Evolution Of The Insects


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Cambridge University Press 0521821495 - Evolution of the Insects David Grimaldi and Michael S. Engel Frontmatter More information

EVOLUTION OF THE INSECTS
Insects are the most diverse group of organisms to appear in the 3-billion-year history of life on Earth, and the most ecologically dominant animals on land. This book chronicles, for the first time, the complete evolutionary history of insects: their living diversity, relationships, and 400 million years of fossils. Whereas other volumes have focused on either living species or fossils, this is the first comprehensive synthesis of all aspects of insect evolution. Current estimates of phylogeny are used to interpret the 400-million-year fossil record of insects, their extinctions, and radiations. Introductory sections include the living species, diversity of insects, methods of reconstructing evolutionary relationships, basic insect structure, and the diverse modes of insect fossilization and major fossil deposits. Major sections cover the relationships and evolution of each order of hexapod. The book also chronicles major episodes in the evolutionary history of insects: their modest beginnings in the Devonian, the origin of wings hundreds of millions of years before pterosaurs and birds, the impact that mass extinctions and the explosive radiation of angiosperms had on insects, and how insects evolved the most complex societies in nature.
Evolution of the Insects is beautifully illustrated with more than 900 photo- and electron micrographs, drawings, diagrams, and field photographs, many in full color and virtually all original. The book will appeal to anyone engaged with insect diversity: professional entomologists and students, insect and fossil collectors, and naturalists.
David Grimaldi has traveled in 40 countries on 6 continents collecting and studying recent species of insects and conducting fossil excavations. He is the author of Amber: Window to the Past and is Curator of Invertebrate Zoology at New York’s American Museum of Natural History, as well as an adjunct professor at Cornell University, Columbia University, and the City University of New York.
Michael S. Engel has visited numerous countries for entomological and paleontological studies, focusing most of his field work in Central Asia, Asia Minor, and the Western Hemisphere. In addition to his positions as Associate Professor in the Department of Ecology and Evolutionary Biology and Associate Curator in the Division of Entomology of the Natural History Museum at the University of Kansas, he is a Research Associate of the American Museum of Natural History and a Fellow of the Linnean Society of London.
David Grimaldi and Michael S. Engel have collectively published more than 250 scientific articles and monographs on the relationships and fossil record of insects, including 10 articles in the journals Science, Nature, and Proceedings of the National Academy of Sciences.

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Cambridge University Press 0521821495 - Evolution of the Insects David Grimaldi and Michael S. Engel Frontmatter More information
Evolution of the Insects
David Grimaldi
American Museum of Natural History
Michael S. Engel
University of Kansas

© Cambridge University Press

www.cambridge.org

Cambridge University Press 0521821495 - Evolution of the Insects David Grimaldi and Michael S. Engel Frontmatter More information

CAMBRIDGE UNIVERSITY PRESS
Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo

Cambridge University Press 40 West 20th Street, New York, NY 10011-4211, USA

www.cambridge.org Information on this title: www.cambridge.org/9780521821490

© David Grimaldi, Michael S. Engel 2005

This book is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press.

First published 2005

Printed in Hong Kong

A catalog record for this publication is available from the British Library.

Library of Congress Cataloging in Publication Data
Grimaldi, David A. Evolution of the insects / David Grimaldi, Michael S. Engel. p. cm. Includes bibliographical references and index. ISBN 0-521-82149-5 (alk. paper) 1. Insects – Evolution. I. Engel, Michael S. II. Title.

QL468.7.G75 2004 595.7Ј138 – dc22

2004054605

ISBN-13 978-0-521-82149-0 hardback ISBN-10 0-521-82149-5 hardback

Cambridge University Press has no responsibility for the persistence or accuracy of URLs for external or third-party Internet Web sites referred to in this book and does not guarantee that any content on such Web sites is, or will remain, accurate or appropriate.

