THE BOOK OF CATERPILLARS THE BOOK OF CATERPILLARS A L I F E - S I Z E G U I D E T O S I X H U N D R E D S P E C I E S F R O M A R O U N D T H E W O R L D EDITED BY DAVID G. JAMES CONTRIBUTORS DAVID ALBAUGH, BOB CAMMARATA, ROSS FIELD, HAROLD GREENEY, JOHN HORSTMAN, DAVID JAMES, SALLY MORGAN, TONY PITTAWAY, JAMES A. SCOTT, ANDREI SOURAKOV, MARTIN TOWNSEND, KIRBY WOLFE THE UNIVERSITY OF CHICAGO PRESS Chicago DAVID G. JAMES is an associate professor in the Department of Entomology, Washington State University, based at the Irrigated Agriculture Research and Extension Center, Prosser. The University of Chicago Press, Chicago 60637 This book was conceived and designed by Ivy Press © 2017 Quarto Publishing plc An imprint of The Quarto Group The Old Brewery, 6 Blundell Street All rights reserved. No part of this book may be used or reproduced London N7 9BH, United Kingdom in any manner whatsoever without written permission, except T (0)20 7700 6700 F (0)20 7700 8066 in the case of brief quotations in critical articles and reviews. www.QuartoKnows.com For more information, contact the University of Chicago Press, 1427 E. 60th St., Chicago, IL 60637. Publisher SUSAN KELLY Creative Director MICHAEL WHITEHEAD Published 2017 Editorial Director TOM KITCH Commissioning Editor KATE SHANAHAN Printed in China Produced by 3REDCARS 26 25 24 23 22 21 20 19 18 17 1 2 3 4 5 Editors RACHEL WARREN CHADD, JOHN ANDREWS Designer JANE MCKENNA ISBN-13: 978-0-226-28736-2 (cloth) Illustrator BILL DONOHOE ISBN-13: 978-0-226-28753-9 (e-book) DOI: 10.7208/chicago/9780226287539.001.0001 JACKET IMAGES Species from the following pages: Bob Cammarata 60, 527, 613; Library of Congress Cataloging-in-Publication Data Ross Field 47, 85, 152, 165, 220, 228, 256; Harold Greeney 67, 107, 213, 226, 237; John Horstman/itchydogimages 74, 195, 209, Names: James, David G., editor. 214, 243, 253, 266, 278, 300, 309, 317, 319, 585; David James 57, Title: The book of caterpillars : a life-size guide to six 105, 125, 135, 138, 170, 179, 191, 241, 260, 282; David Liebman hundred species from around the world / edited by David G. 41, 132; Tony Pittaway 441; James A Scott 286; Shutterstock/ James. Eric Isselee 180; Shutterstock/Sari O’Neal 533; Shutterstock/ Description: Chicago : The University of Chicago Press, 2017. | Kamieniak Sebastian 591; Shutterstock/xpixel 568; Leroy Simon Includes bibliographical references and index. 375, 391, 410, 417, 504; Andrei Sourakov 154, 286; © J Voogd 182, Identifi ers: LCCN 2017013376 | ISBN 9780226287362 (cloth : alk. 540; Wolfgang Wagner 38, 53, 65, 118, 177; Roger Wasley 269, 338; paper) | ISBN 9780226287539 (e-book) Wikimedia Commons/Bernard Dupont 245; Kirby Wolfe 431. Subjects: LCSH: Caterpillars. | Caterpillars—Classifi cation. | Caterpillars—Conservation. LITHOCASE IMAGES Classifi cation: LCC QL542 .B66 2017 | DDC 595.7813/92—dc23 David Liebman 39; Shutterstock/Roger Meerts 509; Wolfgang LC record available at https://lccn.loc.gov/2017013376 Wagner 201. CONTENTS Introduction 6 What is a caterpillar? 10 From eggs to pupation 14 The miracle of metamorphosis 18 Voracious eaters 22 Caterpillar defenses 24 Caterpillars and people 28 Research and conservation 32 The caterpillars 34 BUTTERFLY CATERPILLARS 36 MOTH CATERPILLARS 284 Appendices 640 Glossary 642 Resources 645 Classifi cation of the Lepidoptera 646 Index by common name 648 Index by scientifi c name 651 Notes on contributors 654 Acknowledgments 656 INTRODUCTION 6 ABOVE The Box Tree Caterpillars—the immature stage of moths and butterflies—are diverse Moth caterpillar (Cydalima perspectalis) and remarkable, with an extraordinary range of survival techniques that is one of many species that has have helped make the Lepidoptera one of the most successful insect groups. become widespread outside its native After beetles, it is the second largest order on the planet; at least 160,000 range, having been introduced to Europe species have been identified and described,with thousands more from eastern Asia with imports of its undescribed. Lepidoptera are also very widespread, occupying every host plant, box (Buxus continent except Antarctica, in habitats ranging from rocky mountain spp.). In Asia, natural predators, including slopes to tropical rain forests, and from waste ground to woollen clothes. the Asian Hornet (Vespa velutina), help Their ecological significance, too, is immense. As larvae, they are mostly control its numbers. prodigious herbivores, hosts for parasitic flies and wasps, and potential food for birds, reptiles, and mammals. As adults, they are vital pollinators. The myriad colors, forms, patterns, and sizes of different caterpillars are all part of their arsenal against predation as they grow, pupate, and perform the magic trick of metamorphosis—transformation into a butterfly or moth. Some caterpillars are cleverly disguised in the colors of their habitat, and others are strikingly colored and patterned, announcing to predators that they are unpalatable or even toxic. Certain species have stinging spines, others can pull mammal-like faces, while many Papilionidae butterflies can puff up their front end to look like a snake’s head, complete with eyespots and an everted organ that mimics a forked tongue. All caterpillars, however, share the same basic body plan of a large head, small thorax with six true legs, a comparatively huge ten-segment abdomen, and a large gut where all the material they consume is processed. In most species, a pair of thick, fleshy prolegs is present on half of the abdominal segments, enabling the caterpillar to move around, while INTRODUCTION 7 breathing is conducted through tiny pores, called spiracles, on the sides of TOP A moth caterpillar of the Lasiocampidae the body. A caterpillar feeds for much of its life, using scissor-like jaws, or family is convincingly camouflaged as a mandibles, to snip off and