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Insect Morphology PRINCIPLES OF INSECT MORPHOLOGY BY R. E. SNODGRASS United States Department of Agriculture Bureau of Entomolo(JY and Plant Quarantine FIRST EDITION SECOND IMPRESSION McGRA W-HILL BOOK COMPANY, INC. NEW YORK AND LONDON 1935 McGRAW-HILL PUBLICATIONS- IN THE ZOOLOGICAL SCaNCES A. FRANKLIN SHULL, CONSULTING EDITOR PRINCIPLES OF INSECT MORPHOLOGY COPYRIGHT, 1935, BY THE l\1CGRAW-HILIi BOOK COMPANY, INC. PRINTED IN THE UNITED STATES OF AMERICA All rights reserved. This book, or parts thereof, may not be reproduced in any form without permission oj the publishers. \ NLVS/IVRI 111111111 II 1111 1111111111111 01610 TaE MAPLE PRESS COMPANY, YORK, PA. PREFACE The principal value of fa cis is that they give us something to think about. A scientific textbook, therefore, should contain a fair amount of reliable information, though it may be a matter of choice with the author whether he leaves it to the reader to formulate his own ideas as to the meaning of the facts, or whether he attempts to guide the reader's thoughts along what seem to him to be the proper channels. The writer of the present text, being convinced that generalizations are more important than mere knowledge of facts, and being also somewhat partial to his own way of thinking about insects, has not been able to refrain entirely from presenting the facts of insect anatomy in a way to suggest relations between them that possibly exist only in his own mind. Each of the several chapters of this book, in other words, is an attempt to give a coherent morphological view of the fundamental nature and the apparent evolution of a particular group of organs or associated struc­ tures. It is more than likely, practically certain, that many of the generalizations here offered will soon be modified or superseded by o.ther generalizations, but they will have served their purpose if they induce critical students to make a wider and more thorough study of the prob­ lems of insect morphology. Two notable books have:: appeared recently in entomology: one, "Lehrbuch der Entomologie," by Dr. Hermann Weber of Danzig, in its first edition; the other, "A General Textbook of Entomology," by Dr. A. D. Imms of Cambridge, England, in its third edition. In preparing the present text the writer has made a special effort to concur with the authors of these books in the. matter of anatomical terms, in order that students may as far as possible be spared confusion in turning from one treatise to another. Unfortunately, however, there is still much unavoid­ able discrepancy in the use and application of anatomical names in entomology. The trouble, in large measure, can be blamed on the insects themselves, since they will not entirely conform with any plan of nomenclature or with any scheme we can devise for naming their parts consistently. To make clear the meaning of terms as used in this text, therefore, a ·glossary of definitions is appended to each chapter, wherein, also, will be found the German equivalents of many of our English and Latinized technical names. In maldng acknowledgments, the writer must first of all declare his indebtedness to the Bureau of Entomology, United States Department v vi PREFACE of Agriculture, for the experience and information acquired in the course of his many years of official service. Most of the illustrations accom­ panying the text that are not accredited to particular sources are the property of the Bureau of Entomology and Plant Quarantine, and many of them have been published in the Miscellaneous Collections and tp.e" Annual Reports of the Smithsonian Institution of Washington, D.C. For the use of these figures the writer hereby expresses his thanks both to the Bureau and to the Smithsonian Institution. With regard to illustrations borrowed from other works, the writer is particularly indebted to Professor Hermann Weber, of Danzig, for permission to use figures from his "Biologie der E(emipteren" and "Lehrbuch der Entomologie." The rest of the illustrations, each accredited to its proper source in.the scientific journals, have been freely drawn from the common heritage of entomology contributed by the many workers in many lands who have devoted themselves to the study of insects. To my wife, Ruth H. Snodgrass, credit is due for the typing of the manu­ script and for much of the work of indexing and proofreading. R. E. SNODGRASS. WASHINGTON', D. C. May, 1935. CONTENTS PAGE PREFACE ••••••••••• V CHAPTER I INTRODUCTION • • • • . • • 1 CHAPTER II GENERAL ORGANIZATION AND DEVELOPMENT . 14 Glossary of Embryological Terms. 44 CHAPTER III THE BODY WALL AND ITS DERIVATIVES . 48 1. The Body Wall .......... 48 2. External Processes of the Body Wall . 55 3. Sense Organs ........... 59 4. Ectodermal Glands, Corpora Allata, Oenocytes. 60 5. Muscle Attachments on the Body Wall .. 62 6. Moulting . 64 Glossary of Terms Applied to the Body Wall. 68 CHAPTER IV Body Regions, Sclerites, and Segmentation. 70 Glossary of General Terms Applied to the Body Segments and the Skeletal Plates. 81 CHAPTER V THE SEGMENTAL ApPENDAGES OF ARTHROPODS . 83 Glossary of Terms Applied to the Appendages 98 CHAPTER VI THE HEAD .