© Cambridge University Press

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Cambridge University Press 0521821495 - Evolution of the Insects David Grimaldi and Michael S. Engel Frontmatter More information

An orthopteran of the extinct family Elcanidae in 120 MYO limestone from Brazil’s Santana Formation. AMNH; length of elcanid (including antennae) 98 mm(3.8 in.).

© Cambridge University Press

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Cambridge University Press 0521821495 - Evolution of the Insects David Grimaldi and Michael S. Engel Frontmatter More information
For the entomophiles, winged and larval

© Cambridge University Press

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Cambridge University Press 0521821495 - Evolution of the Insects David Grimaldi and Michael S. Engel Frontmatter More information

CONTENTS
Preface Commonly Used Abbreviations
1. Diversity and Evolution
Introduction SPECIES: THEIR NATURE AND NUMBER
Drosophila Apis How Many Species of Insects?
RECONSTRUCTING EVOLUTIONARY HISTORY
Systematics and Evolution Taxonomy, Nomenclature, and Classification Paleontology
2. Fossil Insects
INSECT FOSSILIZATION
Types of Preservation
DATING AND AGES MAJOR FOSSIL INSECT DEPOSITS
Paleozoic Mesozoic Cenozoic
3. Arthropods and the Origin of Insects ONYCHOPHORA: THE VELVET WORMS TARDIGRADA: THE WATER BEARS ARTHROPODA: THE JOINTED ANIMALS Marellomorpha: The Lace Crabs Arachnomorpha: Trilobites, Arachnids, and Relatives Crustaceomorpha Mandibulata The Invasion of Land HEXAPODA: THE SIX-LEGGED ARTHROPODS Entognatha: Protura, Collembola, and Diplura
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1
1 6 7 9 11 15 15 33 36
42 42 43 62 65 65 70 84
93 94 96 97 98
98 107 107 109 111
111

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Cambridge University Press 0521821495 - Evolution of the Insects David Grimaldi and Michael S. Engel Frontmatter More information

viii

CONTENTS

4. The Insects

119

MORPHOLOGY OF INSECTS

119

General Structure

119

The Head

121

The Thorax

125

The Abdomen

131

DEFINING FEATURES OF THE INSECTS

137

RELATIONSHIPS AMONG THE INSECT ORDERS

137

A Brief History of Work

137

A Roadmap to the Phylogeny of Insects

144

5. Earliest Insects

148

ARCHAEOGNATHA: THE BRISTLETAILS

148

DICONDYLIA

150

ZYGENTOMA: THE SILVERFISH

150

RHYNIOGNATHA

152

6. Insects Take to the Skies

155

PTERYGOTA, WINGS, AND FLIGHT

155

Insect Wings

156

EPHEMEROPTERA: THE MAYFLIES

160

METAPTERYGOTA

166

PALAEODICTYOPTERIDA: EXTINCT BEAKED INSECTS

168

Palaeodictyoptera

170

Dicliptera

170

Megasecoptera

171

Diaphanopterodea

172

Paleozoic Herbivory

173

ODONATOPTERA: DRAGONFLIES AND EARLY RELATIVES

173

Geroptera

174

Holodonata: Protodonata and Odonata

174

Protodonata: The Griffenflies

175

Order Odonata: The Dragonflies and Damselflies

178

7. Polyneoptera

188

NEOPTERA

188

WHAT ARE POLYNEOPTERA?

