisms typical of highly reabsorptive digested products by repeatedly sucking up and
cells. Water conservation by the rectum results in reinjecting the liquefied juices into the food.
the relatively dry frass or fecal pellets characteris- Refluxing mixes the secretions and fluids and
tic of many terrestrial insects. extends the effective life of the digestive enzymes,
The highest degree of specialization in the and is particularly effective when the food con-
hindgut occurs in those insects that digest cellu- tains a limiting boundary, such as the shell of a
lose, such as termites. In termites, the hindgut is seed or the cuticle of an insect that acts as a con-
usually divided into several chambers harboring tainer for the liquefying body contents.
either bacteria or protozoa that secrete all or part
of the cocktail of enzymes needed to digest cellu-
lose. Glucose, liberated from cellulose digestion, Carbohydrate Digestion
may be fermented by the resident microorgan-
isms, with the end products being short chain Starch and sucrose are the typical carbohydrates
fatty acids (principally acetic acid) that can be that insects digest from plant food, and glycogen
absorbed by the termite and used as an energy and various sugars are present in animal tissue
source. Some termites release methane, a green- eaten by carnivorous insects. Cellulose, the major
house gas, from the metabolic activities of their complex polysaccharide present in plant tissue,
microorganisms, but whether this is a significant cannot be digested by most insects. Carbohydrate
natural source of methane is a topic of debate by digestion begins with the action of -amylase, an
various scientists. enzyme present in the salivary gland secretions of
many insects. Amylase works best at slightly acid
pH, and hydrolyzes interior glucosidic linkages
Digestive Enzyme Secretion ofstarch and glycogen, resulting in a mixture of
shorter dextrins. In the midgut -glucosidase and
Midgut cells secrete and release digestive enzymes oligo-1, 6-glucosidase (isomaltase) digest smaller
in several ways. They may enclose digestive dextrins, releasing glucose. Many insects also have
enzymes in small vesicles surrounded by a mem- one or more - or -glycosidases that digest
brane and then release the enzymes into the ali- a broad range of small carbohydrates, such as
mentary canal by fusing the vesicle membrane maltose, sucrose, trehalose, melezitose, raffinose,
with the cell membrane. In some insects, parts of stachyose, melibiose, raffinose, and stachyose.
the cell (some of the microvilli) or the entire Some insects can secrete trehalase in the gut to
midgut cell may disintegrate and release enzymes digest trehalose, the principal blood sugar typi-
into the gut lumen. Of course, when the entire cell cally in high concentration in insects. -glucosidase,
breaks down, the cell must be repaired or replaced. -galactosidase, and -fructofuranosidase act
Replacement occurs through the growth of the upon various substrates to release simple sugars
regenerative cells. in the gut. An insect usually has only a few of
Extraoral digestion (digestion outside the these carbohydrate digesting enzymes, depending
insect body) occurs in some insects, including upon the food it eats. For example, Apis mellifera
seed feeders and some predatory insects. By inject- honeybees have several -glucosidases or sucrases
ing enzymes from the salivary glands and midgut that act rapidly upon sucrose, usually the princi-
into the food source (animal or plant material) pal carbohydrate in the nectar taken by honey-
and then sucking back the liquefied digestion bees. They utilize the resulting glucose and
products, insects can utilize very high percentages fructose for an immediate energy source and for
Alimentary Canal and Digestion
A 115

making honey. Termites, some beetles, a few cock- including insects, get their amino acids from
roaches, and woodwasps in the family Siricidae digestion of dietary proteins. Within insects as a
digest cellulose with aid (usually) from fungi, bac- group, there are several different types of protein
teria, or protozoa, which produce some or all of digesting enzymes, some of which act at acid pH,
the complement of three enzymes necessary for others at slightly alkaline pH, and some at highly
cellulose digestion. alkaline pH. Usually a particular species will have
several different proteinases, but no insect is
known to have both an acid-effective proteinase
Lipid Digestion and an alkaline-effective proteinase. The pH of
the alimentary canal is important to the action
The major storage forms of lipids (fats) in both of any digestive enzyme, and no insect is known
plants and insects are triacylglycerols, esters of to have an alimentary canal that is strongly acid
fatty acids with glycerol. Midgut cells, and in some in one part and strongly alkaline in another
cases symbionts, secrete lipases, which are enzymes part.
that hydrolyze triacylglycerols and release fatty Proteinases are classified broadly as serine,
acids and glycerol. Amino acids, proteins, and fatty cysteine, aspartic acid, and metallo-proteinases
acylamino complexes act as emulsifiers in the depending upon the amino acid or metal at the
midgut of some insects facilitating the digestion of active site of the enzyme. Trypsin and chymotrypsin
fats. The glycocalyx layer, a viscous protein and are two endoproteinases with alkaline pH optima
carbohydrate complex that often lies on the sur- (about pH 8) that are common in many insects and
face of the microvilli, probably aid in emulsifying which attack large proteins internally at the linkage
fats and in promoting contact between lipases and between certain amino acid, thus breaking the pro-
triacylglycerols. Fatty acids released from triacylg- tein into smaller polypeptides. Most insects appear
lycerols are resynthesized into the insects own to have several types of exopeptidases that remove
triacylglycerols and stored in fat body cells. Imma- the terminal amino acid from a protein or peptide
ture insects typically store relatively large amounts chain. Thus, through the concerted action of both
of triacylglycerols, and use some of the released types of digestive enzymes, a protein can be com-
fatty acids during pupation and for egg develop- pletely digested with release of its component amino
ment. Some insects, for example Orthoptera, Lepi- acids. Cysteine- and aspartic acid-proteinases have
doptera, and some aphids, mobilized fatty acids mildly acid pH optima, and are called cathepsins by
rapidly enough to use fatty acid metabolism to some authors. All members of a taxonomic group
support flight, but other groups such as Diptera may not have the same type of proteinases. Many
and Hymenoptera cannot release fatty acids from beetles have cysteine proteinases most active at
the fat body and transport them to the flight mus- slightly acid pH, while some scarabeid beetles
cles rapidly, and so they only use carbohydrates for secrete serine-proteinases that act at the high
flight energy. They still can use lipids during pupa- midgut pH typical of these insects, and they have
tion, and for other metabolic processes that occur no detectable cysteine-proteinases. Lepidoptera
more slowly. typically secrete trypsin-like enzymes active at
alkaline pH.
One defense mechanism that has evolved
Protein Digestion against herbivory in many plants is the presence of
proteinase inhibitors, some of which inhibit serine
All animals must have a pool of amino acids proteinases while others act upon cysteine pro-
available for synthesis into proteins, and for teinases. Experimentally, it has been shown that
repair of tissues and organs. Most animals, some insects secrete multiple trypsin enzymes
A Alimentary Canal and Digestion

(isozymes of trypsin) and others just secrete larger Gut pH

amounts of the same few isozymes after consum-
ing a trypsin inhibitor. This counter action by the The pH of the alimentary canal is highly variable
insect probably allows some protein digestion to in different species of insects. The pH of a gut seg-
escape ingested inhibitors, but transgenic plants ment influences the action of enzymes secreted
designed to have proteinase inhibitors have been into or carried with the food into the gut, influ-
tested and proven to have adverse effects upon the ences solubility and toxicity of toxins and plant
growth of some insects. allelochemicals, and may alter the population of
gut microorganisms. In most insects, the crop has
little or no presence of buffering agents and tends
Absorption of Digested Products to be slightly acidic, a factor favoring carbohydrate
digesting enzymes. Larvae of Lepidoptera and Tri-
Few details are known about the absorption of choptera tend to have a very high midgut pH,
digested products by insects. In many verte- varying from about 8 to 10, promoted by goblet
brates, glucose absorption from the alimentary cells that secrete potassium and bicarbonate into
canal requires an active mechanism involving the lumen of the midgut. They have protein digest-
ATP to supply the energy. In those insect that ing enzymes that are favored by the high pH, and
have been studied, glucose from digestion of car- the high pH may afford some protection from tan-
bohydrates is rapidly absorbed passively by a pro- nins and other allelochemicals that they ingest
cess known as facilitated diffusion, and with their plant food.
involvement of ATP is not necessary. Fat body
cells on the hemolymph side of the gut rapidly
synthesize absorbed glucose into the disaccharide Illustrative Examples of Diversity
trehalose, keeping the hemolymph concentration in Food, Form and Function of the
of glucose low in most insects. Consequently, Alimentary Canal
even low concentrations of glucose in the gut
have a favorable diffusion pathway to the hemo- The following examples are not intended to be a
lymph and continue to be absorbed passively. In comprehensive review of foods, and alimentary
larvae of a few lepidopterans that have been canal structure and physiology, but will merely
carefully studied (Manduca sexta, Philosamia highlight interesting diversity.
cynthia, Bombyx mori), amino acids are actively Opportunistic feeders may have evolved modi-
absorbed by transport proteins in the apical fications to capture and store food when available,
membranes of midgut columnar cells. Energy and thus survive lean periods when food is not avail-
for absorption comes from the high K+ concen- able. For example, the foregut of the praying mantis,
tration in the gut lumen and high transepithelial Tenodora sinensis, is long and wide, and occupies
potential created by the proton-ATPase pump nearly the entire length of the body, apparently an
active in midgut goblet cells. The transport pro- adaptation for storage of prey when it can be cap-
teins in these membranes show strong specificity tured. The midgut, eight gastric caeca, and hindgut
for particular amino acids, and transport sys- are compressed into the last three abdominal seg-
tems for at least six different amino acids are ments. Probably much of the digestion occurs in
known, and transport systems for other amino theposterior part of the crop with enzymes passed
acids probably will be discovered. In the Colo- forward from the small midgut.
rado potato beetle, Leptinotarsa decemlineata, Considering the universal presence of cellu-
transport proteins for leucine and tyrosine have lose in plant tissues, relatively few insects evolved
been demonstrated in midgut tissue. the ability to use cellulose as a source of nutrients.
Alimentary Canal and Digestion
A 117

Termites, some beetles, a few hymenopterans, and protein poor. Fungi growing in the dead wood and
a few cockroaches do use cellulose as a carbohy- ingested by the larvae may provide additional
drate source. The hindgut of termites is highly spe- nutrients and/or enzymes for digesting the wood.
cialized for housing gut microbiota that provide Hemiptera take xylem or phloem sap, both of
the cellulase enzymes needed to digest cellulose, which are poor in amino acids and protein, but
although there is some evidence that certain ter- usually rich in sucrose (150 to more than 700 mM).
mites may be able to produce some or all of the They typically excrete a copious, dilute fluid, and
several enzymes necessary to completely digest in some, such as aphids, the fluid contains so much
cellulose. Gut variation exists among the castes in a sugar that it is called honeydew. They have to
colony; for example, soldiers in the family Rhinot- ingest large volumes of fluid to get the amino acids,
ermitidae are fed liquid food by the worker caste, and then they have to get rid of the excess water
and do not have to digest cellulose so they have and sucrose. A characteristic evolutionary feature
reduced gut structure. The workers are responsible of the gut in Hemiptera is the filter chamber in
for colony construction and nutrition, and they which a loop of the hindgut is in direct contact
have highly evolved hindgut chambers to hold var- with part of the foregut and a great deal of the
ious types of microbiota. Termites hatch without ingested fluid diffuses directly into the hindgut
their gut microbiota, and lose most of their gut from the foregut without passing through the
symbionts at each molt, but they become reinfested midgut. This, of course, causes loss of some
by feeding upon fluid and excreta from older amino acids and other components that may be
nymphs. Termites in the family Termitidae have needed, but water and sucrose, both of which are
symbiotic bacteria in the hindgut, while termites in in excess of needs, are the major components lost.
some other families have flagellate protozoans as The filter chamber is able to concentrate gut fluid
well as bacteria in a multi-compartmented hindgut, up to 10-fold in some xylem feeders (Cicadoidea
and they get some or all of their cellulase(s) from and Cercopoidea), but only about 2.5-fold in
their symbionts. Some termites, the Macrotermi- members of the Cicadelloidea, which are phloem
tinae, cultivate fungus gardens in their under- feeders. Xylem feeders probably need to concen-
ground nests and get their cellulases from the trate xylem fluids more because of the lower amino
conidiophores of the fungus. Symbionts in the acid content (310 mM amino acids per liter in
hindgut of some termites can capture atmospheric xylem fluid) than do phloem feeders (1565 mM
nitrogen in an organic form, which is probably amino acids per liter in phloem fluid).
quite important to many termites because their Pre-oral digestion with enzymes secreted into
diet of wood is relatively low in proteins. Some the prey occurs in many of the predacious beetles.
fungus-growing termites convert some of the Seed feeders also employ pre-oral digestion by
protons (H+) and carbon dioxide from the initial injecting salivary secretions and possibly regurgi-
fermentation of glucose into methane (CH4), and tated midgut enzymes into the seed, allowing these
some investigators have suggested that termites are to liquefy part of the seed, and then sucking the
a significant environmental source of methane, a nutrients and enzymes back. Pre-oral digesters
greenhouse gas. often reflux the liquefied contents by repeatedly
Larvae of the woodwasp (Hymenoptera, imbibing and then reinjecting the mixture of
Symphyta, Siricidae, genus Sirex) acquire cellulase enzymes and digested nutrients into the seed.
and xylanase from fungi ingested with the wood Refluxing likely conserves enzymes longer, gives
on which they feed. Larvae of cerambycid beetles them more opportunity to function, and allows
and of some other beetles feed upon wood in the insects to use more of their potential food.
down or dying trees; they generally have long life In honeybees and some other related
cycles because wood is so nutrient poor, especially hymenopterans, the midgut is closed off from the
A Alimentary System

hindgut by a plug of cellular tissue during larval midgut so that digestion can begin. Feeding is
development, and any food that cannot be nearly continuous when plenty of food is avail-
digested and absorbed into the body must remain able, and larvae may ingest more than their body
in the midgut. Just before pupation the connec- weight in food daily. Food moves rapidly through
tion between midgut and hindgut is opened, and the relatively straight gut and frass droppings are
accumulated undigested residue, such as the frequent in phytophagous caterpillars. Because
shells of pollen grains, is excreted into the cell. the larval and adult forms of Lepidoptera have
Adult honeybees clean the cell and the larva very different life histories and food habits, the
pupates inside the cell. adult gut is quite different from that of the larva.
Nectar taken by male and female mosquitoes Many adult Lepidoptera feed only upon nectar,
is stored in a large, sac-like crop that is a diver- which is stored in the crop and slowly released
ticulum from the foregut, but blood meals taken into the midgut for digestion to simple sugars.
by the females are passed directly into the midgut Some adult Lepidoptera have vestigial mouth-
for the beginning of digestion. The midgut is dif- parts and do not feed at all; they survive and
ferentiated functionally into an anterior and a (females) produce eggs at the expense of body
posterior region. The anterior part secretes carbo- substance, and they generally live only a few
hydrate digesting enzymes, and nectar compo- days. An unusual food utilized by Tineola bissel-
nents are digested as fluid from the crop and is liella larvae (clothes moth) is wool, and larvae
passed into the anterior midgut. This arrange- have a very strong reducing action in the midgut
ment keeps possible trypsin inhibitors that may that breaks disulfide bonds between adjacent
be present in nectar away from the site of protein loops of the proteins, causing the wool proteins
digestion, which occurs in the posterior midgut. to lose their three-dimensional shape and unfold.
Simple sugars resulting from digestion, or those This allows more access for protein digesting
already in the nectar, are absorbed in the anterior enzymes.
midgut. The posterior midgut cells secrete trypsin-
like enzymes and protein (blood) digestion and
absorption occur in the posterior midgut. The References
posterior midgut cells, more so than anterior
midgut cells, have extensive microvilli and basal Chapman RF (1998) The insects: structure and function.
infoldings characteristic of secretion and absorp- Cambridge University Press, Cambridge, UK, 770 pp
Klowden MJ (2002) Physiological systems in insects. Aca-
tive processes. The midgut cells in this region get demic Press, New York, NY, 415 pp
stretched by the large volume of blood that a mos- Nation JL (2002) Insect physiology and biochemistry. CRC
quito takes if it is allowed to feed to repletion. Press, Boca Raton, FL, 485 pp
Consequently, the cells have several types of con-
necting structures (desmosomes) between cells to
help hold them together and prevent excessive Alimentary System
leaking of materials in or out between cells while
they are stretched. The alimentary system (canal) is a system of tubu-
Larvae of Lepidoptera have a very short lar structures that takes in food at the mouth,
foregut, a large, long, relatively straight midgut, stores the food, fosters digestion and absorption of
and a short hindgut. There is no storage or diges- nutrients, and allows excretion of waste materials
tion in the short, nearly vestigial foregut. Nearly from the rectum. It is conveniently divided into
all lepidopterous larvae are phytophagous feed- the foregut, midgut, and hindgut (Fig.35).
ers, and the gut modifications appear to be an Alimentary Canal and Digestion
adaptation to pass food quickly into the long Foregut
Allegheny Mound Ant, Formica exsectoides (Hymenoptera: Formicidae)
A 119

foregut midgut hindgut

esophagus crop ventriculus pylorus ileum rectum

buccal anus
cavity proventriculus
mouth Malpighian tubule

Alimentary System , Figure 35 Generalized insect alimentary system (adapted from Chapman, The
insects: structure and function).

