What would happen to termites if trichonympha disappeared

Biology

Malaria remains a true scourge of humankind, with perhaps the greatest social and economic impact of any parasitic disease; the cover art for this edition of FOUNDATIONS OF PARASITOLOGY reflects this situation. Anopheles species capable of transmitting malaria are common throughout much of the world, especially in the tropics and sub-tropics, and their presence, coupled with living conditions that expose people to their bites both day and night, virtually ensures that human populations in these regions will be at risk. Our cover design is intended to convey the idea that vectors are the primary factor in sustaining risk of acquiring many infections, but as every parasitology student either knows or soon learns, complex life cycles also are common among parasites. Terefore, disease control efforts can focus on any stage that is vulnerable to disruption, and that claim is true for any parasite species with a complex life cycle. Finally, ecological settings in which vectors thrive and in which humans encounter both vectors and parasites are crucial to the maintenance of risk, regardless of the disease. Our cover is thus a reminder of the multi-faceted lives of many parasites, especially those that encounter various host tissues during development. Te mosquito on our cover is Anopheles feeborni, a New World species; the other figures include, counterclockwise from the bottom, a pre-erythrocytic schizont, ring stages in a multiply-infected erythrocyte, an erythrocytic schizont, gametocytes, and oocysts on the gut of an experimentally infected mosquito. For parasitology students wishing to explore the biology of malarial parasites on the Internet, we recommend the Malaria Atlas Project site (http://www.map.ox.ac.uk/)

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Enlargement of liver and spleen (hepatospleenomegaly) in a laboratory mouse infected with the rodent malaria parasite, Plasmodium berghei . The liver and spleen enlarge and darken from the accumulation of parasite hemozoin pigment granules in reticuloendothelial cells of these organs (p. 153). Hepatospleenomegaly can also occur in humans infected with their malaria parasites. The mouse- P. berghei system, like other non-human models of parasitic diseases, has been important in improving understanding of the complex interactions between hosts and parasites.

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ninth edition

Gerald D. Schmidt & Larry S. Roberts’

foundations of parasitology

larry s. roberts Texas tech university emeritus

john janovy, jr. university of nebraska–lincoln

steve nadler university of california, davis

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GERALD D. SCHMIDT & LARRY S. ROBERTS’ FOUNDATIONS OF PARASITOLOGY NINTH EDITION

Published by McGraw-Hill, a business unit of The McGraw-Hill Companies, Inc., 1221 Avenue of the Americas, New York, NY 10020. Copyright © 2013 by The McGraw-Hill Companies, Inc. All rights reserved. Previous editions © 2009, 2005, 2000, and 1996. No part of this publication may be reproduced or distributed in any form or by any means, or stored in a database or retrieval system, without the prior written consent of The McGraw-Hill Companies, Inc., including, but not limited to, in any network or other electronic storage or transmission, or broadcast for distance learning.

Some ancillaries, including electronic and print components, may not be available to customers outside the United States.

This book is printed on acid-free paper.

1 2 3 4 5 6 7 8 9 0 QPD/QPD 0 9 8

ISBN 978–0–07–352419–1 MHID 0–07–352419–0

Senior Vice President, Product & Markets: Kurt L. Strand Vice President, Content Production & Technology Services: Kimberly Meriwether David Publisher: Michael Hackett Brand Manager: Rebecca Olson Director of Development: Elizabeth Sievers Marketing Manager: Patrick Reidy Director, Content Production: Terri Schiesl Senior Project Manager: Joyce Watters Buyer: Nichole Birkenholz Media Project Manager: Prashanthi Nadipalli Cover Designer: Studio Montage, St. Louis, MO. Cover Image: Mosquito photo by James Gathany (CDC, used by permission); background scene, pre-erythrocytic stages, and mosquito gut by John Janovy, Jr.; ring stages, schizont, and gametocytes from Plate 3, by A. Wilcox, U. S. Government Printing Office, Washington, D.C., 1960. Typeface: 9.5/11 Times Roman Compositor: S4 Carlisle Publishing Services Printer: Quebecor World Dubuque, IA

Library of Congress Cataloging-in-Publication Data

Roberts, Larry S., 1935-

Foundations of parasitology / Larry S. Roberts, Texas Tech University, Emeritus, John Janovy, Jr., University of Nebraska-Lincoln, Steve Nadler, University of California, Davis. —Ninth edition.

pages cm

Includes index.