189

Plecopterida

192

Orthopterida

193

PLECOPTERA: THE STONEFLIES

194

EMBIODEA: THE WEBSPINNERS

196

ZORAPTERA: THE ZORAPTERANS

199

ORTHOPTERA: THE CRICKETS, KATYDIDS, GRASSHOPPERS, WETAS, AND KIN

202

Ensifera

208

Caelifera

210

PHASMATODEA: THE STICK AND LEAF INSECTS

211

TITANOPTERA: THE TITANIC CRAWLERS

215

CALONEURODEA: THE CALONEURODEANS

217

DERMAPTERA: THE EARWIGS

217

GRYLLOBLATTODEA: THE ICE CRAWLERS

222

MANTOPHASMATODEA: THE AFRICAN ROCK CRAWLERS

224

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Cambridge University Press 0521821495 - Evolution of the Insects David Grimaldi and Michael S. Engel Frontmatter More information
CONTENTS
DICTYOPTERA
Dictyopteran Relationships Blattaria: The Roaches Citizen Roach: Isoptera (Termites) The Predatory Roachoids: Mantodea (Mantises) Ages of the Dictyoptera
8. The Paraneopteran Orders PSOCOPTERA: THE BARK LICE PHTHIRAPTERA: THE TRUE LICE Fossils and Ages FRINGE WINGS: THYSANOPTERA (THRIPS) Feeding Habits Social Behavior Diversity and Relationships Fossils and Origins THE SUCKING INSECTS: HEMIPTERA Sternorrhyncha: Aphids, Whiteflies, Plant Lice, and Scale Insects Auchenorrhyncha: The Cicadas, Plant Hoppers, and Tree Hoppers Coleorrhyncha Heteroptera: The “True Bugs”
9. The Holometabola
PROBLEMATIC FOSSIL ORDERS
Miomoptera Glosselytrodea
THE ORIGINS OF COMPLETE METAMORPHOSIS
ON WINGS OF LACE: NEUROPTERIDA Raphidioptera: The Snakeflies Megaloptera: The Alderflies and Dobsonflies Neuroptera: The Lacewings, Antlions, and Relatives
10. Coleoptera and Strepsiptera
EARLY FOSSILS AND OVERVIEW OF PAST DIVERSITY ARCHOSTEMATA ADEPHAGA MYXOPHAGA POLYPHAGA
STREPSIPTERA: THE ENIGMATIC ORDER Diversity Relationships to Other Orders Fossils
11. Hymenoptera: Ants, Bees, and Other Wasps
THE EUHYMENOPTERA AND PARASITISM ACULEATA
The Ants The Bees (Anthophila)
EVOLUTION OF INSECT SOCIALITY
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227 228 230 238 252 260
261 261 272 275 280 283 283 284 285 287
289
303 312 314
331 331 331 332 333 335 337 340 341
357 360 363 366 370 371 399 402 402 403
407 413 429 440 454 464
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Cambridge University Press 0521821495 - Evolution of the Insects David Grimaldi and Michael S. Engel Frontmatter More information

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CONTENTS

12. Panorpida: Antliophora and Amphiesmenoptera

468

PANORPIDA

468

ANTLIOPHORA: THE SCORPIONFLIES, TRUE FLIES, AND FLEAS

468

MECOPTERIDA: MECOPTERANS AND SIPHONAPTERA

470

Early History

470

Recent Diversity and Relationships

474

The Fleas

480

Evolution of Ectoparasites and Blood Feeders of Vertebrates

489

DIPTERA: THE TRUE FLIES

491

The Brachycera

514

The Cyclorrhapha

531

13. Amphiesmenoptera: The Caddisflies and Lepidoptera

548

TRICHOPTERA: THE CADDISFLIES

548

LEPIDOPTERA: THE MOTHS AND BUTTERFLIES

555

Mesozoic Fossils

556

Basal Groups

560

Ditrysia

573

The “Higher” Ditrysians: Macrolepidoptera

581

Butterflies and Their Relatives (Rhopalocera)

590

Mimicry

602

14. Insects Become Modern: The Cretaceous and Tertiary Periods

607

THE CRETACEOUS

607

Flowering of the World: The Angiosperm Radiations

607

Plant Sex and Insects: Insect Pollination

613

Radiations of Phytophagous Insects

622

Austral Arthropods: Remnants of Gondwana?

625

Insects, Mass Extinctions, and the K/T Boundary

635

THE TERTIARY

637

Mammalian Radiations

638

Pleistocene Dispersal and Species Lifespans

642

Island Faunas

642

15. Epilogue

646

WHY SO MANY INSECT SPECIES?