Midgut likely have other effects as well, but they are still
Hindgut relatively unknown (contrast with allatostatins).
Juvenile Hormone

The notal plate of the meso- or metathorax in a
pterygote insect. A family of beetles (order Coleoptera). They com-
Thorax of Hexapods monly are known as comb-clawed beetles.
The portion of the thorax to which the wings are Allegheny Mound Ant, Formica
attached. exsectoides (Hymenoptera:
Thorax of Hexapods Formicidae)

gregory s. paulson
Allatostatins Shippensburg University, Shippensburg, PA, USA

These are neuropeptides from neural and non- The Allegheny mound ant, Formica exsectoides
neural tissues that affect the corpora allata, inhib- Forel, is a common mound-building ant of the
iting production of juvenile hormone. They likely northeastern and central United States. Workers
have other effects as well, but they are still rela- are approximately 3/8 inch long (1 cm) with a
tively unknown (contrast with allatotropins). reddish-tan head and thorax, and a dark brown
Juvenile Hormone abdomen. In suitable habitat, F. exsectoides will
form dense populations, their presence easily dis-
cernible due to conspicuous mound-type nests
Allatotropins which can be as large as 15 feet (4.6 m) in diame-
ter and 4 feet (1.2 m) high. At one site near
These are neuropeptides from neural and non- Altoona, Pennsylvania, researchers counted more
neural tissues that stimulate the corpora allata, than 30 large mounds per acre. Despite the con-
resulting in synthesis of juvenile hormone. They spicuous nature of F. exsectoides nests and its wide
A Allegheny Mound Ant, Formica exsectoides (Hymenoptera: Formicidae)

geographic range, relatively few papers concern- Habitat has a significant effect on physical char-
ing this species have been published since H.C. acteristics of F. exsectoides nests. Forest nests tend to
McCooks first paper in 1877. be significantly larger in height, width, length, nest
Nests tend to be clustered in the habitat footprint and volume than nests in meadows. In
(Fig.36). Nests in each cluster often will share for- general, the nests are round, but it is not uncommon
aging trails and resources. Although there is little to find elongated nests that are orientated to the sun.
aggression between workers from different nests or The shape of a nest and its orientation may help the
nest clusters, the workers still show fidelity to a ants to maintain a relatively constant internal tem-
home nest. Nests of F. exsectoides have multiple perature and relative humidity.
queens (polygynous) although there is tremendous
variation in the number of queens per mound.
Over 1,400 queens were reported in one mound,
but that is probably an anomaly. Most mounds
probably contain fewer than 20 queens. Due to the
Andrews EA (1925) Growth of ant mounds. Psyche
large number of queens, the reproductive output of 32:7587
a nest can be prodigious resulting in large numbers Andrews EA (1927) Ant mounds as to temperature and sun-
of workers in a colony. More than 250,000 workers shine. J Morphol Physiol 44:120
have been found in some nests. Reproductive forms Bristow CM, Cappaert D, Campbell NJ, Heise A (1992) Nest
structure and colony cycle of the Allegheny mound ant,
(alates) are present in the nests from mid-summer Formica exsectoides. Forel (Hymenoptera: Formicidae)
until early fall. Activity of the ants is related to Insectes Sociaux 39:385402
ambient conditions but, in general, workers are Bristow CM, Yanity E (1999) Seasonal response of workers of
the Allegheny mound ant, Formica exsectoides.
active from late March until November. Formica
(Hymenoptera: Formicidae) to artificial honeydews of
exsectoides are generalist predators, scavengers and varying nutritional content. Great Lakes Entomol
collect honeydew from symbiotic hemipterans. 32:1527

Allegheny Mound Ant, Formica Exsectoides (Hymenoptera: Formicidae), Figure 36 Three large nest
mounds of Formica exsectoides in a Pennsylvania forest clearing.
A 121

McCook HC (1877) Mound-making ants of the Alleghenies, of years, plants have done everything possible to
their architecture and habits. Trans Entomol Soc Am
make life miserable for insects. On the other hand,
Rowe HC, Bristow CM (1999) Sex ratio and sexual dimor- insects have returned the favor many times over. In
phism in Formica exsectoides, the Allegheny mound ant recent geological time, plants and insects have
(Hymenoptera: Formicidae). Great Lakes Entomol changed their spots (plant-feeding strategies),
resulting in an incredible point-counterpoint rela-
tionship of these organisms that is characterized by
some remarkable developments.
Allele As far as nutrients are concerned, different
kinds of plants (species) are fairly similar and can
One of two or more alternative forms of a gene at provide an insect herbivore with the basic nutri-
a particular locus. If more than two alleles exist, ents required for growth and development. These
the locus is said to exhibit multiple allelism. compounds (chemicals) are called primary com-
pounds because they are required for the insects
growth, development, and reproduction. All insects
Allelochemic require these compounds and in theory they
should be readily available from a wide variety of
A non-nutritional chemical produced by one plant species. But most species of insects, rather
species (often a plant) that affects the growth, than feeding on many different kinds of plants,
health, or behavior of another species (often an limit their plant menu to a relatively small number
herbivore). of plant species (monophagy), most of which are
Allelochemicals related. Significantly, the limited preferences that
insects have for their food plant species are due to
non-nutritive compounds that usually vary from
Allelochemicals one plant group to another. These compounds are
not related to the primary compounds identified
murray s. blum with growth and development, and it is apparent
University of Georgia, Athens, GA, USA that these plant-derived compounds (allelochemi-
cals) generally have functions related to other spe-
For thousands of years insects and plants have been cies of organisms. These compounds are obviously
locked in a battle for which survival is the ultimate not primary compounds but rather secondary
prize. For insects, plants constitute food sources for compounds (non-nutritive) whose manufacture
growth and development, and in some cases, sites has been described as secondary metabolism.
for reproduction. On the other hand, plants attempt Indeed, these allelochemicals appear to be respon-
to counter insects feeding on their tissues (her- sible for both the associations and non-associations
bivory) so that their own vigorous growth and that insects have with specific groups of plant spe-
development will occur and lead to reproductive cies. In essence, it would be no exaggeration to state
success. The consequences of this warfare are great that the host-plant preferences of insects really
and for humankind the outcomes of these battles reflect the ability of an insect species to either tol-
may be of major economic significance in terms of erate, or be repelled by, an allelochemical. Alle-
the production of various foods. The welfare of lochemicals are not mysterious compounds but
various human populations can be threatened if rather are a very important part of the everyday
hordes of ravenous insects consume specific crops world, especially in terms of human food prefer-
that are the mainstays of these populations. But ences. In a sense, the strong food preferences
plants do not take this lying down. Over thousands exhibited by insects are not so different from those
A Allelochemicals

of humans, with one striking exception. Many icotine in the leaves before they are ten years old.
insect species are locked in to specific food plants, Nitrogen-containing compounds (alkaloids) pro-
and these insects will reject a foreign plant species duced by opium poppies are powerful repellents
and die in the absence of their normal food plant. for a wide range of insect species, and there is no
On the other hand, there is little evidence that doubt that compounds such as morphine and her-
human beings will subject themselves to starvation oin, which are powerful human narcotics, were
if their favorite foods are not readily available. evolved to deter herbivores rather than to func-
tion as narcotics for humans.
Alkaloids such as nicotine have been adapted
A World of Allelochemicals to function as insecticides, and a variety of plant
products such as derris, rotenone, ryania, and
Fruits and vegetables possess characteristic odors sabadilla are also used as insecticides in different
and tastes that create desire (preference) for these cultures. In some cases, allelochemicals such as
foods. Significantly, these tastes and smells are prunasin in cherry leaves cause poisoning in
notidentified with the primary compounds respon- livestock. Not to be outdone, humans have fre-
sible for plant growth and development, such as quently utilized the alkaloid strychnine to murder
sugars, fats, and proteins. Therefore, the plant has people. However, it would be a mistake to lose
invested in producing a variety of chemicals that track of the fact that, human abuses notwithstand-
will not help it grow or reproduce. While an onion ing, these allelochemicals were evolved as plant
may possess a distinctive odor and taste for both protectants long before humans appeared. Obvi-
insects and humans (not necessarily the same odor ously, allelochemicals do not provide plants with
and taste for both), this fact hardly justifies the absolute protection against herbivores. Indeed,
onion spending its energy and resources to produce probably all plants containing allelochemicals
an onion fragrance. On the other hand, if the taste arefed upon by insects, and in many cases these
and odor of onions combine to make this vegetable herbivores are only found on a limited number of
distasteful and repellent to most plant-feeding host plant species. For example, monarch butterfly
insects, then these allelochemicals perform a very caterpillars are limited to the milkweed species.
vital function. In essence, it is generally believed Bark beetles limit their attacks to pines and related
that these secondary compounds are responsible conifers, developing in environments that are rich
for protecting plants from herbivores and possibly in toxic turpentines. These insects have breached
pathogens as well. A brief examination of some the chemical defenses of their hosts, and in so
well-characterized allelochemicals offers a means doing, they have captured specific kinds of food
of examining these compounds as agents of defense plants that are either repellent or highly toxic to
both as toxins and as repellents. most other species of insects. Guaranteed these
Oleander, which has a very limited number of forbidden fruits, these herbivores should have to
herbivores, is extremely toxic because of the pres- share their food resources with very limited num-
ence of allelochemicals that are somewhat related bers of competitors. Barring an ecological disaster
to cholesterol. The odor of the plant probably con- does not devastate the populations of their host
stitutes an early-warning system that makes poten- plant species, this specialization should have much
tial herbivores aware of the danger of feeding on to recommend it. On the other hand, many insect
this plant. The same can be said for the tobacco species choose a lifestyle which is characterized by
plant which, like oleander, does not have too many feeding on a variety of unrelated plant species.
insect herbivores. Leaves of the tobacco plant are Insects like the monarch butterfly and bark
quite toxic, but in some South American popula- beetles that are restricted to a limited number of
tions young children become addicted to the related plant species are referred to as specialists.
A 123

These herbivores have become resistant to the grasshoppers, aphids, lacewings, beetles, wasps,
toxic effects of their host plant allelochemicals, butterflies and moths. In general, these insects are
and in many cases they appear to be completely brightly (= warningly) colored, a characteristic
immune to the plant toxins they ingest. In the case described as aposematic. Armed with the toxins
of monarch caterpillars feeding on milkweeds, it from their food plants, large insects such as bril-
has been demonstrated that these larvae actually liantly colored grasshoppers move very slowly, as
grow more rapidly on milkweed plants containing if to advertise their poisonous qualities to the
the highest concentration of toxins. world. Obviously the term toxic is relative, since
Indeed, allelochemical concentrations may these insects routinely sequester these allelochem-
begenerally quite high, often averaging 510% of icals during normal feeding. However, since these
the dry weight of the plant. By contrast, plant- specialists are physiologically adapted for ingest-
feeding generalists feed on a wide range of plant ing these compounds, their ability to tolerate these
species, often unrelated. However, in general, allelochemicals is really not surprising. On the
theseherbivores select plant species in which the other hand, non-adapted species (e.g., predators)
concentrations of allelochemicals are not too would certainly encounter toxic reactions if they
high, enabling them to process low levels of a ingested these toxic plant products.
widevariety of plant toxins. The fates of allelochemicals, which are usually
The immunity of specialists to the toxic effects present in mixtures, are not at all predictable after
of the allelochemicals in their diets demonstrates ingestion by an adapted herbivore. Although many
that for these insects these compounds can no lon- compounds are sequestered immediately after
ger be considered poisons. Surprisingly, the basis ingestion, others may be metabolized before being
for this important allelochemical resistance, which stored, or even eliminated after being metabolized.
has great economic significance, was only under- In other cases selected allelochemicals in a mixture
stood about thirty years ago. may be absorbed and sequestered whereas other
compounds in the mixture may be eliminated
immediately. An examination of the options for
Sequestration and its initially processing ingested allelochemicals
Consequences emphasizes the versatility of specialists in treating
the toxic compounds produced by their food
Insects such as the monarch butterfly store com- plants.
pounds in their tissues that render them unpalatable
to predators. These compounds, the cardenolides,
were ingested by the larvae from their milkweed food Sequestration of Insect Toxins by
plants, and retained in their bodies into the adult Vertebrates: A Significant
stage. The storage of these milkweed compounds is Allelochemical Phenomenon
called sequestration, and constitutes a widespread
phenomenon among specialists feeding on alle- It has become evident that the allelochemical rela-
lochemical-rich plants. In a sense, sequestration rep- tionship of insects and plants is paralleled by a
resents the insects success in utilizing the plants similar relationship of amphibians and insects. It is
chemical defenses for its own purposes. Indeed, now recognized that the sequestration of ingested
sequestration can be regarded as a form of detoxica- toxic insect compounds by vertebrates differs little
tion since potentially toxic compounds are removed from this phenomenon in insect herbivores and
from the circulation and stored in the tissues. plants. In essence, a variety of insect toxins is
Sequestration has been detected in at least sequestered by amphibians and these compounds
seven orders of insects including species of toxic have similar protective functions for frogs and
A Allelochemicals

insects (see Allelochemicals as phagostimulants). rocessing it.An insect species may utilize a vari-
Frogs exploit insect allomones (defensive com- ety of adaptive strategies for processing a single
pounds) as if they were animal allelochemicals, compound that is characteristic of the host plant
and it seems worthwhile to emphasize this congru- defense.
ency in examining the scope of allelochemistry.
Frogs in the genus Dendrobates contain mono-,
di-, and tricyclic alkaloids which are clearly of ant Immediate Allelochemical Excretion
origin. The alkaloids, termed pumiliotoxins, appear
to be products of ant species in the genera Brachy- Some insects essentially fail to absorb ingested
myrmex and Paretrechina and constitute the only allelochemicals from the gut. These compounds
known dietary source of alkaloids of these frogs, are excreted directly and are concentrated in
not unlike the specialist insects feeding on narrow the feces. A lymantriid moth larva that is a
plant diets enriched with allelochemicals. The same specialist on the coca plant, which is the source
phenomenon has been described for the myrmicine of the alkaloid cocaine, rapidly excretes this
ant Myrmicaria melanogaster which synthesizes ten compound with only traces being found in the
alkaloids. Some of these alkaloids have previously blood. However, cocaine may still have defen-
been identified in a dendrobatid frog and a toad. sive value for the larva as part of an oral regur-
Neurotoxic steroidal alkaloids, the batrachoto- gitate that is externalized when the larva is
xins, have been isolated from New Guinea birds in disturbed.
the genera Pitohui and Ifrita. These compounds are Three different species of moth larvae that
among the most toxic natural substances known, feed on tobacco plants rapidly excrete nicotine,
and they are not produced by captive birds, sug- a very toxic and reactive alkaloid. There is no
gesting a dietary source. Recently, the batrachotox- evidence that nicotine is absorbed from the gut
ins were identified in beetles in the genus Chloresine of any tobacco feeder, but as is the case for the
(Melyridae) which are normally fed on by the bird moth larva excreting cocaine, nicotine in oral or
species. Since the genus Chloresine is cosmopoli- anal exudates constitutes an excellent defensive
tan, it is the possible source of some of the avian compound.
alkaloids found in birds in different areas.
Vertebrate sequestration of alkaloids from
insects has only recently been explored. Clearly Allelochemical Metabolism
this chemical storage has a common denominator
with sequestration of alkaloids by insects (see Alle- Many insect specialists rapidly metabolize
lochemicals as pheromonal precursors) and should ingested allelochemicals which are then seques-
be examined as a paradigm of comparative physi- tered, or in some cases excreted. Nicotine, which
ology. Clearly, insects are pivotal to both systems, is both highly reactive and very toxic, is con-
either as food for vertebrates or food for insects, verted to a non-toxic metabolite called cotinine
with sequestration the major common feature. by both tobacco-feeding insects and those that
are not tobacco feeders. Since cotinine has virtu-
ally no toxicity to insects, it is probable that its
Initial Processing of production from nicotine constitutes true
Allelochemicals by Specialists detoxication.
Cabbage-feeding insects feed on plants
Once an adapted insect has ingested an alle- that are rich in sinigrin, a compound that yields
lochemical, a menu of options is available for a highly toxic mustard oil when metabolized.
A 125