ISBN 978–0–07–352419–1 — ISBN 0–07–352419–0 1. Parasitology. I. Janovy, John, 1937—II. Nadler, Steve. III. Roberts, Larry S, 1935—Gerald D. Schmidt & Larry S. Roberts’ foundations of parasitology. IV. Title.

QL757.R585 2013

616.9'6—dc23 2012027864

www.mhhe.com

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v

LARRY S. ROBERTS

Larry S. Roberts, professor emeritus of biology at Texas Tech University, was profes- sor of zoology at University of Massachusetts, Amherst, and was adjunct professor of biol- ogy at Florida International University and the University of Miami, where he had exten- sive experience teaching para- sitology, invertebrate zoology, marine biology, and develop- mental biology. He received his Sc.D. in parasitology at the Johns Hopkins University and has coauthored Foundations of Parasitology from the first edition through this, the ninth edition. He is also coauthor of Integrated Principles of Zoology, Biology of Animals, and Animal Diversity, and is author of The Underwater World of Sport Diving.

Dr. Roberts has published many research articles and reviews. He has served as president of the American Society of Parasitologists, the Southwestern Society of Parasitologists, the Southeastern Society of Parasitologists, and the Helminthological Society of Washington. He received the Henry Baldwin Ward Medal from the American Society of Parasitologists. His hobbies include scuba diving, underwater photography, and tropical horticulture.

Dr. Roberts can be contacted at Lroberts1@compuserve .com

JOHN JANOVY, JR.

John Janovy, Jr . (PhD University of Oklahoma, 1965) is Professor Emeritus at the University of Nebraska where he was the Paula and D. B. Varner Distinguished Professor of Biological Sciences for much of his career. His re- search interest is parasitol- ogy, with particular focus on parasite ecology and life cycles. He has been di- rector of the Cedar Point Biological Station, interim director of the University of Nebraska State Museum, an assistant dean of Arts and Sciences, and secretary-treasurer of the American Society of Parasitologists. He is currently (2012) President-Elect

ab out the authors

of the American Society of Parasitologists. His scholarly and cre ative accomplishments consist of approximately 100 scientific papers and book chapters; 14 books, including Keith County Journal, On Becoming a Biologist, Teaching in Eden, Outwitting College Professors , and Foundations of Parasitology (with Larry Roberts and Steve Nadler); the screenplay for the televised version of Keith County Journal (Nebraska Public Television); and numerous popular articles. His teaching experiences include almost continuous ser- vice in the large- enrollment freshman biology course; Field Parasitology (BIOS 487/887) at the Cedar Point Biological Station; Invertebrate Zoology (BIOS 381); Parasitology (BIOS 385); a decade in BIOS 103/204 (Organismic Biology/ Biodiversity); and numerous honors seminars. He has super- vised 18 MS students, 14 PhD students, and approximately 50 undergraduate researchers, including 10 Howard Hughes scholars. His honors include the University of Nebraska Distinguished Teaching Award (1970), University Honors Program Master Lecturer (1986), American Health magazine book award (1987, for Fields of Friendly Strife ), University of Nebraska Outstanding Research and Creativity Award (1998), The Nature Conservancy Hero recognition (2000), and the American Society of Parasitologists Clark P. Read Mentorship Award (2003).

GERALD D. SCHMIDT

Gerald D. Schmidt was professor of biology at the Univers i ty of Northern Colorado (UNC) when he passed away. He received his PhD from Colorado State University. He was active in research and promoting re- search activities at UNC, and he published more than 160 research articles in scien- tific journals, as well as six books. He received awards from UNC for outstand- ing teaching and for distin- guished scholarship. He was a board member of the World Federation of Parasitologists; a Fellow of the Royal Society of Tropical Medicine and Hygiene, London; and a Fellow of the Royal Society of South Australia.

Dr. Schmidt served the American Society of Parasitologists as secretary-treasurer for seven years. He was co-author of Foundations of Parasitology through the first four editions. His hobbies were hunting and fishing, espe- cially fishing, and he wrote a book on fishing. Dr. Schmidt died on 16 October 1990; many more details of his life can be found in the Journal of Parasitology, 78:757–773.

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vi About the Authors

six years. Dr. Nadler was an associate editor of the Journal of Parasitology, and president of the American Society of Parasitologists (2007–08). His scholarly accomplish- ments include approximately 90 scientific papers, and his research has been supported by grants from the U.S. National Science Foundation and the National Institutes of Health. He currently serves on the editorial boards of the journals Parasitology, Systematic Parasitology, Zookeys, and Animal Cells and Systems. His research laboratory is supported by efforts of undergraduate and graduate students, along with visiting scientists and postdoctoral scholars. At UC Davis his undergraduate and graduate teaching includes courses in parasitology, nematology, and molecular phylogenetics.

