Springer Se ries in Microbiology

Editor: Mortimer P. Starr

Armand Maggenti

General Nematology

With 135 Figures

Springer-Verlag New York Heidelberg Berlin

Armand Maggenti Department of Nematology University of California College of Agricultural and Environmental Sciences Davis, California 95616 U.S.A.

Series Editor: Mortimer P. Starr Department of Bacteriology University of California Davis, California 95616 U.S.A.

Front cover illustration. Left: Heth, a parasite of myriapods and diplopods. The genus was first described by Cobb in 1898. Right: A composite, three-dimensional drawing of a plant parasitic nematode.

Library of Congress Cataloging in Publication Data

Maggenti, Armand R. General nematology.

(Springer series in microbiology) Bibliography: p. Inc1udes index. 1. N ematoda. I. Title. 11. Series.

QL391.N4M213 595.1'82 81-8863 AACR2

© 1981 by Springer-Verlag New York, Inc. Softcover reprint of the hardcover 1 st edition 1981

All rights reserved. No part of this book may be translated or reproduced in any form without written permission from Springer-Verlag, 175 Fifth Avenue, New York, New York 10010, U.S.A.

Tbe use of general descriptive names, trade names, trademarks, etc. in this pub­lication, even if the former are not especially identified, is not to be taken as a sign that such names, as understood by the Trade Marks and Merchandise Marks Act, may accordingly be used freely by anyone.

9 8 7 6 5 432 1

ISBN-13: 978-1-4612-5940-4 e-ISBN-13: 978-1-4612-5938-1 001: 10.1007/978-1-4612-5938-1

To Mary Ann

Preface

This text is an overall view of nematology because I believe the science should be treated as a unified discipline. The differences in the biological habits of nematodes do not justify the separation of plant nematologists and animal nematologists, since the separation is not a reflection of any difIerences inherent to nematodes. Therefore, the book is arranged with a format that in the beginning chapters illustrates the similarities and se­quence of development of morphological characters among nematodes regardless of their biological habits. The later chapters illustrate the in­tegration of the evolutionary development of the parasitie habit from related free-living forms.

Nematology is probably the last major discipline to establish its inde­pendence from the parent science of zoology. This natural evolvement of nematology has occurred because of the overwhelming accumulation of sophisticated information and research that reflects the unique relation­ships of nematodes to other forms of plant and animal life as weH as their relationships in other facets of the environment. N ematodes are inverte­brate animals that, like insects, are unusual in their great numbers and varieties, their smaH size (generally microscopic), their high degree of internal organization, and their virtually ubiquitous distribution. They oc­cupy almost every ecological niehe, often causing disease of humans, other animals, and plants. These activities often result in debility, death, or in the impairment and loss of food supply with consequent loss to producers and consumers.

Hopefully tbis book will intrigue teachers, students, nematologists, plant pathologists, parasitologists, and zoologists. Each will approach the book from their own level of needs; some will read it superficially, some will delve into its speculation, and all, I hope, will learn to appreciate the science itself.

viii Preface

In presenting my understanding of nematology, I hope to comply with the admonition of Thomas Huxley: "My business is to te ach my aspira­tions to conform themselves to fact, not to try and make facts harmonize with my aspirations .... Sit down before fact as a little child, be prepared to give up every preconceived notion, follow humbly wherever and to what­ever abysses nature leads, or you shall learn nothing." May the reader approach my effort with the same attitude.

I wish to acknowledge but not blame for this effort: D. J. Raski, who first introduced me to nematology; M. W. Allen for his teaching; and more immediately my wife Mary Ann to whom the book is dedicated, who put aside her own work to bring this book to fruition; W. H. Hart, Nahum Marban-Mendoza, and EUa Mae Noffsinger for their reviews and com­ments; Gaylen Paxman, nematologist, librarian, critic, and friend; my graduate students Fawzia Adbel-Rahman and Steve Martinez, who were interested and involved; all my students in Introductory Nematology, on whom the concepts presented here were first tried; R. Giblin; I. Cid deI Prado; and especially my son Peter, who in critical moments eased the pressure by preparing penciI sketches for inking. My obligation is un­bounded to B. G. Chitwood. He is a constant stimulus and his memory an abi ding inspiration.

In 1967 B. G. Chitwood offered me the opportunity to revise the classic An Introduction to Nematology (B. G. and M. B. Chitwood). I feit then and I still beIieve that classic should stand. To modify it is to redo the Mona Lisa in crayons. I urge all students of nematology to read, read, and reread Chitwood. To have known the man was a pleasure, to have had hirn as amentor exasperating, to have had hirn as a friend magnificent.

This is a short preface because the purpose of the book is to read it. Enjoy now these magnificent animals as I have.

Davis, California July 1981

ARMAND MAGGENTI

Contents

CHAPTER 1 History of the Science 1

I. Introduction ........................................ 1 H. Ancient Times to the Eighteenth Century . . . . . . . . . . . . . . . . . 2

IH. History and Development in Nineteenth and Twentieth Century Europe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

IV. History and Development in America . . . . . . . . . . . . . . . . . . . . . 7

CHAPTER 2 Nematodes and Their Allies 9

I. Phylum Rotifera ..................................... 12 H. Phylum Gastrotricha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

IH. Phylum Kinorhyncha ................................. 23 IV. Phylum Nematomorpha ............................... 27 V. Phylum Nemata ..................................... 32

CHAPTER 3 Nematode Integument. . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 42

I. External Cutic1e ..................................... 43 11. Internal Body Cutic1e ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 46

IH. Cuticular Structures .................................. 48 IV. Hypodermis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 68 v. Excretory System. . . . . .. . . .. . . . . . . . . . . . . . . . . . . .. .. . .. 74

VI. Molting ............................................ 80

CHAPTER4

Internal Morphology 86

I. Somatic Musculature ................................. 86

x Contents

II. Alimentary Canal .................................... 92 III. Nervous System ...................................... 122

CHAPTER 5 Reproductive System 131

I. Introduction ........................................ 131 11. Female Reproductive System .................. . . . . . . . .. 135

111. Male Reproductive System ............................. 142 IV. Spermatogenesis ..................................... 147 V. Oogenesis .......................................... 151

VI. Embryology ........................................ 154 VII. Postembryonie Changes ............................... 156

CHAPTER 6 Plant Parasitism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 158

I. Adenophorean Plant Parasitism ... . . . . . . . . . . . . . . . . . . . . .. 160 11. Secementean Plant Parasitism .......................... 165

CHAPTER 7 Invertebrate Parasitism and Other Associations 218

I. Introduction ........................................ 218 11. Facultative Parasitism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 221

111. Obligate Body Cavity and Tissue Parasites ................ 225 IV. Essential Nonparasitic Associations Inc1uding Vectors of

Insect Diseases ...................................... 241 V. Obligate Associations of the Alimentary Tract of

Invertebrates ........................................ 244

CHAPTER 8 Vertebrate Parasitism 245

I. Introduction ........................................ 245 II. Adenophorean Parasites of Vertebrates ................... 251

111. Secementean Parasites of Vertebrates .................... 258

CHAPTER 9 Classification of Nemata ................................... 305

I. Oassification of Nemata ............................... 306

Selected References ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 347

Index .................................................. 363