Although sinigrin can be sequestered without Selective Biomagnification of

generating the reactive mustard oil in a variety Allelochemicals in Tissues
of cabbage-feeding species, cabbage butterflies
(whites) actually break down sinigrin and seques- There is little indication that the profiles of insect-
ter the highly reactive mustard oil. For these stored allelochemicals in any way mirror those of
butterflies, the mustard oil is more suitable for their host plant. In a sense, each insect species treats
storage than sinigrin. ingested allelochemicals distinctively, so that a com-
Larvae of the tiger moth Seirarctia have pound totally excreted by one species may consti-
evolved a novel strategy for coping with the toxic tute the main sequestration product of another.
effects of MAM, a compound derived from cycasin The very toxic grasshopper Poekilocerus bufo-
which is a constituent in the cycad leaves upon nius sequesters only two of the cardenolides that it
which they feed. When larvae encounter MAM, ingests from its milkweed food plant. Similar
they convert it to cycasin which is absorbed through selectivity is shown by moth larvae (Syntomeida
the gut wall before being sequestered. Since the species) which sequester oleandrin, the main ste-
enzyme that produces cycasin or MAM is only roid found in the leaves of oleander. On the other
found in the gut, once cycasin crosses the gut wall hand, a variety of other insects feeding on olean-
into the blood prior to sequestration there is no der leaves do not sequester oleandrin.
chance of MAM being generated from cycasin. Similar unpredictability characterizes the
Many species of moths, butterflies, and sequestration of pyrrolizidine alkaloids by moth
grasshoppers feed on plant species that produce larvae. Tiger moths (Amphicallia species) seques-
extremely toxic compounds known as pyrroliz- ter the alkaloids crispatine and trichodesmine,
idine alkaloids. These alkaloids, present as mix- whereas the main alkaloid present is crosemper-
tures, are frequently sequestered by these ine. Another tiger moth (Tyria species) concen-
specialist insects and, in some cases, metabolized trates senecionine in its tissues in spite of the fact
plant compounds are the preferred storage forms. that this compound is a trace constituent in the
For example, larvae of the tiger moth (Tyria spe- leaves. Tyria is no less curious as a sequestrator
cies) feed on ragwort and primarily sequester because it stores jacobine, jacozine, and jacoline as
the alkaloid seneciphylline, although this com- minor constituents in adults, yet these three com-
pound is present in the plant as the N-oxide. pounds are major alkaloids in the leaves.
Conversely, the grasshoppers of the Zonocerus
species convert the ingested alkaloid monocro-
taline to its N-oxide before sequestering the The Diverse Functions of
compound. Captured Allelochemicals
Insects feeding on milkweed metabolize the
toxic cardenolides (steroids) produced by these While highly concentrated allelochemicals may
plants, converting them into compounds that constitute a major deterrent to non-adapted
can be readily sequestered. The milkweed bug insects, these compounds can represent a real
(Oncopeltus species) oxidizes cardenolides as a treasure trove for species for which these plant
mechanism for converting these steroids into products are non-toxic. Indeed, in the course of
compounds that can be efficiently sequestered. exploiting for their own protection compounds
Similarly, larvae of the monarch butterfly store that are repellent or toxic to most insect species,
metabolized cardenolides in tissues after oxidiz- specialists have gone beyond the point of simply
ing these compounds into suitable chemical being resistant to allelochemicals. In many cases, a
forms for sequestration. variety of specialist species have utilized the rich
A Allelochemicals

allelochemical pool that is available in order to association is typical of a variety of insect species
develop a menu of remarkable functions. and their bacterial symbiotes, then a multitude
of insect-bacterial relationships may require re-
evaluation of possible examples of insect seques-
Insect Sequestration of Bacterial tration of bacterial allelochemists.
Compounds and their Glandular Non-pathogenic bacteria are commonly housed
Secretion in insects and, in a sense, these prokaryotes are
sequestered by their insect hosts. Furthermore, if the
Prokaryotes (bacterial types) are almost every- bacteria synthesize toxic compounds which may be
where and their widespread association with externalized from a defensive gland (prothoracic
insects is certainly well established. But the bases glands of dytiscids), then the bacterial products may
for these diverse bacteria-insect relationships are, be regarded as bacterial allelochemicals that have
for the most part, terra incognita. However, very been sequestered. Indeed, bacterial compounds of
recent research suggests one very surprising func- symbiotic bacteria of insects clearly constitute an
tion for bacteria in insect glands. unrecognized group of allelochemicals.
All major types of metabolism evolved in
prokaryotes and the success of these organisms
was both cause and effect of changing environ- Additives in Defensive Glands
ments on earth. If these bacteria are sequestered in
insect secretory glands, their great metabolic abili- Milkweed bugs (Oncopeltus species) add cardeno-
ties could be utilized to biosynthesize bacterial lides, derived from their milkweed host plants, to
allelochemicals which could be used as potent their thoracic defensive gland secretion which
defensive compounds. This possibility appears to considerably enhances the deterrency of their
have been realized as a product of the virtual ubiq- secretion. Similarly, a warningly colored generalist,
uity of both insects and their biosynthetically ver- the lubber grasshopper (Romalea guttata) incor-
satile prokaryotes. porates a large number of allelochemicals derived
Predaceous diving beetles (Dytiscus species) from a variety of plant species into its thoracic
are distinguished by their ability to produce gland secretion. This grasshopper generally feeds
defensive steroids, some of which are novel ani- on plants with low concentrations of allelochemi-
mal products that are limited to species of diving cals, but if it is fed high concentrations of plants
beetles. Furthermore, insects do not synthesize with known repellents (e.g., onion), the odorous
cholesterol which in insects must be obtained secretion can be highly deterrent.
from exogenous sterols. However, it now appears Another toxic grasshopper, Poekilocerus bufo-
that the surprising steroidal versatility of dytisc- nius, utilizes allelochemicals as the mainstay of its
ids may reflect the biosynthetic elegance of bacte- defensive secretion. This aposematic (very warn-
ria rather than insects. ingly colored) insect sequesters two of six carde-
Adult diving beetles may contain concentra- nolides from its milkweed diet which are the major
tions of at least 10 bacterial species, mostly detected irritants in the secretion when it is sprayed at
in a variety of organs. Culturing individual bacte- adversaries. Utilization of allelochemicals as
rial species resulted in the identification of diverse defensive gland constituents is particularly pro-
steroids that had previously been characterized in nounced in the swallowtail larvae of Atrophaneura
the prothoracic defensive glands of the adults. The alcinous, which feed on leaves that are rich in toxic
steroid-rich secretions of these glands function as aristolochic acids. Seven aristolochic acids are
vertebrate deterrents that can cause emesis of fish sequestered by the larvae and all are transferred
that swallow these beetles. If the dytiscid-bacterial tothe defensive gland in the head. The acids are
A 127

concentrated in the gland and are the major deter- Allelochemicals as Inhibitors of Toxin
rents for birds. Production

Some plant toxins are present in plants in an

Regurgitation and Defecation of inactive form only to be converted to toxic
Allelochemicals compounds after ingestion by herbivores. This is
particularly true for many cyanogens (cyanide-
The intestines of stimulated grasshoppers can dis- containing toxins) that generate cyanide when
charge ingested plant products which may serve as the leaf surface is broken as would occur with a
repellents for predators. Regurgitated allelochem- plant feeder. It now appears that cyanogenesis
icals can effectively repel ants, as is the case for (producing cyanide) in damaged leaves may be
anal discharges from the hind gut. When tactually inhibited by allelochemicals that are compart-
stimulated, the milkweed bug, Oncopeltus fascia- mentally isolated from the cyanogens in the
tus, also defecates a solution containing repellent intact leaves.
allelochemicals. In this case, they are cardenolides Leaves of papaya, Carica papaya, contain two
ingested from their milkweed food plant. cyanogens that yield hydrogen cyanide after enzy-
matic attack. However, tannins, which are widely
distributed in plants, inhibit the release of cyanide
Allelochemicals as Tissue Colorants caused by the action of enzymes that attack the
cyanogens. Insects attacking plants containing
The cuticular (skin) coloration of many insects is cyanogens may have adapted tannins to prevent
diet-dependent and is highly adaptive since it cyanide release, a strategy that may be suitable for
enables the insect to respond in a positive way to other plant groups that yield toxic products after
its background color. Diet-induced changes may leaf damage.
result in the insect being cryptic (background
matching), whereas aposematic species can be
background contrasting. Background quality,
which is of great survival value, appears to be Allelochemicals as Pheromonal
controlled by allelochemicals that are widespread Precursors
in the diets of moths, butterflies and true bugs.
These insects are particularly sensitive to the Bark beetles (Scolytidae) in the genera Dendroc-
carotenoids (e.g., tomato red) that fortify their tonus and Ips convert the hydrocarbons pro-
host plants. duced by their pine hosts into alcohols that are
If the large white butterfly, Pieris brassicae, is utilized as either aggregation or sex pheromones
reared on its normal diet of cabbage leaves, the (communication compounds) by the attacking
pupae are green and contrast with their back- beetles. Similarly, butterflies in the family
ground. This toxic insect contains high concentra- Nymphalidae and moths in the family Arctiidae
tions of carotenoids, and the carotenoid lutein is convert pyrrolizidine alkaloids (PAs) into sex
concentrated in the cuticle. On the other hand, if pheromones that are especially critical during
these insects are reared on an artificial diet lacking courtship. The PAs may be collected from dam-
carotenoids, they possess a turquoise-blue color- aged plants by males to be transformed into
ation and exhibit no response to background. In sexual pheromones that constitute the key to
the absence of carotenoids, these insects are quite reproductive success. For these males, the alle-
conspicuous on their background and could be lochemicals (PAs) are identified with reproduc-
readily detected by predators. tive fitness.
A Allelochemicals

Allelochemicals as Structural Paint Beetle larvae in the genus Chrysomela also

convert a toxic allelochemical into a metabolite
Some insects actually paint structures with with considerable importance in growth and
ingested compounds possessing considerable bio- development. These larvae feed on leaves of wil-
logical activity. Larvae of the parsnip webworm, low (Salix), a rich source of salicin, a toxic metab-
Depressaria pastinacella, apply ingested alle- olite. Metabolism of salicin yields a very effective
lochemicals to silk-webbed flowers that serve as defensive compound that is sequestered by the
housing units. The applied compounds are derived larvae in defensive glands. In addition, this metab-
from wild parsnip, a food plant that is rich in olism generates enough glucose to account for
highly toxic furanocoumarins. These compounds about one-third of the daily caloric requirements
are sequestered in the silk glands before being of the larvae. Salicin should be regarded as an alle-
applied to the flowers in which the larvae reside. lochemical nutrient.
Since the larvae are quite sensitive to ultraviolet
light, the presence of UV-absorbing furanocou-
marins on their silken housing is highly adaptive. Allelochemicals as Agents of Sexual
In addition, because these allelochemicals possess Development
pronounced antimicrobial activity against bacte-
ria and fungi, their presence on the silk can act as Tiger moths in the genus Creatonotus feed on
a major barrier to pathogens. plant species that produce high concentrations of
pyrrolizidine alkaloids (PAs). These compounds
are converted to sex pheromones by the males.
Allelochemicals as Metabolites in Additionally, these allelochemicals control the
Primary Metabolic Pathways development of important secondary sexual char-
acters called coremata. The coremata are eversible
Some specialist herbivores metabolize the charac- andraconial (male) organs that are the source of
teristic allelochemicals in their host plants into the volatile sex pheromones of the males, and their
compounds that are of major significance in degree of development is controlled by the amount
growth and development. In essence, these spe- of PAs ingested by the developing larvae. In effect,
cialists exploit their food plants by utilizing not PAs are functioning as male hormones that regu-
only their primary nutrients for growth and devel- late both sex pheromone production and develop-
opment, but their allelochemicals as well. ment of the coremata.
Larvae of the bruchid beetle, Carydes brasil-
iensis, develop exclusively on seeds of a legume
(pea family) that contains canavanine, a foreign Allelochemical Discharge from Non-
amino acid related to arginine. Canavanine is glandular Reservoirs
highly toxic when incorporated into proteins by
non-adapted herbivores. On the other hand, lar- Some insects sequester ingested allelochemicals in
vae of C. brasiliensis metabolize canavanine into non-glandular reservoirs that can be evacuated
products of great metabolic significance. Large upon demand. Gregarious larvae of the European
amounts of ammonia are generated for fixation pine sawfly, Neodiprion sertifer, sequester toxic tur-
into organic compounds, and an amino acid is pentine terpenes in foregut pouches. These pine-
produced from canavanine for ready metabolism. derived compounds can be discharged upon
Thus, the very toxic allelochemical of the legume demand to function as highly effective predator
has been thoroughly exploited by the beetle larvae deterrents. Similarly, lygaeids such as the milkweed
as a source for key nutrients. bug, Oncopeltus fasciatus, sequester cardenolides
A 129

from their milkweed hosts in dorsolateral spaces alkaloid sources, the copulatory bonus ensures that
on the thorax and abdomen. Significantly, high these toxic allelochemicals will be available to protect
concentrations of cardenolides are stored in these both the female and her eggs. It is also very significant
spaces, resulting in a concentrated deterrent dis- that the resistance of the spermatozoa to the known
charge which repels potential predators. toxic effects of the pyrrolizine alkaloids enables the
copulatory bonus strategy to be highly adaptive.

Allelochemicals as Defensive Agents

ofEggs Allelochemicals as Synergists
Insects ingesting allelochemicals often utilize these
compounds as protectants for the next generation The intimate relationship of specialist insects and
of insects. These plant compounds may be seques- their food plants is exemplified by the turnip aphid,
tered in the eggs in order to provide a formidable Lipaphis erysimi, and its alarm pheromone. This
defense against predators and pathogens. The aphid is typical of many aphid species. Paired glands
insect embryo must be resistant to the toxic effects near the tip of the abdomen secrete an alarm phero-
of the allelochemicals that have been sequestered mone that causes both adults and larvae to disperse
in the reproductive system. For example, chry- and drop off of the food plant. The alarm phero-
somelid beetle adults feeding on willow and pop- mones synthesized by the aphids are key communi-
lar sequester the toxic allelochemical salicin which cations chemicals that enable these insects to
is used to fortify the eggs. Salicin has different abandon ship when threatened by a predator. Sur-
functions in the embryo and the larvae. For the prisingly, (E)-B-farnesene, the major alarm agent
embryo, salicin is a deterrent toxin which can kill for a large variety of aphid species, is only weakly
ants. For the young larvae, salicin is converted to active when secreted by the turnip aphid. However,
salicylaldehyde, a powerful repellent that is not the activity of this pheromonal secretion is increased
frequently encountered in insects. A wide variety appreciably by allelochemicals that act as powerful
of allelochemicals are sequestered in insect eggs synergists for the major alarm pheromone. These
which includes pyrrolizidine alkaloids, aristo- synergists are derived from typical food plant com-
lochic acids, cannabinoids, quinones, cardenolides pounds that have been modified by the aphids.
and mustard oils. It is evident that the females of a
large number of species have appropriated their
host-plant defenses (allelochemicals) for protec- Allelochemicals as Phagostimulants
tion of their eggs.
The close relationship of insect specialists and
their allelochemicals is further demonstrated by
Allelochemicals as a Copulatory Bonus some species of sawflies and chrysomelid beetles
which feed on very bitter food plants. Adults of
Females may obtain allelochemicals suitable for their the turnip sawfly, Athalia rosae, feed on the sur-
own protection and that of their eggs from the semi- face of a plant that is not a larval food plant. Com-
nal ejaculate. For example, males of ithomiine but- pounds in the leaf surface that are responsible for
terflies gather pyrrolizidine alkaloids (PAs) from their bitter taste are powerful phagostimulants for
flowers and decomposing foliage and about half of A. rosae. In addition, these bitter compounds are
the PAs are channeled to the spermatophore (sperm incorporated into the cuticle, thus providing these
packet) that is transferred to the female during copu- sawflies with a cuticular set of armor to protect
lation. Since the females are rarely found feeding on against aggressive predators.
A Allelochemicals

Similarly, species in three genera of chry- eridania, from a host of allelochemicals. Unrelated
somelid beetles utilize cucurbitacins, compounds plant compounds rapidly induce enzymatic in
found in their squash and pumpkin hosts, as creasesof 2 to 3-fold in larvae. Significantly, the rise
phagostimulants that are biomagnified in their in P-450 activity is immediate and proceeds rapidly
bodies. The beetles are rendered distasteful and, over much of its course during the first few hours.
as is the case for the sawflies, the allelochemicals These results strongly suggest that P-450 induction
possess dual roles that both induce ingestion is critical to allelochemical tolerance.
and promote sequestration of highly distasteful
Allelochemicals as Allomonal Precursors