•.......•..•..... 100 1. General Morphology of the Arthropod Head. 100 2. Structu're of the Definitive Insect Head . 104 3. Special Modifications in the Structure of the Head . 118 Glossary of Terms Applied to the Head . 127 CHAPTER VII THE HEAD ApPENDAGES . 130 1. ·Preantennal Appendages. 130 2. The Antennae ~ . 131 3. The Postantennal Appendages . 133 4. The Mandibles. 133 5. The Superlinguae. 139 6. The Maxillae ... 141 7. The Labium. 145 8. Glands of the Head Appendages. 153 Glossary of Terms Applied to the Head Appendages. 155 Vll Vlll CONTENTS CHAPTER VIn PAGE THE THORAX ........ 157 1. Evolution of the Thorax .... 157 2. The Neck .......... 159 3. General Structure of the Thorax . 160 4. The Prothorax. 172 5. The Pterothorax . 173 6. The Thoracic Muscles ..... 186 Glossary of Terms Applied to the Thorax 190 CHAPTER IX THE THORACIC LEGS. 193 1. The Structure of the Legs . 193 2. Muscles and Mechanism of the Legs . 200 Glossary of Terms Applied to the Parts of an Insect's Leg 209 CHAPTER X THE WINGS ............... 211 1. Origin and Evolution of the Wings . 211 2. Development of the Wings .. 214 3. Structure of the Wings. 215 4. The Wing Muscles . 228 5. The Wing Movements ... 233 6. Insect Flight. 240 Glossary of Terms Applied to the Wings. 243 CHAPTER XI THE ABDOMEN . • • . • • . • . 246 l. General Structure of the Abdominal Segments . 247 2. The Abdominal Musculature. 257 3. The Abdominal Appendages. 267 CHAPTER XII THE ORGANS OF INGESTION. 280 l. The Preoral Cavity. 281 2. The Cephalic Stomadaeum. 284 3. The Feeding Mechanism of N europtera and Coleoptera. 286 4. The Feeding Mechanism of Hymenoptera . 295 5. The Feeding Mechanism of Lepidoptera. 302 6. The Feeding Mechanism of Diptera .... 311 7. The Mouth Parts of Siphonaptera . 325 8. The Feeding Mechanism of Thysanoptera. 326 9. The Feeding Mechanism of Hemiptera . 328 10. The Feeding Mechanism of Anoplura. 344 CHAPTER XIII THE ALIMENTARY CANAL ........... 347 l. Development of the Alimentary Canal . 347 2. General Structure of the Alimentary Canal 348 3. The Stomodaeum. 349 4. The Mesenteron . 359 5. The Proctodaeum. 374 CONTENTS ix PAGE 6. The Filter Chamber. 383 Glossary of Terms Applied to the Alimentary Canal. 387 CHAPTER XIV THE ORGANS OF DISTRIBUTION, CONSERVATION, AND ELIMINATION. 389 1. The Blood. 389 2. The Organs of Circulation. 397 3. The Fat-body . 407 4. The Oenocytes. 410 5. The Corpora allata. 411 6. The Organs of Elimination. 413 Glossary of Terms Used in This Chapter. 420 CHAPTER XV .. THE RESPIRATORY SYSTEM ......... 422 1. The Integument as a Respiratory Organ. 423 2. Blood Gills . 424 3. The Tracheal System. , 425 4. General Mechanism of Tracheal Respiration. 452 Glossary of Terms Applied to the Respiratory System. 461 CHAPTER XVI THE NERVOUS SYSTEM. ..... 464 1. General Structure, Organization, and Function of the Nervous System. 465 2. The Central Nervous System ... 472 3. The Stomodaeal Nervous System. 501 4. The Peripheral Nervous System . .. 503 Glossary of Terms Applied to the Nervous System 507 CHAPTER XVII THE SENSE ORGANS ..................... 510 1. General Structure and Ciassification of Insect Sense Organs . 512 2. The Hair Organs. 515 3. T'he Campaniform Organs. 521 4. The Plate Organs ..... 523 5. The Scolopophorous Organs . 525 6. The Eyes ........ 528 Glossary of Terms Applied to the Sense Organs. 548 CHAPTER XVIII THE INTERNAL ORGANS OF REPRODUCTION. 550 1. Thc Female Organs. , . 552 2. The Male Organs. ..... 567 3. General Morphology of the Reproductive Organs. 573 Glossary of Terms Appli~d to the Internal Reproductive Organs 578 CHAPTER XIX THE ORGANS OF COPULATION AND OVIPOSITION . 581 1. The Male Genitalia. 582 2. The Female Genitalia. 607 Glossary of Terms Applied to the External Genitalia 620 REFERENCES . 625 INDEX( ..... 647 PRINCIPLES OF INSECT MORPHOLOGY CHAPTER I INTRODUCTION Morphology, in the biological sense, is the science of form in living organisIns. Anatomy is the determination of structural facts. Morphol­ ogy seeks to find the reason for structure, and to understand the relation of different structural forms to one another. Morphology, therefore, must be intimate with function, since it must see forms as plastic physical adaptations to the work to be performed. A few physiological functions are basic to all organisms; they are essential to the continuance of matter in a living state. The various structural types of organisms are special ways of accomplishing these functions, that is, for doing the same things in different ways or under different circumstances. Some represent improvements in the machinery along established lines; others represent changes or new ideas developed along new and divergent lines. The morphologist, therefore, though primarily a comparative anatomist, in order properly to develop his subject, must give attention to the working of the physical mechanisms with which he deals in his aI!.atomical studies, he must look for the significance of structural modifications Rnd innova­ tions, and he must understand the basic physiological functions that .underlie organic form.
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