Encyclopedia of Entomology Encyclopedia of Entomology Edited by John L. Capinera University of Florida Second Edition Volume 4 S–Z Professor John L. Capinera Dept. Entomology and Nematology University of Florida Gainesville FL 32611–0620 USA 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. springer.com Editor: Zuzana Bernhart, Dordrecht/ Sandra Fabiani, Heidelberg Development Editor: Sylvia Blago, Heidelberg Production Editor: le-tex publishing services oHG, Leipzig Cover Design: Frido Steinen-Broo, Spanien Printed on acid-free paper SPIN: 11757993 2109 — 5 4 3 2 1 0 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 Boevé 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 Acknowledgments 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 edition. 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. Preface 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. x Preface 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 Entomology 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.