Allelochemicals as Inducers of In some cases, insects have produced powerful

Detoxifying Enzymes repellents from allelochemicals in their food plants
and have thus exploited the plants defensive chem-
Both generalist and specialist insects can encoun- istry in a very efficient way. Such a strategy is par-
ter a diversity of allelochemicals with varying ticularly adaptive because the insect has benefited
degrees of toxicity. For generalists this is particu- both nutritionally and defensively from feeding on
larly true since a generalist diet can sample a wide its host.
variety of plant species containing a large diversity Host plant exploitation is particularly pro-
of allelochemicals. On the other hand, specialists nounced in some chrysomelid beetle larvae in the
may encounter fewer allelochemicals but it is likely genera Chrysomela and Phratora. The larvae feed
that these compounds will be at high concentra- on willow and poplar leaves, both of which con-
tions. In the case of both feeding modes, it is tain salicin, a well known feeding deterrent for
obviously necessary to possess mechanisms for non-adapted species. The beetle larvae convert
blunting the toxic properties of the ingested allelo- salicin to salicylaldehyde and glucose, utilizing the
chemicals. Detoxication would appear to consti- former for defense and the latter for growth. For
tute the key process for neutralizing the toxicities these chrysomelid larvae, the conversion of salicin
of ingested allelochemicals. The enzymes chiefly to salicylaldehyde is doubly beneficial. Very little
identified with converting allelochemicals into energy is used to synthesize salicylaldehyde, com-
less toxic compounds are the mixed-function pared to what is required to produce other defen-
oxidases, particularly cytochrome P-450. sive compounds that must be totally synthesized.
Mixed-function oxidases metabolize fat-soluble Because salicylaldehyde is a far more effective
toxins into water-soluble ones that can be excreted. repellent than salicin, the beetle larvae receive a
The level of these enzymes may determine the toler- very important double bonus by converting the
ance of an insect for a particular allelochemical. allelochemical into a compound that can be read-
For a generalist ingesting a large diversity of ily stored and secreted from the defensive glands.
allelochemicals derived from many plant species,
the induction of a variety of these oxidases would
promote the possibility of detoxifying many kinds Allelochemicals as Communicative
of plant compounds. For a specialist, fewer oxidases Jamming Agents
at very high levels would enable the herbivore to
detoxify the very high concentrations of allelochem- In theory, plant species could reduce or eliminate
icals in its restricted food plants. herbivory if the plants generated volatile com-
Mixed-function oxidases play a key role in pounds identical to or similar to the pheromones
protecting the southern armyworm, Spodoptera utilized by herbivores as signals. If these signals
A 131

were behaviorally disruptive, feeding could be commonly exploited plant compounds as key ele-
appreciably diminished, to say the least. ments in their phytochemical defenses.
The wild potato, Solanum berthaultii, has A compound commonly produced by conifers
effectively jammed the pheromonal alarm signal is a-pinene, which inhibits diverse microorganisms
of its potential aphid herbivore. (E)-B-farnesene, including the insect pathogen Bacillus thuringien-
an alarm pheromone of the aphid Myzus persicae, sis. Along with several related compounds, -pinene
is also produced by wild potatoes, resulting in reduces the infectivity of B. thuringiensis for larvae
repellency and dispersion of the aphids. In effect, of the Douglas fir tussock moth, Orgyia pseudot-
the potato has exploited the aphids herbivory by sugata. At concentrations approximating those
utilizing a highly disruptive compound that has found in fir needles, a-pinene increases the 50%
been evolved by aphids as a warning signal. lethal dose for B. thuringiensis by 700-fold.
A pathogenic fungus, Nomuraea rileyi, fre-
quently attacks lepidopterous (moth) larvae such
Quenchers of Phototoxic Allelochemicals as the corn earworm, Helicoverpa zea. However,
thepathogenicity to this larva can be reduced if the
Diverse plant species produce photo-activated moth ingests a tomato alkaloid, -tomatine. If the
compounds that are highly toxic to insects after larvae ingest -tomatine prior to exposure to fungal
digestion. In essence, these compounds generate conidia, it increases larval survivorship considerably.
highly toxic species of oxygen that attack key bio- The alkaloid is a further asset to H. zea because it is
chemicals such as nucleic acids. On the other quite toxic to larval parasites of the corn earworm.
hand, if the herbivore simultaneously ingests The pathogenicity of viral pathogens of H. zea
allelochemicals that are effective quenchers of can also be compromised by host plant allelochem-
toxic oxygen species along with the phototoxins, icals. Chlorogenic acid, a common plant compound,
then survival and prosperity are possible. The is oxidized to chlorogenoquinone by plant enzymes,
availability of these allelochemical antioxidants and this oxidation product binds to a nuclear poly-
has enabled some insect species to utilize food hedrosis virus. Binding to this baculovirus results in
plants that are forbidden fruits for most a reduction in digestibility and a decrease in infec-
herbivores. tivity. Furthermore, it appears that the liberation of
Larvae of the tobacco hornworm feed on a infective virons in the midgut, which is a require-
variety of plant species that contain the phototoxin ment for successful infection, is impaired by the
-terthienyl, a constituent of many species of asters binding of chlorogenoquinone to the baculovirus.
(Asteraceae). However, the additional ingestion of
-carotene reduces mortality from 55% (controls)
to 3% (+carotene) during 48 h. -carotene, an Specialists and Generalists: Two
effective quencher of toxic oxygen species, is con- Selected Case Studies
centrated in the tissues of the larvae where it can
serve as a potent antioxidant for photoactivated Although specialists and generalists may be highly
toxins found in its food plant. efficient sequestrators, the storage characteristics of
both groups differ considerably. Some insights into
how these insects manipulate the allelochemicals in
Antibiotic Functions of Allelochemicals their diets have been provided by recent studies of
the fates of a variety of ingested plant chemicals. An
The demonstrated range of allelochemicals against analysis of these studies demonstrates that the par-
insect-associated viruses, fungi and bacteria ticulars of sequestration are, if nothing else, very
makes it probable that these arthropods have unpredictable.
A Allelochemicals

The Monarch Butterfly, Danaus plexippus in the gut fluid but high concentrations in the
hemolymph. However, before wing expansion in
The monarch is a specialist that feeds exclusively the newly developed adult, the cardenolide level
on different species of milkweeds. Milkweeds con- in the hemolymph is at its lowest, only to
tain steroids called cardenolides, which are some- increase to the highest level in any life stage.
what related to vertebrate hormones such as The presence of high levels of cardenolides in
testosterone. These compounds are toxic and the blood of the adult demonstrates that these
highly emetic, vomiting often following their compounds are not locked in tissues but rather are
ingestion by non-adapted species. circulating freely, possibly to be utilized upon
Polar (water soluble) cardenolides are seques- demand. The warningly colored (aposematic)
tered in the large volume of gut fluid possessed by adult monarch utilizes a defensive system based
the larvae. Sequestration is much more efficient on compounds that it did not ingest as an adult.
from plants with low level cardenolide concentra- Although the complexities of cardenolide seques-
tions than with high concentrations. Significantly tration in this species are evident, it is highly sig-
for the monarch, and not necessarily for other nificant to understand these ingested steroids are
milkweed feeders, it is the large volume of gut fluid an extraordinarily dynamic state.
that makes it possible to feed and develop on these
The cardenolide-rich gut fluid, which may The Lubber Grasshopper, Romalea
exceed one-third of the larvas total liquid volume, microptera (also known as R. guttata)
is withdrawn at pupation to become part of the
hemolymph (blood) pool, stored primarily under This large grasshopper found in the southeastern
the wings. Subsequently, the wing scales (bird United States is quite conspicuous because of it red,
predators beware), along with the hemolymph, black and yellow coloration. It is one of the most
become the richest sources of cardenolides in the aposematic (warningly colored) species in its habi-
body after being withdrawn from the gut fluid. The tat. This brightly colored grasshopper is especially
volume of gut fluid decreases before pupation only distinctive because it is a generalist that feeds on a
to increase again before pupal molting. Again, gut very wide range of plants belonging to a variety of
fluid diminishes during pupal development only to species. Lubber grasshopper is known to feed on
increase again in the new adult. The cardenolide- 104 plant species belonging to 38 families, many of
rich gut fluid is again converted to hemolymph which produce toxic allelochemicals. Both imma-
during adult development so that very little remains ture and mature grasshoppers are capable of caus-
to be lost when the newly developed adult evacu- ing emesis in predators such as lizards,demonstrating
ates accumulated waste products from its gut. that all stages of these insects are protected from at
The polar cardenolides in the gut fluid least some predatory vertebrates.
clearly are the source of the defensive com- Immature individuals of R. microptera produce
pounds manipulated by the monarch at all defensive compounds that cause emesis in both liz-
stages. The larval and pupal exuviate (cast skins) ard and bird predators. Additionally, mature and
eliminated after molting are an excretory form adult grasshoppers secrete defensive compounds
for the cardenolides, as is the case for these from paired tracheal (respiratory) glands in the
compounds in the wing scales. Excretion not- metathorax. These glands only become active near
withstanding, the ability of all life stages to the adult period and their secretion can be extremely
manipulate the cardenolide pool is quite pro- repellent to small predatory insects such as ants. At
nounced. This is evident in 2-day-old pupae least 50 compounds are produced by the defensive
that contain low concentrations of cardenolides glands, the secretions varying intraspecifically, so
A 133

that components of females of the same age and mode in the presence of a preferred host plant is
population sometimes differ by 70-fold, with some not unreasonable, and can result in a secretion
compounds being absent in certain individuals. with a high a concentration of sequestered alle-
However, in addition to the compounds synthesized lochemicals as is characteristic of some specialist
in the metathoracic glands, a number of allelochem- insects.
icals are sequestered in these glands as a reflection Lubber grasshopper is quite unpalatable and
of an individual grasshoppers diet. Indeed, the emetic to a variety of vertebrates, especially birds.
composition of the metathoracic gland secretion of Diverse bird species have been demonstrated to
each grasshopper appears to be unlike that of any vomit after ingestion of these grasshoppers, pre-
other grasshopper, since no two of these generalist sumably as a consequence of Romalea-synthesized
grasshoppers have identical diets from which to toxins that fortify their bodies. While Romalea
sequester allelochemicals. For a predator, each lub- would appear to be completely defended against
ber secretion may be sufficiently distinctive to make birds, as is often the case, the best defense has been
it impossible to learn an olfactory pattern that overcome by a better offense. Shrikes, predatory
clearly identifies the prey as lubber grasshopper. birds that impale their insect prey on spines or even
Lubber grasshopper is unusual in being a barbed wire, capture lubber grasshoppers and
polyphagous (eating many plant species) insect impale them. However, the birds wait for about 48 h
species that sequesters allelochemicals. In general, before they remove and eat the grasshoppers.
monophagous (feeding on one group of plant spe- Though shrikes store all their food in this manner,
cies) and stenophagous (feeding on a limited range in all likelihood the emetic toxin(s) produced by
of plant species) insect herbivores characteristi- Romalea decomposes during the time the grass-
cally sequester plant compounds, but not general- hopper is impaled.
ist feeders. Furthermore, if R. microptera is
presented with a restricted diet (specialist feeding
mode), the number of compounds in the secre-
tions and their concentrations are reduced, and References
the relative composition of the secretion is mark-
edly different from that of field-collected grass- Blum MS (1981) Chemical defenses of arthropods.
Academic Press, New York, NY
hoppers. Significantly, if grasshoppers are presented Blum MS (1983) Detoxication, deactivation, and utilization
with only a single-host plant as a food source, they of plant compounds by insects. In: Hedin P (ed), Plant
frequently feed readily, sequestering host-plant resistance to insects. American Chemical Society,
Washington, DC, pp 265275
volatiles, and exhibit no immediate ill effects. Lub-
Bowers MD (1990) Recycling plant natural products for
bers feeding only on wild onion sequester a large insect defense. In: Evans DL, Schmidt JO (eds), Insect
number of onion volatiles which impart a strong defenses. Adaptive mechanisms and strategies of prey
onion odor to the secretion. The secretion is a and predators. State University of New York, Albany, NY,
pp 353386
powerful repellent to hungry ants and is consider- Evans DL, Schmidt JO (eds) Insect defenses. Adaptive mecha-
ably more active than the secretions of field- nisms and strategies of prey and predators. State Uni-
collected grasshoppers. Compounds in other versity of New York, Albany, NY, 482 pp
single-plant diets (e.g., catnip) produce secretions Gibson RW, Pickett JA (1983) Wild potato repels aphids by
release of aphid alarm pheromone. Nature 302:608609
that are similarly active. Pasteels JM, Gregoire JC, Rowell-Rahier M (1983) The chemical
The secretion of lubber grasshopper clearly ecology of defense in arthropods. Annu Rev Entomol
has both a dietary and an individual origin that 28:263289
Whitman DW (1988) Allelochemical interactions among
correlates with great variations in secretory com-
plants, herbivores, and their predators. In: Barbosa P,
ponents. The possibility that these grasshoppers Letourneau D (eds), Novel aspects of insect-plant inter-
can temporarily switch to a monophagous feeding actions. Wiley, New York, NY, pp 1164
A Allelopathy

Allelopathy Allomone
The ability of a plant species to produce substances A chemical that is released by one species that
that are toxic to certain other plants. Allelopathic influences the behavior or physiology of a differ-
chemicals may affect germination, growth or ent species. The organism releasing the substance
reproduction of plants. usually benefits. Allomones are a type of
semiochemical used in warning.
Chemical Ecology
Allens Rule
Among mammals and birds, individuals of a spe- Allopatric
cies occurring in colder climates tend to have
shorter appendages, and a correspondingly lower Having separate and mutually exclusive areas of
surface to volume ratio, than members of the same distribution (contrast with sympatric).
species living in warmer climates. This trend
results from the need to conserve heat in cold cli-
mates but to eliminate excess heat in hot climates. Allopatric Speciation
A variant of this is Bergmanns rule. These rules do
not apply to ectothermic animals such as insects. A mechanism of speciation resulting from geo-
Bergmanns Rule graphic separation of populations, particularly
Thermoregulation physical barriers such as mountains and oceans.
Speciation Processes Among Insects

Allochronic Speciation
Allophagic Speciation
A mechanism of speciation wherein new species
develop in the same place but are separated due to A mechanism of speciation wherein new species
their tendency to occur at different times. develop in the same place, but are separated by
Speciation Processes Among Insects their preference for different food.
Speciation Processes Among Insects
Allogenic Succession
A temporal succession of species that is driven by
processes from outside the community (contrast
Allozymes are a subset of isozymes. Allozymes are
with autogenic succession).
variants of enzymes representing different allelic
alternatives of the same locus.
Allometric Growth
A growth pattern in which different parts of an Almond Seed Wasp, Eurytoma
organism grow at defined rates. In some cases, the amygdali Enderlein
body parts remain proportional (isometric (Hymenoptera: Eurytomidae)
growth), in other cases they do not. Departure
from isometric growth is used to explain castes of nikos a. kouloussis
social insects, which may have disproportionately Aristotle University of Thessaloniki, Thessaloniki,
large heads, mandibles, etc. Greece
Almond Seed Wasp, Eurytoma Amygdali Enderlein (Hymenoptera: Eurytomidae)
A 135

The almond seed wasp is a serious pest of almonds, the translucent nucellar tissue. After oviposition, the
Prunus amygdalus Batch, in several countries of female deposits onto the fruit surface a host-marking
southeastern Europe and the Middle East, and pheromone. This pheromone enables females to dis-
also in Armenia, Azerbaijan, and Georgia. The criminate between the infested and uninfested fruit,
adult female is 68 mm long and has a black head and to select the latter for oviposition. Thus, a uni-
with dark brown eyes. The thorax and the spindle- form distribution of eggs among available fruits is
shaped abdomen are shiny black. The tibiae and achieved, and an optimal use of the available fruit for
tarsi are light brown while the remaining parts of larval development. The newly hatched larva bores
the leg are black. The male is usually smaller than through the nucellus and the embryo sac to feed on
the female (46 mm long). The larva is whitish, the developing seed embryo. The larva attains full size
legless, tapering in both ends, curved and clearly in midsummer, and enters diapause within the seed
segmented. Its head is light brown and very small. integument of the destroyed almond, which usually
Its length when fully grown is about 6 mm. remains on the tree in a mummified condition.
Almond seed wasp is a univoltine species, with Owing to oviposition by the wasp certain
a small part of the population completing its life varieties suffer a heavy premature drop. In most
cycle in two or more years because of prolonged dia- varieties though, the main damage consists in
pause. The diapause terminates during the winter. theconsumption of the seed by the larvae. This
Pupation takes place inside the fruit in late winter to damage varies depending on the variety. Certain
early spring. Adults emerge after boring a circular soft-shelled varieties may lose up to 90% of their
exit hole through the hard pericarp with their man- crop. Others are nearly immune because by the
dibles. Shortly after adult emergence, virgin females time females emerge in spring their pericarp has
release a volatile sex pheromone to attract males for become too thick and endocarp too hard for the
mating. Within a few days they mate and the females ovipositor to penetrate. Though not a common
start ovipositing (Fig.37) into unripe, green almonds. practice, planting of resistant varieties might be
Using her long ovipositor, the female drills through an effective strategy against this pest.
the pericarp of unripe, green almonds and the integ- The pest can be controlled by collection
ument of the seed, and deposits a stalked egg within anddestruction of mummified fruits before adult

Almond Seed Wasp, Eurytoma Amygdali Enderlein (Hymenoptera: Eurytomidae), Figure 37 Female
Eurytoma amygdali ovipositing into an almond.
A Alpha Taxonomy

emergence in spring, which is an effective measure Beta Taxonomy

if applied in large areas by multiple growers. How- Gamma Taxonomy
ever, the method most commonly used is the appli-
cation of systemic insecticides against the neonate
larvae within the oviposited almonds. This strategy Alternate Host
is meant for varieties that do not suffer fruit drop
because of oviposition. Recent studies have indi- One of the hosts of a pathogen or insect where a
cated that a single spraying can be effective if applied portion of the life cycle occurs. Often this term is
when 1050% of the eggs have hatched. This per- used to refer to a weed host of a crop pest or
centage can be determined by dissecting sampled disease.
almonds under a binocular microscope. Estimates
of egg hatch can be obtained by knowing the time Alternation of Generations
the first adults emerge from infested almonds in
spring. This can be determined by following the exit Some insects undergo reproduction that involves
of adult wasps from infested almonds kept in cages alternation of sexual and asexual generations. Typ-
in the orchard, or by following the population of ically, females produce both males and females but
males with the use of sex pheromone traps contain- at some point females cease producing males and
ing live virgin females as lures. produce only females parthenogenetically. Later
generations then commence production of males
again, allowing sexual reproduction to occur before
References the parthenogenetic cycle begins again. This occurs
most often in Hymenoptera and Hemiptera.
Katsoyannos BI, Kouloussis NA, Bassiliou A (1992) Monitor- Aphids (Hemiptera: Aphididae)
ing populations of the almond seed wasp, Eurytoma Gall Wasps (Hymenoptera: Cynipidae)
amygdali, with sex pheromone traps and other means,
and optimal time of chemical control. Entomologia
Experimentalis et Applicata 62:916
Kouloussis NA, Katsoyannos BI (1991) Host discrimination
and evidence for a host marking pheromone in Eury-
toma amygdali. Entomologia Experimentalis et Appli- Self destructive behavior that is performed for the
cata 58:165174 benefit of others; sacrifice.
Plaut HN (1971) On the biology of the adult of the almond
seed wasp, Eurytoma amygdali End. (Hym., Eurytomi-
dae) in Israel. Bull Entomol Res 61:275281
Plaut HN (1972) On the biology of the immature stages of the Alucitidae
almond wasp, Eurytoma amygdali End. (Hym., Euryto-
midae) in Israel. Bull Entomol Res 61:681687 A family of moths (order Lepidoptera). They com-
Tzanakakis ME, Papadopoulos NT, Katsoyannos BI, Drakos
monly are known as many-plumed moths.
GN, Manolakis E. Premature fruit drop caused by Eury-
toma amygdali (Hymenoptera: Eurytomidae) on three Many-Plumed Moths
almond varieties. J Econ Entomol 90:16351640 Butterflies and Moths