STEVE NADLER

Steve Nadler (PhD in Medical Parasitology, Louisiana State University Medical Center, New Orleans) is Professor of Nematology in the Depart- ment of Entomology and Nematology at the University of California, Davis. His re- search interests concentrate on the systematics and evolution- ary biology of nematodes, in- cluding both free-living and parasitic species. He served as chair of the Department of Nematology at UC Davis for

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23 Nematodes: Trichinellida and Dioctophymatida, Enoplean Parasites 377

24 Nematodes: Tylenchina, a Functionally Diverse Clade 391

25 Nematodes: Rhabditomorpha, Bursate Roundworms 397

26 Nematodes: Ascaridomorpha, Intestinal Large Roundworms 411

27 Nematodes: Oxyuridomorpha, Pinworms 425

28 Nematodes: Gnathostomatomorpha and Spiruromorpha, a Potpourri 431

29 Nematodes: Filarioidea: Filarial Worms 441

30 Nematodes: Dracunculomorpha, Guinea Worms, and Others 457

31 Phylum Nematomorpha, Hairworms 465

32 Phylum Acanthocephala: Thorny-Headed Worms 473

33 Phylum Arthropoda: Form, Function, and Classification 489

34 Parasitic Crustaceans 513

35 Pentastomida: Tongue Worms 535

36 Parasitic Insects: Phthiraptera, Chewing and Sucking Lice 543

37 Parasitic Insects: Hemiptera, Bugs 555

38 Parasitic Insects: Fleas, Order Siphonaptera 563

39 Parasitic Insects: Diptera, Flies 575

40 Parasitic Insects: Strepsiptera, Hymenoptera, and Others 599

41 Parasitic Arachnids: Subclass Acari, Ticks and Mites 611

Glossary 631

Index 653

Preface xv

1 Introduction to Parasitology 1

2 Basic Principles and Concepts I: Parasite Systematics, Ecology, and Evolution 9

3 Basic Principles and Concepts II: Immunology and Pathology 23

4 Parasitic Protozoa: Form, Function, and Classification 41

5 Kinetoplasta: Trypanosomes and Their Kin 61

6 Other Flagellated Protozoa 87

7 The Amebas 105

8 Phylum Apicomplexa: Gregarines, Coccidia, and Related Organisms 119

9 Phylum Apicomplexa: Malaria Organisms and Piroplasms 143

10 Phylum Ciliophora: Ciliated Protistan Parasites 167

11 Microsporidia and Myxozoa: Parasites with Polar Filaments 175

12 The Mesozoa: Pioneers or Degenerates? 185

13 Introduction to Phylum Platyhelminthes 191

14 Trematoda: Aspidobothrea 201

15 Trematoda: Form, Function, and Classification of Digeneans 209

16 Digeneans: Strigeiformes 235

17 Digeneans: Echinostomatiformes 253

18 Digeneans: Plagiorchiformes and Opisthorchiformes 265

19 Monogenoidea 283

20 Cestoidea: Form, Function, and Classification of Tapeworms 299

21 Tapeworms 325

22 Phylum Nematoda: Form, Function, and Classification 349

br i e f contents

vii

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Antibodies 28 Lymphocytes 29 Subsets of T Cells 29 T-Cell Receptors 30 Generation of a Humoral Response 30 Cell-Mediated Response 32 Inflammation 32 Acquired Immune Deficiency

Syndrome (AIDS) 34 Immunodiagnosis 34 Pathogenesis of Parasitic Infections 35 Accommodation and Tolerance in

the Host-Parasite Relationship 37 The Microbial Deprivation Hypothesis 38 Overview 38

Learning Outcomes 40

References 40

Additional Readings 40

4 Parasitic Protozoa: Form, Function, and Classification 41

Form and Function 41 Nucleus and Cytoplasm 42 Locomotor Organelles 44 Reproduction and Life Cycles 48 Encystment 50 Feeding and Metabolism 51 Excretion and Osmoregulation 52 Endosymbionts 52 Classification of Protozoan Phyla 52 Characters Generally Shared by Amebas 55 Stramenopiles 56

Learning Outcomes 58

References 59

Additional Readings 59

5 Kinetoplasta: Trypanosomes and Their Kin 61

Forms of Trypanosomatidae 61 Genus Trypanosoma 64 Section Salivaria 65 Section Stercoraria 71 Genus Leishmania 77 Cutaneous Leishmaniasis 79 Visceral Leishmaniasis 83 Other Trypanosomatid Parasites 85