646

Age

646

Design

646

Capacity for High Speciation Rates

647

Low Rates of Natural Extinction

647

THE FUTURE

647

Glossary

651

References

662

Index

733

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Cambridge University Press 0521821495 - Evolution of the Insects David Grimaldi and Michael S. Engel Frontmatter More information

PREFACE

Writing a book on a subject as vast as the evolution of the most diverse lineage of organisms had one simple justification for us: it was needed. Having taught Insect Diversity and Insect Systematics at the City University of New York, Columbia University, Cornell University, and the University of Kansas, we became acutely aware of a gaping hole in entomology. No volume integrates the unprecedented diversity of living and extinct insects, particularly within the evolutionary framework of phylogeny. Some excellent texts, popular books, and field guides cover insect identification, structure, and living diversity, as well as physiology, behavior, and general biology, of which The Insects of Australia (Naumann, 1991a) is perhaps the best example. For our lectures to students we thus found ourselves pulling an extremely scattered literature together. Instead of trudging through the insect families – interesting as they are – we found that students were fascinated by an approach of folding Recent insect diversity into one large context of phylogeny, biogeography, ecology, and the fossil record. The big picture engaged them. After four years of intensive literature research and writing, study and imaging of important museum specimens, and thousands of figures, we like to think we’ve succeeded in our goal.
Our approach to the volume was tempered by our own experience and interests with fossil insects. Entomologists typically ignore fossils, and since we too work on speciose groups of living insects, we have always been intrigued by the dismissiveness of most entomologists. Why ignore such illuminating parts of evolutionary history? We hope that this book will reveal to our colleagues the significance, and even esthetics, of insect fossils. There are several comprehensive treatments of the insect fossil record, particularly the hexapod section of the Treatise on Invertebrate Paleontology (Carpenter, 1992) and the more recent History of Insects by Rasnitsyn and Quicke (2002). But these volumes are devoted entirely to fossil insects, so something more inclusive, and accessible, was needed.
Compiling a book like this is humbling, not only because of the scope of the subject, but also because discoveries and new work reported every month in paleontology and insect

systematics continually revise the field. As this book was nearing completion, for example, two large projects were launched. One of these is the U.S. National Science Foundation’s Tree of Life project, which seeks to examine the phylogeny of major groups of organisms using all existing data and vast new morphological and DNA data. The other is the Dresden conference on insect phylogeny, which met for the first time in 2003 (e.g., Klass, 2003), and which is intended to meet every few years. Like the insects themselves, our understanding is thus evolving. As more genes become sequenced for hundreds more species of insects, for example, phylogenetic hypotheses will be revised, or at least discussed. But, thirty years ago a book like this would have been very different and much slimmer. Our knowledge of insect relationships has advanced tremendously over this period of time, and dozens of spectacular fossil deposits of insects have been discovered. Tomorrow’s discoveries will reinforce, revise, and entirely redefine our present knowledge, but one needs to start somewhere. The optimal moment is always elusive. We hope that thirty years from now – indeed, twenty – much of what we present here will not fall far from the mark. Should we be so fortunate, new editions of this volume will attempt to keep abreast of developments.
Working at the American Museum of Natural History has also given us a keen appreciation for appealing to the nascent naturalist and scientist, not only to the landed professional. We were very deliberate in developing a volume that would be visually engaging to insect and fossil collectors, general naturalists, botanists, and other biologists, as well as to student and professional entomologists. Although we tried to avoid the thick jargon of entomology and systematics, it was not entirely avoidable (some of the jargon is useful), and we hope our colleagues will understand this was done deliberately to make the subject more digestible. The nearly 1,000 images were also included to make the book more engaging. Should the images and captions whet the reader’s appetite, a healthy meal of text is also available.
A volume like this would not have been possible without the assistance of authoritative colleagues around the world,
xi

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