Alpha Taxonomy Alula

The identification of organisms, and particularly The expanded membrane at the base of the trail-
the description and naming of organisms (species) ing edge of the front wing.
new to science. Wings Of Insects
Amber Insects: DNA Preserved?
A 137

Amazonian Primitive Ghost vertebrates that got stuck in the sticky exudate.
Moths (Lepidoptera: The hardened resin was preserved in the earth for
Neotheoridae) millions of years, especially in regions where it
was deposited in dense, wet sediments such as clay
john b. heppner or sand that formed in the bottom of an ancient
Florida State Collection of Arthropods, Gaines- lagoon or river delta. For thousands of years, peo-
ville, FL, USA ple have collected amber. Many people use amber
as a gem, but scientists find amber a magnificent
Amazonian primitive ghost moths, family way to identify ancient organisms.
Neotheoridae, are defined on the basis of a single Amber can be found in a variety of sites
species from the Amazonian area of southern around the world. The composition, color, clar-
Brazil, although two additional species have been ity, and other properties of amber vary accord-
discovered for the family recently. The family is ing to age, conditions of burial and type of tree
part of the superfamily Hepialoidea, in the that produced the resin. The oldest amber is
infraorder Exoporia. Adults medium size (38 mm from the Carboniferous (360285 million years
wingspan), with head roughened; haustellum ago, mya) and can be found in the United King-
short and vestigial mandibles present; labial palpi dom and in Montana in the USA. Permian
long, porrect and 3-segmented; maxillary palpi amber is 185145 million years old and found
very small and 2-segmented. Wing maculation is most often in Russia. Triassic amber (245215
dark and unicolorous. Biologies and larvae remain mya) can be found in Austria, and Jurassic
unknown. amber (215145 mya) is found in Denmark.
Cretaceous amber (65140 mya) is found in
many locations around the world and represents
References the time when dinosaurs reigned and flowering
plants evolved along with a variety of insects.
Kristensen NP (1978) A new familia of Hepialoidea from For example, the rich amber deposits in central
South America, with remarks on the phylogeny of New Jersey in the USA are from the Turonion
the subordo Exoporia (Lepidoptera). Entomologia
Germanica 4:272294
period of the Upper Cretaceous, about 92 mya.
Kristensen NP (1999) The homoneurons Glossata. In: Other Cretaceous-period amber is found in
Kristensen NP (ed), Lepidoptera, moths and butterflies, North Russia and Japan. Baltic amber is found
vol 1: evolution, systematics, and biogeography. Handbuch in the Baltic sea where amber has been collected
der Zoologie. Band IV. Arthropoda: Insecta. Teilband
35:5163. W. de Gruyten, Berlin and made into decorative objects for at least
13,000 years. In the Dominican Republic, amber
deposits 2330 million years old are found in
rock layers. Dominican amber is particularly
Amber Insects: DNA Preserved? rich in insect inclusions. This amber was formed
from the resin of an extinct tree in the legume
marjorie a. hoy family. Tertiary amber deposits are found in
University of Florida, Gainesville, FL, USA several locations around the world and are from
1.6 to 65 million years old. Tertiary deposits in
Amber is a polymerized form of tree resin that the USA are found in Arkansas. Some websites
was produced by trees as a protection against dis- with photographs showing amber inclusions
ease agents and insect pests. The resin hardened can be viewed at:
and, sometimes, captured insects, seeds, feathers, Amber Inclusions at: http://www-user.uni-bremen.
microorganisms, plants, spiders, and even small de/~18m/amber.html;
A Amber Insects: DNA Preserved?

Amber on-line at: amber-preserved insects has yet to be reproduced

html; in independent laboratories, despite multiple
American Museum of Natural History at: www.amnh. attempts to do so, which has cast doubt on the
org/exhibitions/amber/; or authenticity of the reports.
The Amber Room at: One of the most controversial claims involved
AMBER.HTM. the isolation of a living bacterium from the abdo-
men of amber-entombed bee. Bacterial DNA from
a 25-million-year-old bee was obtained and
Insect DNA in Amber? sequenced and a bacterial spore was reported to
be revived, cultured, and identified. The classifica-
The ability to amplify dinosaur DNA from insects tion of the bacterium is controversial because the
preserved in amber in the film Jurassic Park cap- bacterium could have come from a currently
tured the imagination of the public. Subsequently, undescribed species of the Bacillus sphaericus
the PCR was used to amplify DNA fragments from complex. The modern B. sphaericus complex is
insects preserved in ancient amber, but these incompletely known, so the new sequence
results have been controversial, as have been the obtained could be that of a modern, but previously
results from amplifying dinosaur DNA. unidentified, bacterium because this group of bac-
Why the controversy? Is amber a special form teria often is isolated from the soil.
of preservative that allows DNA to persist for Other claims of amplifying ancient DNA have
unusually long periods of time (millions of years)? been disproved. For example, the mitochondrial
Amber entombs insect specimens completely, cytochrome b sequence of an 80-million-year-old
after which they completely dehydrate so the tis- dinosaur from the Upper Cretaceous in Utah was
sue is effectively mummified. Terpenoids, which later discovered to be, most probably, of human
are major constituents of amber, could inhibit origin. Likewise, a 20-million-year-old magnolia
microbial decay. Certainly, preservation of amber- leaf produced sequences that were similar to those
embedded insects seems to be exceptional and of modern magnolias. The authenticity of the
insect tissues in amber appear comparable in qual- magnolia sequences were cast into doubt because
ity to the tissues of the frozen wooly mammoth they were exposed to water and oxygen during
(which is only 50,000 years old). But is the DNA preservation and DNA is especially vulnerable to
in these tissues preserved? degradation under such conditions.
DNA has been extracted from a variety of The most common ancient DNA analyzed is
insects in amber, including a fossil termite Masto- usually mitochondrial DNA because it is so abun-
termes electrodominicus estimated to be 2530 dant; however, this abundance makes it easy to
million years old, a 120- to 130-million year old contaminate the ancient sample with modern
conifer-feeding weevil (Coleoptera: Nemonychi- mtDNA. The amplification of ancient DNA
dae) and a 25- to 40-million year old bee. These remains highly controversial because the technical
are extraordinary ages for DNA! difficulties are great.
The DNA sequences obtained from all amber- DNA is a chemically unstable molecule that
preserved insects meet several, but not all, criteria decays spontaneously, mainly through hydroly-
of authenticity; the fossil DNA sequences make sis and oxidation. Hydrolysis causes deamina-
phylogenetic sense and DNA has been isolated tion of the nucleotide bases and cleavage of
from a number of specimens in several cases base-sugar bonds, creating baseless sites. Deami-
(although the weevil example was derived from a nation of cytosine to uracil and depurination
single specimen). However, the extraction and (loss of purines adenine and guanine) are two
amplification of fossil DNA sequences from types of hydrolytic damage. Baseless sites weaken
Ambrosia Beetles
A 139

the DNA, causing breaks that fragment the DNA selection of well-preserved specimens, choice of
into smaller and smaller pieces. Oxidation leads tissue samples that are likely to have best DNA
to chemical modification of bases and destruc- preservation, and surface sterilization to elimi-
tion of the ring structure of base and sugar resi- nate surface contamination. The operations
dues. As a result, it is almost always impossible should be carried out in a laboratory dedicated to
to obtain long amplification products from work on ancient specimens and work on ancient
ancient DNA. DNA should be separated from that on modern
PCR products from ancient DNA often are DNA. Most importantly, multiple negative con-
scrambled. This is due to the phenomenon trols should be performed during DNA extrac-
called jumping PCR, which occurs when the tion and PCR set up, although a lack of positives
DNA polymerase reaches a template position in the negative controls is not definitive proof of
which carries either a lesion or a strand break authentic ancient DNA. Another crucial step is
that stops the polymerase. The partially extended the authentication of the results. Putatively ancient
primer can anneal to another template fragment DNA sequences should be obtained from differ-
in the next cycle and be extended up to another ent extractions of the same sample and from dif-
damaged site. Thus, in vitro recombination can ferent tissue samples from different specimens.
take place until the whole stretch encompassed The ultimate test of authenticity should be inde-
by the two primers is synthesized and the ampli- pendent replication in two separate laboratories.
fication enters the exponential part of the PCR. So far, this type of replication has not been
This phenomenon makes it essential that clon- achieved for DNA from amber-preserved arthro-
ing and sequencing of multiple clones be pod specimens.
carried out to eliminate this form of error in
Most archeological and paleontological speci- References
mens contain DNA from exogenous sources such
as bacteria and fungi, as well as contaminating Austin JJ, Ross AJ, Smith AB, Fortey RA, Thomas RH (1997)
DNA from contemporary humans. Aspects of Problems of reproducibility does geologically ancient
burial conditions seem to be important in DNA DNA survive in amber-preserved insects? Proc R Ento-
mol Soc London B 264:467474
preservation, especially low temperature during Hofreiter M, Serre D, Poinar HN, Kuch M, Paabo S (2001) Ancient
burial. The oldest DNA sequences reported, and DNA. Nat Rev Genet 2:353359
confirmed in other laboratories, come from the Poinar G Jr, Poinar R (2001) The amber forest: a reconstruc-
remains of a wooly mammoth found in the Sibe- tion of a vanished world. Princeton University Press,
Princeton, NJ
rian permafrost; these sequences are only 50,000 Poinar HN, Stankiewicz BA (1999) Protein preservation and
years old rather than millions of years old. DNA retrieval from ancient tissues. Proc Natl Acad Sci
Theoretical calculations and empirical USA 96:84268431
Yousten AA, Rippere KE (1997) DNA similarity analysis of a
observations suggest DNA should only be able
putative ancient bacterial isolate obtained from amber.
to survive, in a highly fragmented and chemi- FEMS Microbiol Lett 152:345347
cally modified form, for 50,000100,000 years.
Because only tiny amounts of DNA usually can
be extracted from an archeological specimen,
stringent precautions and multiple controls are Ambrosia Beetles
required to avoid accidental contamination with
modern DNA. Some members of the subfamily Scolytinae (order
A methodology to deal with ancient speci- Coleoptera, family Curculionidae).
mens has been proposed that includes careful Beetles
A Ambush Bugs

Ambush Bugs Reference

Members of the family Reduviidae (order Konradsen F, de Silva A, van der Hoek W (2005) Felix P.
Hemiptera). Amerasinghe. American Entomologist 51:191

American Butterfly Moths

(Lepidoptera: Hedylidae)
john b. heppner
A family of mayflies (order Ephemeroptera). Florida State Collection of Arthropods,
Mayflies Gainesville, FL, USA

American butterfly moths, family Hedylidae, total

Amelitidae only 40 known species, all Neotropical. The family
is in the superfamily Geometroidea, in the section
A family of mayflies (order Ephemeroptera). Cossina, subsection Bombycina, of the division
Mayflies Ditrysia. Adults medium size (3565 mm wing-
span), with head scaling normal; haustellum
naked; labial palpi upcurved; maxillary palpi 1 to
2-segmented; antennae filiform. Wings triangular,
Amerasinghe, Felix P with forewings somewhat elongated and often
with apex emarginated (Fig.38); hindwings usually
Felix Amerasinghe was a noted Sri Lankan medi- more rounded. Body usually narrow. Maculation
cal entomologist. He was known for his work on somber hues of brown and gray, often with apical
the taxonomy and ecology of disease-transmitting dark patch and some speckling, plus pale or hya-
arthropods. Amerasinghe graduated from the line patches (rarely mostly pale or hyaline). Adults
University of Peradeniya, Sri Lanka, and received nocturnal. Larvae are leaf feeders. Host plants are
his Ph.D. from the University of Bristol, United recorded in Euphorbiaceae, Malvaceae, Sterculi-
Kingdom, in 1977. He made important long-term aceae, and Tiliaceae.
studies in the effects of irrigation on mosquito
populations and malaria transmission, and became
an authority on Japanese encephalitis. The devel-
opment of keys for the identification of South
Asian mosquitoes was one of his important con-
tributions, greatly enhancing disease surveillance
Amerasinghe worked principally at the Uni-
versity of Peradeniya, but also at the University of
Sri Lanka, and in later years joined the Interna-
tional Water Management Institute as research
leader, initiating studies on the socioeconomic American Butterfly Moths (Lepidoptera:
impact of malaria, malaria parasitology, and Hedylidae), Figure 38 Example of American
molecular biology. He died in Colombo, Sri Lanka, butterfly moths (Hedylidae), Macrosoma lucivittata
on June 7, 2005. (Walker), from Ecuador.
American Grasshopper, Schistocerca Americana (Drury) (Orthoptera: Acrididae)
A 141


Aiello A (1992) Nocturnal butterflies in Panama, Hedylidea

(Lepidoptera: Rhopalocere). Quintero D, Aiello A (eds),
Insects of Panama and Mesoamerica. Oxford University
Press, Oxford, pp 549553
Scoble MJ (1986) The structures and affinitxies of the Hedy-
loidea: a new concept of the butterflies. Bull Br Mus Nat
Hist Entomol 53:251286
Scoble MJ (1990) An identification guide to the Hedylidae
(Lepidoptera: Hedyloidea). Entomologica Scandinavica American False Tiger Moths (Lepidoptera:
21:121158 Dioptidae), Figure 39 Example of American false
Scoble MJ (1998) Hedylidae. In Lepidopterorum Catalogus, tiger moths (Dioptidae), Josia gigantea Druce,
(n.s.). Fasc. 93. Association for Tropical Lepidoptera,
Gainesville, FL, 9 pp from Mexico.
Scoble MJ, Aiello A (1990) Moth-like butterflies (Hedylidae:
Lepidoptera): a summary, with comments on the egg. J References
Nat Hist 24:159164
Bryk F (1930) Dioptidae. Lepidopterorum catalogus, 42:165.
W. Junk, Berlin, Germany
Miller JS (1987) A revision of the genus Phryganidia Packard,
American Dog Tick with description of a new species (Lepidoptera: Diopti-
dae). Proc Entomol Soc Wash 89:303321
Ticks Prout LB (1918) A provisional arrangement of the Dioptidae.
Novitates Zoologicae 25:395429
Seitz A (ed) (19251927) Familie: Dioptidae. In: Die Gross-
Schmetterlinge der Erde. 6. Die amerikanischen Spinner
American False Tiger Moths und Schwrmer, pl. 6771. A. Kernen, Stuttgart,
(Lepidoptera: Dioptidae) Germany, pp 499534
Todd EL (1981) The noctuoid moths of the Antilles Part I
(Lepidoptera: Dioptidae). Proc Entomol Soc Wash
john b. heppner 83:324325
Florida State Collection of Arthropods,
Gainesville, FLa, USA

American false tiger moths, family Dioptidae, total American Grasshopper,

507 species, primarily Neotropical (505 sp.); actual Schistocerca americana (Drury)
fauna likely exceeds 800 species. Two subfamilies are (Orthoptera: Acrididae)
known: Dioptinae and Doinae. Some specialists
place the family within the Notodontidae. The fam- john l. capinera
ily is in the superfamily Noctuoidea, in the section University of Florida, Gainesville, FL, USA
Cossina, subsection Bombycina, of the division
Ditrysia. Adults medium size (2258 mm wingspan) This grasshopper is found widely in eastern North
(Fig.39). Maculation mostly very colorful, with var- America, from southern Canada (where it is an
ious patterns of large spotting, and some lustrous. occasional invader) south through Mexico to
Larvae and pupae often also colorful Adults are northern South America. In the midwestern states,
mostly nocturnal, but some are diurnal or crepuscu- where it is common, the resident population
lar. Larvae are leaf feeders, particularly toxic plants receives a regular infusion of dispersants from
in families like Aristolochiaceae, Euphorbiaceae, southern locations. In the southeast it is quite
Passifloraceae, and Violaceae, but also on various common, and one of the few species to reach epi-
others like Fagaceae. Very few are economic. demic densities. It is native to North America.
A American Grasshopper, Schistocerca Americana (Drury) (Orthoptera: Acrididae)