Learning Outcomes 86

References 86

Additional Readings 86

viii

c ontents

Preface xv

1 Introduction to Parasitology 1 Relationship of Parasitology to Other

Sciences 1 Some Basic Definitions 2 Interactions of Symbionts 2 Parasitology and Human Welfare 4 Parasites of Domestic and Wild Animals 6 Parasitology for Fun and Profit 7 Careers in Parasitology 7

References 8

Additional Readings 8

Parasitology on the World Wide Web 8

2 Basic Principles and Concepts I: Parasite Systematics, Ecology, and Evolution 9

Systematics and Taxonomy of Parasites 9 Parasite Ecology 10 The Host as an Environment 10 A Parasite’s Ecological Niche 10 Parasite Populations 12 Trophic Relationships 14 Adaptations for Transmission 15 Epidemiology and Transmission Ecology 17 Theoretical Parasitology 18 Parasite Evolution 18 Evolutionary Associations Between Parasites

and Hosts 18 Parasitism and Sexual Selection 19 Evolution of Virulence 21

Learning Outcomes 21

References 21

Additional Readings 21

3 Basic Principles and Concepts II: Immunology and Pathology 23

Susceptibility and Resistance 24 Innate Defense Mechanisms 24 Cell Signaling 24 Cellular Defenses: Phagocytosis 27 Adaptive Immune Response

of Vertebrates 28 Basis of Self and Nonself Recognition

in Responses 28

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6 Other Flagellated Protozoa 87 Order Retortamonadida 87 Family Retortamonadidae 87 Order Diplomonadida 88 Family Hexamitidae 88 Genus Giardia 88 Trichomonads (Class Trichomonada, Order

Trichomonadida) 93 Family Trichomonadidae 93 Family Monocercomonadidae 98 Order Hypermastigida 101 Order Opalinida 101 Family Opalinidae 101

Learning Outcomes 103

References 103

Additional Readings 103

7 The Amebas 105 Amebas Infecting Mouth and Intestine 105 Family Entamoebidae 105 Genus Iodamoeba 113 Amebas Infecting Brain and Eyes 114 Family Vahlkampfiidae 114 Family Acanthamoebidae 116 Amebas of Uncertain Affinities 117

Learning Outcomes 118

References 118

Additional Readings 118

8 Phylum Apicomplexa: Gregarines, Coccidia, and Related Organisms 119

Apicomplexan Structure 119 Class Conoidasida, Subclass

Gregarinasina 120 Order Eugregarinorida 121 Gregarine-Like Apicomplexans:

Cryptosporidium Species 122 Subclass Coccidiasina 124 Order Eucoccidiorida 124 Suborder Adeleorina 124 Suborder Eimeriorina 125

Learning Outcomes 141

References 141

Additional Readings 141

9 Phylum Apicomplexa: Malaria Organisms and Piroplasms 143

Order Haemospororida 143 Genus Plasmodium 143 Genus Haemoproteus 159 Genus Leucocytozoon 160

Order Piroplasmida 160 Family Babesiidae 161 Family Theileriidae 164

Learning Outcomes 165

References 165

Additional Readings 165

10 Phylum Ciliophora: Ciliated Protistan Parasites 167

Class Spirotrichea 167 Order Clevelandellida; Family Nyctotheridae 167 Class Litostomatea 168 Order Vestibuliferida, Family Balantidiidae 168 Order Entodiniomorphida 169 Class Oligohymenophorea 170 Subclass Hymenostomatia, Order Hymenostomatida,

Family Ichthyophthiriidae 170 Subclass Peritrichia 170 Order Sessilida 170 Order Mobilida, Family Trichodinidae 172

Learning Outcomes 173

References 173

Additional Readings 173

11 Microsporidia and Myxozoa: Parasites with Polar Filaments 175

Phylum Microsporidia 175 Family Nosematidae 177 Other Microsporidian Species 177 Epidemiology and Zoonotic Potential 178 Myxozoa 178 Family Myxobolidae 179

Learning Outcomes 184

References 184

Additional Readings 184

12 The Mesozoa: Pioneers or Degenerates? 185

Phylum Dicyemida 185 Class Rhombozoa 185 Phylum Orthonectida 187 Class Orthonectida 187 Phylogenetic Position 188 Host-Parasite Relationships 189