grind up tiny piece after tiny piece of foodstuff. patch of moss on tree bark in Pu’er, As its body expands, it molts, often changing appearance. Most caterpillars Yunnan, China. Many Lepidopteran larvae develop through five instars (stages), shedding their skin at each stage. have an extraordinary ability to blend into From egg hatch to maturity, they will increase in mass by up to 1,000 times. their surroundings. In Lepidoptera, all development occurs at the caterpillar stage, which can take as little as ten days or, if suspended to escape extreme heat or cold, ABOVE The California Sister (Adelpha may last a few years, and up to seven years in the Arctic Woolly Bear californica) caterpillar, when young, rests (Gynaephora groenlandica). When the caterpillar pupates, all the necessary immune from predators on piers cells are present to be reorganized during metamorphosis into a moth or it creates from its frass (excreta). butterfly, whose life is usually much shorter. SELECTION CRITERIA Despite being so numerous, many Lepidoptera are relatively unknown and undescribed, especially at the larval stage. More than 70 families and 55,000 species comprise the “microlepidoptera” group of very small moths, INTRODUCTION with minute caterpillars that have been rarely, if ever, studied or photographed. This book, therefore, focuses on the caterpillars of larger moths and butterflies, which have received most attention from scientists and photographers. The 600 species that are described here reveal the enormous diversity of form, coloration, and adaptation that exists among these creatures. They range in size from large (6 in/150 mm) hawkmoths and Saturniidae larvae, such as the Hickory Horned Devil (Citheronia regalis), to tiny (⅜ in/10 mm) moth caterpillars like the Case-bearing Clothes Moth (Tinea pellionella), with a full panoply of spiny, hairy, striped, and variously patterned and ornamented larvae in between, from every continent where Lepidoptera live. Some of the caterpillars feature unusual 8 adaptations or live in extreme habitats; others are the subject of scientific research, or are culturally significant, or economically important. HOW THE BOOK WORKS The larval life and ecology of 600 species are described in text and images in two sections—Butterfly Caterpillars and Moth Caterpillars. While not strictly a taxonomic division, this reflects common practice, as all butterfly species are generally considered members of the superfamily Papilionoidea, while the more numerous moth species account for all other Lepidoptera. Each caterpillar is shown life size at maturity, together with a line drawing of the adult butterfly or moth. Some have also been magnified to highlight their detail. All images are of live caterpillars, as, unlike adult butterflies and moths, caterpillars cannot be pinned and photographed because they rapidly lose their coloration after death. A distribution map indicates each species’ range. The entry heading may be the species’ common name, accompanied by its Latin name (the genus + species name), or, where there is no accepted common name, only the Latin name. Below the heading, the “authority” is given, that is, who first described the species and the date when it was described. Parentheses are used to show that a genus name has changed since it was first described, while square brackets indicate discrepancies and uncertainties about the author or date. An information box above each entry briefly summarizes key details about the species—its family, range, habitat, host plants or material, a notable fact, and its conservation status. Each species has been checked against the IUCN (International Union for the Conservation of Nature) Red List of Threatened Species, but as relatively few Lepidoptera have been assessed, many species are listed as “Not evaluated,” although this INTRODUCTION 9 is often modified by local expert information and regional or national ABOVE The caterpillar of the Pale Tussock assessments. A few vulnerable species may also be described as being on (Calliteara pudibunda) greatly outshines its an appendix of CITES (Convention on International Trade in Endangered dull-colored adult, with its conspicuous, Species of Wild Fauna and Flora); this means they are subject to an flower-like tufts of yellow hair. These international agreement restricting trade in specimens. are part of its defense mechanism and a warning to potential predators. The hairs SPREADING THE WORD are both urticating and detach easily, Anybody can study Lepidoptera, and finding and keeping caterpillars making the caterpillar distinctly unpalatable. should be as much a feature of a young child’s life as rearing tadpoles. Watching these insects develop and metamorphose can be an inspirational experience. Yet, in many places, species numbers are dwindling as a result of habitat destruction, agricultural development, pesticide use, and climate change. School classroom programs for rearing caterpillars, popular in the United States, Europe, and Australia, do much to stimulate interest in Lepidoptera and create awareness of the threat to their survival. Further research is also required to help better manage their conservation. Very few species cause significant damage, despite their reputation as “pests” for feeding on cultivated plants, and, arguably, any damage is vastly outweighed by the value of butterflies and moths as pollinators. Both adults and caterpillars, in all their wondrous forms, play a further vital role. They live in such a variety of habitats and are so sensitive to change within those habitats, that scientists increasingly view the insects as an important bellwether of environmental health. For without the caterpillar as Lepidoptera progeny, plant regulator, and food for many creatures, ecosystems would collapse.
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