Life History than ventral orientation but the wing tip does not
exceed the first abdominal segment. In the sixth
In warm climates, American grasshopper has two instar (Fig.41) there are 2426 antennal segments
generations per year and overwinters in the adult and the wing tips extend beyond the second
stage. In Florida, eggs produced by overwintered abdominal segment. The overall body length is
adults begin to hatch in April-May, producing about 67, 1213, 1618, 2225, 2730, and 3545
spring generation adults by May-June. This spring mm for instars 16, respectively. Development
generation produces eggs that hatch in August- time is about 46, 46, 46, 48, 68, and 913
September. The adults from this autumn genera- days for the corresponding instars when reared at
tion survive the winter. about 32C.
The eggs of S. americana initially are light The adult (Fig.40) is rather large, but slender
orange in color, turning tan with maturity. They bodied, measuring 3952 and 4868 mm in length
are elongate-spherical in shape, widest near the in the male and female, respectively. A creamy
middle, and measure about 7.5 mm in length and white stripe normally occurs dorsally from the
2.0 mm in width. The eggs are clustered together front of the head to the tips of the forewings. The
in a whorled arrangement, and number 75100 forewings bear dark brown spots, the pronotum
eggs per pod, averaging 85 eggs. The eggs are
inserted into the soil to a depth of about 4 cm and
the upper portion of the oviposition hole is filled
by the female with a frothy plug. Duration of the
egg stage is about 14 days. The nymphs, upon
hatching, dig through the froth to attain the soil
Normally there are six instars in this grass-
hopper though sometimes only five. The young
grasshoppers are light green in color. They are
extremely gregarious during the early instars. At
low densities the nymphs remain green through-
out their development, but normally gain increas- American Grasshopper, Schistocerca Americana
ing amounts of black, yellow, and orange coloration (Drury) (Orthoptera: Acrididae), Figure 40 Adult
commencing with the third instar. Instars can be of American grasshopper, Schistocerca americana
distinguished by their antennal, pronotal, and (Drury).
wing development. The first and second instars
display little wing development but have 13 and
17 antennal segments, respectively. In the third
instar, the number of antennal segments increases
to 2022, the wings begin to display weak evi-
dence of veins, and the dorsal length of the ven-
tral lobe of the pronotum is about 1.5 times the
length of the ventral surface. Instar four is quite
similar to instar three, with 2225 antennal seg-
ments, though the ratio of the length of the dorsal American Grasshopper, Schistocerca Americana
to ventral surfaces of the pronotal lateral lobe is (Drury) (Orthoptera: Acrididae), Figure 41 Sixth
2:1. In instar five there are 2425 antennal seg- instar of American grasshopper, Schistocerca
ments, and the wing tips assume a dorsal rather americana (Drury).
American Grasshopper, Schistocerca Americana (Drury) (Orthoptera: Acrididae)
A 143

dark stripes. The hind wings are nearly colorless. Damage

The hind tibiae normally are reddish. Overall, the
body color is yellowish brown or brownish with Grasshoppers are defoliators, eating irregular
irregular lighter and darker areas, though for a holes in leaf tissue. Under high density conditions
week or so after assuming the adult stage a pinkish they can strip vegetation of leaves, but more com-
or reddish tint isevident. monly leave plants with a ragged appearance.
Adults are active, flying freely and sometimes American grasshopper displays a tendency to
in swarms. They normally are found in sunny swarm, and the high densities of grasshoppers can
areas, but during the warmest portions of the day cause severe defoliation.
will move to shade. Adults are long lived, persist- Because American grasshopper is a strong
ing for months in the laboratory and apparently in flier, it also sometimes becomes a contaminant
the field as well. This can lead to early-season situ- of crops. When the late-season crop of collards
ations where overwintered adults, all instars of in the Southeast is harvested mechanically, for
nymphs, and new adults are present simultane- example, American grasshopper may become
ously. Mild winters favor survival of overwintering incorporated into the processed vegetables.
adults and apparently lead to population increase Although most grasshoppers can be kept from
if summer weather and food supplies also are dispersing into crops near harvest by treating
favorable. the periphery of the crop field, it is much more
Adults of American grasshopper tend to be difficult to prevent invasion by American grass-
arboreal in habit, and a great deal of the feeding hopper because it may fly over any such barrier
by adults occurs on forest, shade, and fruit trees. treatments.
The nymphs, however, feed on a large number of Populations normally originate in weedy
grasses and broadleaf plants, both wild and cul- areas such as fence rows and abandoned fields.
tivated. During periods of abundance, almost no Thus, margins of fields are first affected and this is
plants are immune to attack, and vegetables, where monitoring should be concentrated. It is
grain crops, and ornamental plants are injured. highly advisable to survey weedy areas in addition
American grasshopper consumes bean, corn, to crop margins if grasshoppers are found, as this
okra, and yellow squash over some other vegeta- gives an estimate of the potential impact if the
bles when provided with choices, but free-flying grasshoppers disperse into the crop. Also, it is
adults normally avoid low-growing crops such important to recognize that this species is highly
as vegetables, corn (maize) being a notable dispersive in the adult stage, and will fly hundreds
exception. of meters or more to feed.
The natural enemies of S. americana are not
well known. Birds such as mockingbirds, Mimus
polyglottos polyglottos (Linnaeus), and crows, Cor- Management
vus brachyrhynchos brachyrhynchos Brehm have
been observed to feed on these grasshoppers. Fly Foliar applications of insecticides will suppress
larvae, Sarcophaga sp. (Diptera: Sarcophagidae) grasshoppers, but they are difficult to kill, particu-
are sometimes parasitic on overwintering adults. larly as they mature. Bait formulations are not
Fungi have also been investigated for grasshopper usually recommended because these grasshoppers
suppression and Metarhizium anisopliae var. spend little time on the soil surface, preferring to
acridum kills American grasshopper quickly under climb high in vegetation.
laboratory conditions. This fungus is effective Land management is an important element
under adverse field conditions in Africa, so it may of S. americana population regulation. Grass-
prove to be a useful suppression tool. hopper densities tend to increase in large patches
A American Foulbrood

of weedy vegetation that follow the cessation of (= Bacillus) larvae, form heat- and drought-
agriculture or the initiation of pine tree planta- resistant spores that persist for years and germi-
tions. In both cases, the mixture of annual and nate under favorable conditions. It is expressed in
perennial forbs and grasses growing in fields older larvae and young pupae, though infection
that are untilled seems to favor grasshopper occurs earlier, and young larvae are more suscep-
survival, with the grasshoppers then dispersing tible than older larvae. Infected individuals turn
to adjacent fields as the most suitable plants are darker in color, then black, and eventually collapse
depleted. However, as abandoned fields convert into a hardened mass in the cell. Signs of infection
to dense woods or the canopy of pine planta- include a sour odor, perforated or sunken caps on
tions shades the ground and suppresses weeds, the cells, and the presence of black deposits in the
the suitability of the habitat declines for cells. If foulbrood is present, insertion of a twig or
grasshoppers. probe into a suspect cell will result in a gummy,
Disturbance or maturation of crops may cause stretchy substance being drawn out of the cell,
American grasshopper to disperse, sometimes over often forming a thread or rope and called ropy.
long distances, into crop fields. Therefore, care Field diagnosis is possible by experienced inspec-
should be taken not to cut vegetation or till the soil tors, but is best confirmed microscopically or by
of fields harboring grasshoppers if a susceptible molecular techniques. There are several subspe-
crop is nearby. Planting crops in large blocks cies of P. larvae, and P. larvae ssp. larvae is consid-
reduces the relative amount of crop edge, and the ered responsible for American foulbrood, with
probability that a crop plant within the field will be other subspecies also affecting honey bees.
attacked. Transmission occurs by feeding infected
Grasshopper Pests in North America honey or pollen, by using infected equipment, and
Grasshoppers and Locusts as Agricultural Pests sometimes by installing infected package bees or
Grasshoppers, Katydids and Crickets queens. Feeding bees sugar syrup therefore is pref-
(Orthoptera) erable to feeding them honey, and disinfection of
hive tools is always recommended. Natural trans-
References mission from hive to hive can occur through rob-
bing behavior. Queens and workers can carry the
Capinera JL (1993) Differentiation of nymphal instars in disease. Bee colonies that are infected normally
Schistocerca americana (Orthoptera: Acrididae). Fla are eliminated by burning them. Antibiotics can
Entomol 76:175179 be fed to colonies to prevent infection.
Capinera JL (1993) Host-plant selection by Schistocerca amer-
Honey Bees
icana (Orthoptera: Acrididae). Environ Entomol
22:127133 Apiculture
Capinera JL, Scott RD, Walker TJ (2004) Field guide to the Paenibacillus
grasshoppers, katydids, and crickets of the United States.
Cornell University Press, Ithaca, NY, 249 pp
Kuitert LC, Connin RV (1952) Biology of the American
grasshopper in the southeastern United States. Fla
Entomol 35:2233

Alippi AM, Lpez AC, Aguilar OM (2002) Differentiation

ofPaenibacillus larvae subsp. larvae, the cause of Amer-
American Foulbrood ican foulbrood of honeybees, by using PCR and restric-
tion fragment analysis of genes encoding 16S rRNA.
Appl Environ Microbiol 68:36553660
Historically, this is the most virulent disease
Morse RA, Nowogrodzki R (1990) Honey bee pests, preda-
of honey bees throughout the world. The bacte- tors and diseases, 2nd ed. Cornell University Press,
rium responsible for the disease, Paenibacillus Ithaca, NY, 474 pp
American Serpentine Leafminer, Liriomyza Trifolii (Burgess) (Diptera: Agromyzidae)
A 145

American Serpentine Leafminer, measuring about 1.0 mm long and 0.2 mm wide.
Liriomyza trifolii (Burgess) Initially they are clear, but soon become creamy
(Diptera: Agromyzidae) white in color.

This leafminer has long been found in eastern

North America, northern South America, and the Larva
Caribbean. However, in recent years it has been
introduced into California, Europe, and elsewhere. Body and mouth part size can be used to differenti-
Expanded traffic in flower crops appears to be the ate instars; the latter is particularly useful. For the
basis for the expanding range of this species. Liri- first instar, the mean and range of body and mouth
omyza trifolii (Burgess), sometimes known as the parts (cephalopharyngeal skeleton) lengths are
American serpentine leafminer, readily infests 0.39 (0.330.53) mm and 0.10 (0.080.11) mm,
greenhouses. As a vegetable pest, however, its respectively. For the second instar, the body and
occurrence is limited principally to tropical and mouth parts measurements are 1.00 (0.551.21)
subtropical regions. mm and 0.17 (0.150.18) mm, respectively. For the
third instar, the body and mouth parts measure-
ments are 1.99 (1.262.62) mm and 0.25 (0.22
Life Cycle and Description 0.31) mm, respectively. A fourth instar occurs
between puparium formation and pupation, but
Leafminers have a relatively short life cycle. The this is a nonfeeding stage and is usually ignored by
time required for a complete life cycle in warm authors. The puparium is initially golden brown in
environments is often 2128 days, so numerous color, but turns darker brown with time.
generations can occur annually in tropical cli-
mates. Growth at a constant 25C requires about
19 days from egg deposition to emergence of the Adult
adult. Development rates increase with tempera-
ture up to about 30C; temperatures above 30C Adults (Fig. 42) are small, measuring less than 2 mm
are usually unfavorable and larvae experience high in length, with a wing length of 1.251.9 mm. The
mortality. At 25C, the egg stage requires 2.7 days head is yellow with red eyes. The thorax and abdo-
for development; the three active larval instars men are mostly gray and black although the ven-
require an average of 1.4, 1.4, and 1.8 days, respec- tral surface and legs are yellow. The wings are
tively; and the time spent in the puparium is 9.3 transparent. Key characters that serve to differen-
days. Also, there is an adult preovipostion period tiate this species from the vegetable leafminer,
that averages 1.3 days. The temperature threshold Liriomyza sativae Blanchard, are the matte, grayish
for development of the various stages is 610C, black mesonotum and the yellow hind margins of
except that egg laying requires about 12 C. the eyes. In vegetable leafminer the mesonotum is
shining black and the hind margin of the eyes is
black. The small size of this species serves to dis-
Egg tinguish it from pea leafminer, Liriomyza huidob-
rensis (Blanchard), which has a wing length of
Eggs tend to be deposited in the middle of the 1.72.25 mm. Also, the yellow femora of Ameri-
plant; the adult seems to avoid immature leaves. can serpentine leafminer help to separate it from
The female deposits the eggs on the lower surface pea leafminer, which has darker femora. Oviposi-
of the leaf, but they are inserted just below the epi- tion occurs at a rate of 3539 eggs per day, for a
dermis. Eggs are oval in shape and small in size, total fecundity of200400 eggs. The female makes
A American Serpentine Leafminer, Liriomyza Trifolii (Burgess) (Diptera: Agromyzidae)

American Serpentine Leafminer, Liriomyza Trifolii (Burgess) (Diptera: Agromyzidae), Figure 42 Adult of
American serpentine leafminer, Liriomyza trifolii.

numerous punctures of the leaf mesophyll with crops that are readily infested and which are
her ovipositor, and uses these punctures for feed- known to facilitate spread of this pest include
ing and egg laying. The proportion of punctures chrysanthemum, gerbera, gypsophila, and mari-
receiving an egg is about 25% in chrysanthemum gold, but there are likely many other hosts, espe-
and celery, both favored hosts, but only about 10% cially among the Compositae. Numerous
in tomato, which is less suitable for larval survival broad-leaved weed species support larval growth.
and adult longevity. Although the female appar- The nightshade Solanum americanum, Spanish
ently feeds on the exuding sap at all wounds, she needles, Bidens alba, and pilewort, Erechtites hier-
spends less time feeding on unfavorable hosts. The acifolia, were suitable weed hosts in Florida.
males live only two to three days, possibly because
they cannot puncture foliage and therefore feed
less than females, whereas females usually survive Damage
for about a week. Typically they feed and oviposit
during much of the daylight hours, but especially Punctures caused by females during the feeding
near mid-day. and oviposition processes can result in a stippled
appearance on foliage, especially at the leaf tip
and along the leaf margins. However, the major
Host Plants form of damage is the mining of leaves by larvae,
which results in destruction of leaf mesophyll.
Liriomyza trifolii is perhaps best known as a pest The mine becomes noticeable about three to four
of chrysanthemums and celery, but it has a wide days after oviposition, and becomes larger in size
host range. For example, at least 55 hosts are as the larva matures. The pattern of mining is
known from Florida, including bean, beet, carrot, irregular. Both leaf mining and stippling can
celery, cucumber, eggplant, lettuce, melon, onion, greatly depress the level of photosynthesis in the
pea, pepper, potato, squash, and tomato. Flower plant. Extensive mining also causes premature
American Serpentine Leafminer, Liriomyza Trifolii (Burgess) (Diptera: Agromyzidae)
A 147

leaf drop, which can result in lack of shading and active miners. Adults can be captured by using
sun scalding of fruit. Wounding of the foliage also adhesive applied to yellow cards or stakes.
allows entry of bacterial and fungal diseases.
Although leaf mining can reduce plant growth,
crops such as tomato are quite resilient, and capa- Insecticides
ble of withstanding considerable leaf damage. It is
often necessary to have an average of one to three Chemical insecticides are commonly used to pro-
mines per tomato leaf before yield reductions tect foliage from injury, but insecticide resistance is
occur. Leafminers are most damaging when they a major problem. Insecticide susceptibility varies
affect floricultural crops due to the low tolerance widely among populations, and level of susceptibil-
of such crops for any insect damage. ity is directly related to frequency of insecticide
application. In Florida, longevity of insecticide ef
fectiveness is often only two to four years, and then
Natural Enemies is usually followed by severe resistance among the
treated populations. Rotation among classes of
Parasitic wasps (parasitoids) of the families Braco- insecticides is recommended to delay development
nidae, Eulophidae, and Pteromalidae are impor- of resistance. Reduction in dose level and frequency
tant in natural control, and in the absence of of insecticide application, as well as preservation of
insecticides usually keep this insect at low levels of susceptible populations through nontreatment of
abundance. At least 14 parasitoid species are some areas, are suggested as means to preserve
known from Florida alone. Species of Eulophidae insecticide susceptibility among leafminer popula-
such as Diglyphus begina (Ashmead), D. interme- tions. Insect growth regulators have been more sta-
dius (Girault), D. pulchripes, and Chrysocharis ble, but are not immune from the resistance problem.
parksi Crawford are generally found to be most Insecticides also are highly disruptive to naturally
important in studies conducted in North America, occurring biological control agents, particularly
although their relative importance varies geo- parasitoids. Use of many chemical insecticides
graphically and temporally. Predators and diseases exacerbates leafminer problems by killing parasi-
are not considered to be important, relative to par- toids of leafminers. This usually results when insec-
asitoids. However, both larvae and adults are sus- ticides are applied for lepidopterous insects, and use
ceptible to predation by a wide variety of general of more selective pest control materials such as
predators, particularly ants. Bacillus thuringiensis is recommended as it allows
survival of the leafminer parasitoids. Because para-
sitoids often provide effective suppression of leaf-
Management miners in the field when disruptive insecticides are
not used, there has been interest in release of para-
Sampling sitoids into crops. This occurs principally in green-
house-grown crops, but is also applicable to field
There are many methods to assess leafminer abun- conditions. Steinernema nematodes have also been
dance. Counting mines in leaves is a good index of evaluated for suppression of leaf mining activity.
past activity, but many mines may be vacant. High levels of relative humidity (at least 92%) are
Counting live larvae in mines is time consuming, needed to attain even moderately high (greater than
but more indicative of future damage. Puparia can 65%) levels of parasitism. Adjuvants that enhance
be collected by placing trays beneath foliage to nematode survival increase levels of leafminer mor-
capture larvae as they evacuate mines, and the tality, but thus far nematodes are not considered to
captures are highly correlated with the number of be a practical solution to leafminer infestations.
A American Silkworm Moths (Lepidoptera: Apatelodidae)

Cultural Practices

Because broadleaf weeds and senescent crops may

serve as sources of inoculum, destruction of weeds
and deep plowing of crop residues are recom-
mended. Adults experience difficulty in emerging
if they are buried deeply in soil.
Vegetable Pests and Their Management

American Silkworm Moths (Lepidoptera:
Capinera JL (2001) Handbook of vegetable pests. Academic Apatelodidae), Figure 43 Example of American
Press, San Diego, CA, 729 pp
silkworm moths (Apatelodidae), Apatelodes palma
Leibee GL (1984) Influence of temperature on develop-
ment and fecundity of Liriomyza trifolii (Burgess) Druce, from Ecuador.
(Diptera: Agromyzidae) on celery. Environ Entomol
Minkenberg OPJM (1988) Life history of the agromyzid fly Some researchers consider the family part of
Liriomyza trifolii on tomato at different temperatures.
Bombycidae. The family is in the superfamily
Entomologia Expimentalis et Applicata 48:7384
Minkenberg OPJM, van Lenteren JC (1986) The leafminers Bombycoidea (series Bombyciformes), in the
Liriomyza bryoniae and L. trifolii (Diptera: Agromyzi- section Cossina, subsection Bombycina, of the
dae), their parasites and host plants: a review. Wagenin- division Ditrysia. Adults (Fig. 43) small to
gen Agric Univ Pap 862. 50 pp
Parrella MP, Robb KL, Bethke J (1983) Influence of selected
medium size (2074 mm wingspan), with head
host plants on the biology of Liriomyza trifolii (Diptera: scaling roughened; haustellum absent (rarely
Agromyzidae). Ann Entomol Soc Am 76:112115 vestigial); labial palpi small; maxillary palpi
Schuster DJ, Gilreath JP, Wharton RA, Seymour PR (1991) absent; antennae bipectinate; body robust. Wings
Agromyzidae (Diptera) leafminers and their parasitoids
in weeds associated with tomato in Florida. Environ broadly triangular; hindwings rounded. Macula-
Entomol 20:720723 tion varied but mostly shades of brown or gray,
Zehnder GW, Trumble JT (1984) Spatial and diel activity of rarely more colorful, with various markings.
Liriomyza species (Diptera: Agromyzidae) in fresh mar-
Adults are nocturnal. Larvae are leaf feeders.
ket tomatoes. Environ Entomol 13:14111416
Host plants include various records in Aquifoli-
aceae, Betulaceae, Bignoniaceae, Lauraceae,
Oleaceae, Rosaceae, among others.
American Silkworm Moths
(Lepidoptera: Apatelodidae)

john b heppner References

Florida State Collection of Arthropods,
Gainesville, FL, USA Franclemont JG (1973) Apatelodidae. In: Dominick RB, et al
(eds), The moths of America north of Mexico including
American silkworm moths, family Apatelodidae, Greenland. Fasc. 20.1, Bombycoidea, 1623. Classey EW,
are exclusively New World, and total 252 species, Seitz A (ed) (1929) Familie: Bombycidae. Die Gross-Schmet-
mostly Neotropical (247 sp.). Three subfamilies terlinge der Erde, 6:675711, pl. 89, 140142. A. Kernen.
are known: Apatelodinae, Epiinae, and Phiditiinae. [Apatelodidae], Stuttgart, Germany
American Tropical Silkworm Moths (Lepidoptera: Oxytenidae)
A 149

American Swallowtail Moths Larvae are leaf feeders, but few known biologically.
(Lepidoptera: Sematuridae) Host plants are unrecorded.

john b. heppner
Florida State Collection of Arthropods, References
Gainesville, FL, USA
Seitz A (ed) (1930) Familie: Uraniidae [part]. Die Gross-
American swallowtail moths (Fig. 44), family Schmetterlinge der Erde, 6:829837, pl. 139. A. Kernen,
Sematuridae, total 36 Neotropical species, one of Stuttgart, Germany
Fassl AH (1910) Die Raupe einer Uranide. Zeitschrift fr
which just reaches into the United States, in southern
Wissenschaftliches Insektenbiologie 6:355
Arizona. The family is in the superfamily Uran- Strand E (1911) Zur Kenntnis der Uraniidengattung Coro-
ioidea, in the section Cossina, subsection Bom- nidia Westw. and Homidia Strand n. g. (=Coronidia
bycina, of the division Ditrysia. Adults medium to auct. p.p.) (Lep.). Deutsche Entomologische Zeitschrift
large (42100 mm wingspan), with head rough- Westwood JO (1879) Observations on the Uraniidae, a family
ened and eyes large; haustellum naked; labial palpi of lepidopterous insects, with a synopsis of the family and
upcurved, with long second segment and correctly a monograph of Coronidia, one of the genera of which it
angled short, smooth apical segment; maxillary is composed. Trans Zool Soc London 10:507542, 4 pl

palpi minute, 1-segmented; antennae thickened,

with elongated club (slightly hooked at tip). Wings
triangular, with hindwings tailed (usually hind- American Tropical Silkworm
wings with some emarginations); body sometimes Moths (Lepidoptera: Oxytenidae)
robust. Maculation various shades of darker brown,
with vertical lines and bands, often brightly colored john b. heppner
in the hindwings; often with eyespots on the tails. Florida State Collection of Arthropods,
Adults are nocturnal but some may be crepuscular. Gainesville, FL, USA

American tropical silkworm moths, family Oxyteni-

dae, include 60 species, all Neotropical. Some spe-
cialists consider this family a subfamily of
Saturniidae. The family is in the superfamily Bom-
bycoidea (series Saturniiformes), in the section
Cossina, subsection Bombycina, of the division Dit-
rysia. Adults medium size to large (4598 mm
wingspan), with head vertex somewhat roughened;
haustellum developed; labial palpi very large; max-
illary palpi absent; antennae bipectinate; body
somewhat slender or robust but with hair-like
scales. Wings triangular with with falcate apex but
sometimes rounded; hindwings somewhat angled
and with short tails or sometimes rounded. Macula-
tion mostly white with paired dark gray vertical
American Swallowtail Moths (Lepidoptera: striae and hindwings similar, but some species are
Sematuridae), Figure 44 Example of American dark brown with indistinct markings. Adults noc-
swallowtail moths Sematuridae), Sematura lunus turnal. Larvae are leaf feeders; some mimic snakes.
(Linnaeus), from Costa Rica. Host plants recorded in Rubiaceae.
A Ametabolous

References respectively. The cysts of the honeybee amoeba

are ingested and excyst, releasing slender pri-
Heppner JB (2003) Oxytenidae. Lepidopterorum catalogus, mary trophozoites that penetrate and multiply
(n.s.). Fasc. 115. Association for Tropical Lepidoptera, in the midgut epithelium. Secondary trophozo-
Gainesville, FL, 12 pp ites emerge from these cells and migrate to the
Jordan K (1924) On the Saturnoidean families Oxytenidae
and Cercophanidae. Novitates Zoologicae 31:135193 lumen of the Malpighian tubules. These tro-
Schssler H (1936) Oxytenidae. Lepidopterorum catalogus, phozoites, having pseudopodia, feed in the
W. Junk, The Hague, 75:120 lumen and cause a flattening of the epithelial
layer and a distension of the tubules. The brush
border in contact with the amoeba swells in
Ametabolous size and loses the associated secretory trans-
port vesicles. Infected tubules contain a mix of
Organisms that do not display the process of secondary trophozoites, precysts, and cysts. The
metamorphosis. In ametabolous organisms there primary damage to the host bee is the malfunc-
is little change in body form during growth and tion of the Malpighian tubules. Both numbers
molting. of amoeba and the presence of other disease
Metamorphosis agents determine the severity of the amoebiasis
in the bee. In general, this disease either induces
stress or under appropriate conditions in the
springtime can be debilitative, resulting in hive
A family of mayflies (order Ephemeroptera). dwindling.
Mayflies Malamoeba locustae, also known as Malam-
oeba locusta, has been detected in a wide range of
grasshopper species and in a single Thysanuran
Amino Acid species. Its life cycle is very similar to that observed
with M. mellificae. The host grasshoppers ingest
Chemical compounds that may occur free, or the resistant uninucleate cysts, and excysted pri-
linked by peptide bonds into proteins. mary trophozoites invade the midgut and caecal
tissues. Within these tissues the trophozoites grow
and divide, and within about 10 days release prog-
eny secondary trophozoites into the lumen. These
Ammophilous cells migrate to the lumen of the Malpighian
tubules and undergo additional cell divisions
Sand loving. Organisms inhabiting or preferring (Fig. 45). The vegetative development of this
sandy habitats are called ammophilous (adjective) amoeba damages the serosal membrane of the
or ammophiles (noun). tubules, inhibiting their response to insect diuretic
hormone. The infected tubules become packed
with trophozoites and cysts. At high levels,
M. locustae may inhibit the excretory function of
Amoebae the tubules and cause the grasshoppers to become
lethargic prior to death. The distended, amoeba-
The two best-studied insect amoebae are Mal- infected tubules may rupture, releasing both
pighamoeba mellificae and Malamoeba locustae, trophozoites and cysts into the hemocoel. These
which are associated with the honeybee, Apis amoebas are quickly recognized as non-self
mellifera, and the Melanoplus grasshoppers, and are encapsulated by circulating phagocytic
A 151

Amoebae, Figure 45 Light micrograph of the cysts of Malamoeba locusta released from infected
Malpighian tubules.

hemocytes. This disease, although a problem in lab- Amphienotomidae

oratory cultured grasshoppers, is rarely detected
in natural populations. A family of psocids (order Psocoptera).
Bark-Lice, Book-Lice, or Psocids

Brooks WM (1988) Entomogenous Protozoa. In: Ignoffo C (ed),
Handbook of natural pesticides, vol 5. Microbial insec- A family of psocids (order Psocoptera).
ticides. Part A. Entomogenous protozoa and fungi. CRC
Press, Boca Raton, FL, pp 1149 Bark-Lice, Book-Lice, or Psocids
Liu TP (1985) Scanning electron microscopy of developmen-
tal stages of Malpighamoeba mellificae Prell in the hon-
eybee. J Protozool 32:139144 Amphipterygidae
A family of damselflies (order Odonata).
Amoebiasis Dragonflies and Damselflies

Infection of an insect by amoebae.

Amorphoscelididae A family of moths (order Lepidoptera) also known
as double-eye moths.
A family of praying mantids (Mantodea). Double-Eye Moths
Praying Mantids Butterflies and Moths
A Amphitoky

Amphitoky Anagrus species (Mymaridae), among the smallest

insects known, are endoparasitoids of eggs of
A type of parthenogenesis in which both females Odonata and Hemiptera. The genus is worldwide
and males are produced. and about 60 species is now recognized.

Amphizoidae Taxonomy and Adult Morphology

A family of beetles (order Coleoptera). They com- The metasoma of Anagrus is not constricted basally,
monly are known as trout stream beetles. so it appears broadly sessile, the hypochaeta in front
Beetles of the marginal vein is basal to the first macro-
Wasps, Ants, Bees and Sawflies chaeta, the tarsi are 4-segmented, the posterior
scutellum is longitudinally divided, and the foretibia
has a comb-like spur. Adult males and females are
Amplification similar, differing mainly in their antennae, with nine
segments and clubbed in females (Fig. 46) and 13
In molecular biology, the production of additional segments and filiform in males. Body color is often
copies of a chromosomal sequence, found as either darker in males. The genitalia, both in males (the
intrachromosomal or extrachromosomal DNA. In aedeagus) and in females (the ovipositor), have
medical entomology, the production of increased features of taxonomic importance.
numbers of virus in a host. This is often a prerequisite The genus is subdivided into three subgenera
to acquisition and transmission of the virus by a Anagrella, Anagrus, and Paranagrus.
blood-feeding insect.

Amplification Hosts
Like all holometabolous insects, Anagrus species
Hosts of viruses that allow amplification of the virus, have three distinct immature stages, egg, larva and
usually used in the context of arboviruses. Some pupa. The egg is stalked, with an ovoid body that
hosts do not allow amplification, and so serve as an swells during embryogenesis. There are two, apodous,
end-point in the virus cycle. larval instars (Fig. 47), which appear completely dif-
Dead-end Hosts ferent from one another. The first instar is sacciform
and usually attached to the egg chorion. It does not
show any cuticular structure that could serve to feed,
Ampulicidae breathe or feel. It is completely immobile and proba-
bly obtains nourishment and breathes through its
A family of wasps (order Hymenoptera). cuticle. The second instar is divided weakly into six
body segments, has a mouth and a salivary gland
opening, two mandibles and an anus, and various
Anagrus Fairyflies (Hymenoptera: other, probably sensory, structures. No spiracle is
Mymaridae) present. Second instar larvae are very active and
fight each other when in the same host egg. The
elisabetta chiappini mature larva (prepupa) develops inside the egg
Universit Cattolica del Sacro Cuore, into an exarate pupa and does not spin a cocoon.
Piacenza, Italy When development is complete, adults are
Anagrus Fairyflies (Hymenoptera: Mymaridae)
A 153

Anagrus Fairyflies (Hymenoptera: Mymaridae), Figure 46 Adult female of Anagrus sp.

females insert their ovipositor into the slit made

by the host or through the plant tissue itself,
depending on the species. Adults occur in various
habitats, both natural and cultivated, depending
on where their hosts occur. This includes dry habi-
Anagrus Fairyflies (Hymenoptera: Mymaridae), tats such as vineyards and beet fields to damp or
Figure 47 Larva of Anagrus sp. aquatic ones (ponds) where host eggs are found in
plants such as Cyperus or Nuphar.
r ecognizable through the host egg chorion, through Certain Anagrus species can develop both as
which they chew a hole to exit. After emergence the solitary or gregarious parasitoids in eggs of differ-
adults shed their waste products (meconium). Males ent size, whereas others appear to be much more
are usually protandrous. specialized on eggs of the same size, in which they
always develop as solitary parasitoids.
Many Anagrus are extremely important
Behavior and Ecology because they provide control of potentially serious
pests on many agricultural crops. The most impor-
Reproduction is bisexual or parthenogenetic. The tant examples are against leafhoppers such as
latter reproduction is usually arrhenotokous but, Empoasca vitis Goethe and Zygina rhamni (Fer-
rarely, thelytokous parthenogenesis has been rari) in vineyards in Europe and Erythroneura spp.
recorded. Females are ready to oviposit as soon as in North America, against leaf- and planthoppers
they emerge. Copulation, if it occurs, is usually such as Nilaparvata spp. and Sogatella spp. on rice
very quick (some tens of seconds) and insemi- in eastern Asia, and against Perkinsiella sachari-
nated females generally do not copulate again. cida Kirkaldy on sugarcane in Hawaii.
When fed with sugar water, honey or nectar, adults Some biological supply companies mass pro-
may live for up to 10 days. It is thought that adult duce and sell Anagrus atomus L. for biological
host feeding may occur as in other parasitoids. control. Care must be taken to ensure the sanitary
Anagrus species mainly parasitize leafhoppers conditions of the product as the parasitoid is bred
(Cicadellidae), planthoppers, (Delphacidae) and on the natural host eggs inserted into plant tissue,
damsel- or dragonfly (Odonata) eggs, all of which which could be a potential vehicle for other pests
are embedded in plant tissue. To reach the eggs, or diseases.
A Anajapygidae

References abdominal segment; in caterpillars (Lepidoptera) it

refers to a ventral projection at the tip of the abdo-
Chiappini E, Lin NQ (1998) Anagrus (Hymenoptera: Mymari- men that is used to eject frass; in some beetle
dae) of China, with descriptions of nine new species. (Coleoptera) larvae it refers to cerci-like projections
Ann Entomol Soc Am 91:549571 near the tip of the abdomen.
Chiappini E, Triapitsyn SV, Donev A (1996) Key to the Hol-
arctic species of Anagrus Haliday (Hymenoptera:
Mymaridae) with a review of the Nearctic and Palaearc-
tic (other than European) species and descriptions of Anal Gills
new taxa. J Nat Hist 30:551595
Moratorio MS (1990) Host finding and oviposition behavior Gills found at the tip of the abdomen and usually
of Anagrus mutans and Anagrus silwoodensis Walker
(Hymenoptera: Mymaridae). Environ Entomol
consisting of three to five small clusters.
19:142147 Abdomen of Hexapods
Moratorio MS, Chiappini E (1995) Biology of Anagrus incar-
natosimilis and Anagrus breviphragma. Bollettino di
Zoologia Agraria e di Bachicoltura, Serie II, 27:143162 Anal Hooks
Triapitsyn SV (1997) The genus Anagrus (Hymenoptera:
Mymaridae) in America south of the United States: a
review. Ceiba 38:112 In Lepidoptera, small hook or club-shaped struc-
Triapitsyn SV (1998) Anagrus (Hymenoptera: Mymaridae) tures at the tip of the abdomen that serve to anchor
egg parasitoids of Erythroneura spp. and other leafhop- the pupa to the cocoon or silk pad.
pers (Hemiptera: Cicadellidae) in North America vine-
yards and orchards: a taxonomic review. Trans Am
Entomol Soc 124:77112
Anal Furrow
The suture-like groove in the membrane of the
Anajapygidae wing.
Wings of Insects
A family of diplurans (order Diplura).
Anal Legs (Prolegs)
Anal Angle In holometabolous larvae, especially Lepidoptera
larvae, the appendages of the tenth abdominal
The hind angle of the forewings. segment (the terminal prolegs).
Wings of Insects

Anal Lobe
Anal Cell
The posterior region of the wing, occupied by the
A cell in the anal area (anal lobe) of a wing. anal veins.
Wings of Insects Wings of Insects

Anal Comb Anal Loop

This term is applied to a variety of structures that A cluster of cells between the anal wing veins, or
differ depending on the taxon. In flea (Siphonaptera) between the cubitus and anal vein, in Odonata.
larvae, it refers to several rows of setae on the tenth Wings of Insects
Andean Moon Moths (Lepidoptera: Cercophanidae)
A 155

Anal Plate Andean Moon Moths

(Lepidoptera: Cercophanidae)
The shield-like plate or dorsal covering on the ter-
minal segment in caterpillars, and some other lar- john b. heppner
vae. It usually is dark in color, and is also called the Florida State Collection of Arthropods,
anal shield. Gainesville, FL, USA

Andean moon moths, family Cercophanidae,

include 30 species of mostly austral South Ameri-
Anal Tube
can moths. There are two subfamilies: Cercophani-
nae (four sp.) and Janiodinae (26 sp.). Some
Eversible, tubular organs in the anal region of lar-
specialists consider this family a subfamily of Sat-
val Coleoptera. These organs are armed with
urniidae. The family is in the superfamily Bomby-
microspines and assist in attachment to the
coidea (series Saturniiformes), in the section
Cossina, subsection Bombycina, of the division
Ditrysia. Adults (Fig. 48) medium size to large
(24105 mm wingspan), with head vertex rough-
Anal Vein ened; haustellum absent; labial palpi very large;
maxillary palpi absent; antennae bipectinate; body
Longitudinal unbranched vein, or veins, extending robust, with long hair-like scales. Wings broadly
from the base of the wing to the outer margin of triangular, often with apex falcate, or more rounded;
the wing, below the cubitus vein. hindwings rounded or emarginated but sometimes
Wings of Insects with tails. Maculation various, but mostly shades of
brown with diagonal line and fainter markings, but
some with long tails and lighter tan, and with eyes-
pots. Adults are nocturnal. Larvae are leaf feeders.
Postembryonic development in which additional
abdominal body segments are added at the time of
molting (the opposite of epimorphosis).

A family of wood wasps (order Hymenoptera,
suborder Symphyta). They commonly are known
as incense-cedar wood wasps.
Wasps, Ants, Bees and Sawflies

Andean Moon Moths (Lepidoptera:

Cercophanidae), Figure 48 Example of Andean
A family of fleas (order Siphonaptera). moon moths (Cercophanidae), Cercophana venusta
Fleas (Walker) from Chile.
A Andesianidae

Host plants recorded in Celastraceae, Lauraceae, Australia Department of Agriculture. He began a

Saxifragaceae, and Tiliaceae. study of a weevil pest of fruit trees, Otiorhynchus
cribricollis, that required detailed autecological
studies. In 1933, he moved to Melbourne, appointed
References by the CSIR as assistant research officer, to work on
the autecology of Thrips imaginis, a pest of apple
Angulo AO, Heppner JB (2004) Cercophanidae. Lepidoptero- trees. He worked in the School of Agriculture and
rum catalogus, (n.s.). Fasc. 116. Association for Tropical Forestry at the University of Melbourne. While
Lepidoptera, Gainesville, FL, 8 pp
Jordan K (1924) On the Saturnoidean families Oxytenidae
working, he was able to complete a thesis for which
and Cercophanidae. Novitates Zoologicae 31:135193, he was awarded the degree of Master of Agricul-
pl. 621 tural Sciences. He married, and in 1935 moved to
Schssler H (1936) Cercophanidae. Lepidopterorum catalo- the Waite Agricultural Research Institute in Ade-
gus, W. Junk, The Hague, 76:112
Ureta RE (1943) Revisin del gnero Polythysana Wlk. (Sat- laide. His main duties now turned to a study of
urniidae). Boletin del Museo Nacional de Historia Nat- Austroicetes cruciata, a plague grasshopper, and
ural de Chile 21:5570, 4 pl diapause of its eggs. However, his supervisor, who
Wolfe KL, Balczar LMA (1994). Chiles Cercophana venusta
had been working on Thrips imaginis, died sud-
and its immature stages (Lepidoptera: Cercophanidae)
Trop Lepidoptera 5:3542 denly, leaving copious unanalyzed data, whose
completion and publication fell to Herbert. The
published work was criticized because it con-
cluded that climatic factors were all-important in
Andesianidae the population dynamics of the pest, without
room for action of biotic factors. But it led to
A family of moths (order Lepidoptera) also known collaboration with L.C. Birch on a book (1954)
as valdivian forest moths. The distribution and abundance of animals.
Butterflies and Moths Then, after Herbert moved to the Zoology
Valdivian Forest Moths Department of the University of Adelaide, it led
to a book designed as a textbook for students:
(1961) Introduction to the study of animal pop-
ulations. In 1962, Herbert was appointed chair-
Andrenidae man of the Zoology Department. In the 1960s,
with collaborators, he developed a program for
A family of bees (order Hymenoptera, superfamily control of Dacus tryoni, Queensland fruit fly,
Apoidae). by release of sterile males. His next book, also
Bees co-authored with L.C. Birch was (1984) The
Wasps, Ants, Bees and Sawflies ecological web. He died on January 27, 1992, fol-
lowing his wife by some years, but survived by his
son and daughter.

Andrewartha, Herbert George

Herbert Andrewartha was born in Perth, Australia,

on December 21, 1907. In 1924, he entered the Uni-
Birch LC, Browning TO (1993) Herbert George Andrewartha
versity of Western Australia from which he obtained 19071992. Historical Records of Australian Science
a bachelors degree in agriculture. He was then 9(3), Available at www.asap.unimelb.
appointed as assistant entomologist by the Western aasmemoirs/andrewar.htm Accessed August 2002
Angel Insects (Zoraptera)
A 157

Androconia Angel Insects (Zoraptera)

In Lepidoptera, glandular wing and body scales. This is a small group of minute insects. They are
Scent scales. infrequently encountered, and poorly known. The
order name is based on the Greek words zoros
(pure), a (without), and pteron (wing). There are
Androparae only about 30 species described, all in the family
In aphids, viviparous females that are produced on
the secondary host in the autumn, and then fly to
the primary host to produce males. Characteristics
Angel insects are only about 3 mm long, with a
wing span of 7 mm. They are dimorphic: a wing-
Anemometer less form that lacks eyes, ocelli, and is only
slightly pigmented, and a winged form (Fig. 49)
An instrument used for measuring wind speed, an that bears eyes, ocelli, and is darker in color. They
important tool when considering use of pesticides have chewing mouthparts. The antennae are
because high wind speeds can result in pesticide filiform, and consist of nine segments. The
drift. legs are unspecialized, the tarsi 2-segmented.
The wings have simplified venation, and the
wings can be shed, as is the case with termites.
Anemotaxis The abdomen is cylindrical and consists of
11segments. Very short, 1-segmented cerci
A movement in response to air movement or air occur near the tip of the abdomen. Metamor-
currents. phosis is not pronounced.

Angel Insects (Zoraptera), Figure 49 A diagram of an angel insect showing a dorsal view. The wings are
removed from the left side of the body.
A Angoumois Grain Moth, Sitotroga cerealella (Lepidoptera: Gelechiidae)

Biology In humans the same disease is known as African

sleeping sickness or human trypanosomiasis. It is
Angel insects are found beneath bark, in humus, transmitted by tsetse flies in Africa.
decaying wood, and sometimes in association with Trypanosomes
termites. They are believed to feed on fungi. Appar- Tsetse Flies
ently they swarm, and drop wings after swarming. Sleeping Sickness or African Trypanosomiasis
They are gregarious, but there is no evidence of
social organization.
References A family of web-spinners (order Embiidina).
Arnett RH Jr (2000) American insects, 2nd edn. CRC Press,
Boca Raton, FL, 1003 pp
Riegel GT (1987) Order Zoraptera. In: Stehr FW (ed) Imma-
ture insects, vol 1. Kendall/Hunt Publishing, Dubuque,
Iowa, pp 184185
Gurney AB (1938) A synopsis of the order Zoraptera with A family of flies (order Diptera). They commonly
notes on the biology of Zorotypus hubbardi Caudell. are known as wood gnats.
Proc Entomol Soc Wash 40:5787

Angoumois Grain Moth, Sitotroga Anneal

cerealella (Lepidoptera:
Gelechiidae) The process by which the complementary base
pairs in the strands of DNA combine.
This is an important primary pest of stored grain.
Stored Grain and Flour Insects
A plant that normally completes its life cycle of
seed germination, vegetative growth, reproduc-
Forming an angle.
tion, and death in a single growing season or year.

Anholocyclic Life Cycle

A life cycle in which there is a complete lack of
male insects (generally aphids). In this type of life A family of beetles (order Coleoptera). They com-
cycle only viviparous parthenogenetic females are monly are known as death-watch beetles.
present throughout the year. (contrast with holo- Beetles
cyclic life cycle)
Animal Sleeping Sickness A family of moths (order Lepidoptera). They also
are known as Australian primitive ghost moths.
Also known as nagana, this is a disease of animals Australian Primitive Ghost Moths
caused by protozoans in the genus Trypanosoma. Butterflies and Moths
Antennae of Hexapods
A 159

Anoplura Antennae of Hexapods

A suborder of wingless ectoparasitic insects com- severiano f. gayubo
monly known as sucking lice (order Phthiraptera). Universidad de Salamanca, Salamanca, Spain
It is sometimes treated as an order.
Chewing and Sucking Lice According to the known data on anatomy and
embryology, the antennae are postoral structures
of an appendicular nature that have been dis-
Anostostomatidae placed, and now situated secondarily above the
anterolateral regions of the cranium, in front of
A family of crickets (order Orthoptera). They the mouth.
commonly are known as wetas and king crickets. Taking into account their intrinsic muscula-
Grasshoppers, Katydids and Crickets ture, two fundamental types of antennae can be
The lower of the two divisions of the clypeus. Segmented Type
Mouthparts of Hexapods
Each antennal division (antennomere) possesses
intrinsic musculature (although the last segment
generally lacks it). This type is found in Diplura
Antagonist and Collembola.

An antagonist usually is an organism (usually a

pathogen) that does no significant damage to the Annulated Type
host, but its colonization of the host protects the
host from significant subsequent damage by a pest. Three segments are recognized that, from the basal
to the apical zone of the antenna, are called scape,
pedicel and antennal flagellum (Fig. 50).
Antecosta (pl., antecostae) The scape is a robust segment that unites the
head capsule with a cuticular reinforcement (Fig. 51),
An internal ridge on the anterior portion of a ter- the antennal socket (also called the torulus). In the
gum or sternum. It serves as a point of attachment antennal socket one or two condyles are distin-
for the longitudinal muscles. guished, which serve to articulate the scape. The
second segment is called the pedicel, and it
usually varies in form and development, although
Antenna (pl., antennae) it is generally small. Lastly, the flagellum is usu-
ally divided into several divisions called
The paired segmented sensory organs, borne one flagellomeres.
on each side of the head. The antennae commonly The movement of the antennae is carried
protrude forward. Each antenna (Figs. 50 and 51) out through extrinsic or motor muscles of the
consists of three segments: the basal scape, a small scape, generally forming three or four functional
pedicel, and an elongate flagellum. The flagellum groups. Depending on the insect group, these
is usually subdivided into many sections. muscles are inserted in the head capsule or in
Antennae of Hexapods the tentorium.
A Antennae of Hexapods

lateral ocellus
compound eye
pedicel postocular area

cervical sclerites
tentorial suture
basimandibular sclerite
clypeus labium

labrum labial palpus


maxillary palpus

Antennae of Hexapods, Figure 50 Side view of the head of an adult grasshopper, showing some major

The antennae usually have bristles and antennae with respect to the head, the direction
s ensilla of different types that act as chemore- and force of the wind, or of the water currents in
ceptor-, thermoreceptor- or hygroreceptor-type aquatic insects. In addition, it can act as an audi-
sensory organs. In addition, the antennae of tory organ in male mosquitoes and chirinomids,
males display modifications tending to increase which perceive the sound produced by the
their surface area, which permits harboring a females in flight.
great number of sensilla and acting as detectors The number of flagellomeres is a character
that detect pheromones emitted by the females, that, in certain cases, is related to the sex, as
and enable (usually) the males to locate the occurs in some Aculeate Hymenoptera in which
females for reproductive functions. Certain the males display 11 flagellomeres and the
modifications in the antennae of the males can females 10. In others, it represents an important
also be related to particular courtship behavior taxonomic character, emphasized in this sense
prior to mating. the family Argidae (Hymenoptera: Symphyta)
In relation to the functions carried out by whose individuals display the flagellomere
the antennae, it is necessary to highlight the undivided.
presence, in the pedicel, of Johnstons organ, Various types of antennae exist (Fig. 52), the
which is formed by cordotonal sensilla. It is appearance of which is owed fundamentally to the
fundamentally a proprioreceptor organ that variation in form and development of the flagellom-
provides information about the position of the eres. The most important are:
Antennae of Hexapods
A 161

vertex flagellum

frons pedicel

median ocellus


anterior tentorior pit

basimandibular sclerite
frontoclypeal suture
clypeolabral suture

maxillary palpus

labial palpus
Antennae of Hexapods, Figure 51 Front view of the head of an adult grasshopper, showing some major

Filiform Aristate

The flagellomeres, normally numerous, are nar- The last flagellomere is normally very wide and
row, cylindrical, and of similar size. It is the most bears a conspicuous bristle named the arista.
common type in the insects. Examples are found in Diptera (Syrphidae and


There exists a narrowing in the union of each Stylate

flagellomere, which are more or less spherical, with
the antenna acquiring a rosaried appearance The last flagellomere is prolonged apically in a
(like the beads of a rosary). There are examples in fine and elongated process named the style. Exam-
various families of beetles. ples are found in Diptera (Rhagionidae and


The flagellomeres are extremely fine and dimin- Clavate

ish in diameter gradually toward the tip; the
antenna thus acquires an appearance of seta or The flagellomeres increase in diameter gradually
hair. The antennae of Odonata constitute a typical toward the apex. Examples are found in Coleoptera
example. (Coccinellidae and Tenebrionidae).
A Antennae of Hexapods

Antennae of Hexapods, Figure 52 Some common types of antennal forms: A, filiform; B, moniliform;
C, capitate; D, clavate; E, setaceous; F, serrate; G, pectinate; H, bipectinate; I, plumose; J, aristate;
K, stylate; L, lamellate; M,flabellate; N, geniculate.

Capitate Pectinate

In this case the last flagellomeres are of greater The flagellomeres project laterally, forming a fine
diameter, in contrast with the preceding, forming and more or less elongated projection. When it is
aclub ormace. Examples are found in Coleoptera produced over two sides of each flagellum, the
(Nitidulidae and Silphidae). antennae are called bipectinate. Examples are
found in Coleoptera (Pyrochroidae).

Serrate Flabellate

The flagellomeres display pointed, lateral prolon- The flagellum displays long, flattened or more or
gations, on one side or on both. Examples are less cylindrical expansions. Examples are found in
found in Coleoptera (Elateridae). some species of Coleoptera (Scarabeidae).
Anther Smut of Carnations
A 163

Lamellate Antennal Fossa

Only the last flagellomeres display long, lateral A groove or cavity in which the antennae are
expansions. Examples are found in Coleoptera located or concealed. This is also called the anten-
(Scarabeidae, subfamily Melolonthinae). nal insertion.
Antennae of Hexapods

Antennal Sclerite
Flagellomeres with numerous long hairs are
arranged in a feather-like or whorled form. Exam- A ring into which the basal joint of each antenna
ples are found in male mosquitoes (Diptera). is inserted.
Antennae of Hexapods

The scape is relatively long, forming a clear angle
Sensory or tactile movements with the antennae
with the rest of the antenna (pedicel plus flagel-
that result in contact of the antennae with an object.
lum). Examples are found in Hymenoptera (For-
micidae and Chalcidoidea), and in Coleoptera
(Lucanidae). Within this type of antenna, particu-
lar variations can exist, as in the case of the
Ormyridae (Hymenoptera: Chalcidoidea), in A small antennal or feeler-like process.
which the first divisions of the flagellum are of a
lenticular type (lens shaped, or double convex).