Learning Outcomes 190

References 190

Additional Readings 190

13 Introduction to Phylum Platyhelminthes 191

Platyhelminth Systematics 192 Turbellarians 196

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x Contents

Acoels 196 Rhabditophorans 196 Temnocephalideans 197 Alloeocoels 197 Tricladids 197 Polycladids 198

Learning Outcomes 199

References 199

Additional Readings 199

14 Trematoda: Aspidobothrea 201 Form and Function 201 Body Form 201 Tegument 201 Digestive System 202 Osmoregulatory System 202 Nervous System 202 Reproductive Systems 203 Development 204 Aspidogaster conchicola 206 Rugogaster hydrolagi 207 Stichocotyle nephropsis 207 Phylogenetic Considerations 207

Learning Outcomes 208

References 208

Additional Readings 208

15 Trematoda: Form, Function, and Classification of Digeneans 209

Form and Function 209 Body Form 209 Tegument 210 Muscular System 213 Nervous System 214 Excretion and Osmoregulation 215 Acquisition of Nutrients and Digestion 217 Reproductive Systems 218 Development 219 Embryogenesis 220 Larval and Juvenile Development 220 Development in a Definitive Host 225 Trematode Transitions 226 Summary of Life Cycle 227 Metabolism 227 Energy Metabolism 227 Synthetic Metabolism 230 Biochemistry of Trematode Tegument 230 Phylogeny of Digenetic

Trematodes 230 Learning Outcomes 233

References 233

Additional Readings 233

16 Digeneans: Strigeiformes 235 Superfamily Strigeoidea 235 Family Diplostomidae 235 Family Strigeidae 236 Superfamily Schistosomatoidea 237 Family Schistosomatidae: Schistosoma Species

and Schistosomiasis 238 Control 248

Learning Outcomes 251

References 251

Additional Readings 251

17 Digeneans: Echinostomatiformes 253 Superfamily Echinostomatoidea 253 Family Echinostomatidae 253 Echinostomatids as Models in Experimental

Parasitology 255 Family Fasciolidae 256 Other Fasciolid Trematodes 259 Family Cathaemasiidae 261 Superfamily Paramphistomoidea 262 Family Paramphistomidae 262 Family Diplodiscidae 262 Family Gastrodiscidae 262

Learning Outcomes 263

References 263

Additional Readings 263

18 Digeneans: Plagiorchiformes and Opisthorchiformes 265

Order Plagiorchiformes 265 Suborder Plagiorchiata 265 Suborder Troglotrematata 269 Order Opisthorchiformes 275 Family Opisthorchiidae 275 Family Heterophyidae 279

Learning Outcomes 280

References 280

Additional Readings 281

19 Monogenoidea 283 Form and Function 284 Body Form 284 Tegument 285 Muscular and Nervous Systems 286 Osmoregulatory System 287 Acquisition of Nutrients 289 Male Reproductive System 289 Female Reproductive System 290 Development 291 Oncomiracidium 291

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Contents xi

Subclass Polyonchoinea 292 Subclass Polystomatoinea 294 Subclass Oligonchoinea 294 Phylogeny 295 Classification of Class Monogenoidea 296

Learning Outcomes 297

References 297

Additional Readings 297

20 Cestoidea: Form, Function, and Classification of Tapeworms 299

Form and Function 299 Strobila 299 Scolex 300 Tegument 301 Calcareous Corpuscles 305 Muscular System 306 Nervous System 307 Excretion and Osmoregulation 307 Reproductive Systems 310 Development 312 Larval and Juvenile Development 313 Effects of Metacestodes on Hosts 315 Development in Definitive Hosts 316 Metabolism 317 Acquisition of Nutrients 317 Energy Metabolism 318 Synthetic Metabolism 320 Hormonal Effects of Metabolites 320 Classification of Class Cestoidea 321

Learning Outcomes 323

References 323

Additional Readings 323

21 Tapeworms 325 Order Diphyllobothriidea 325 Family Diphyllobothriidae 325 Diphyllobothrium Species 325 Other Diphyllobothriideans Found in Humans 329 Sparganosis 329 Order Caryophyllidea 329 Order Spathebothriidea 330 Order Cyclophyllidea 330 Family Taeniidae 330 Other Taeniids of Medical Importance 335 Family Hymenolepididae 340 Family Davaineidae 342 Family Dilepididae 342 Family Anoplocephalidae 343 Family Mesocestoididae 343 Family Dioecocestidae 344 Order Proteocephalata 344 Order Tetraphyllidea 345

Order Trypanorhyncha 345 Subcohort Amphilinidea 347 Cohort Gyrocotylidea 347

Learning Outcomes 347

References 348

Additional Readings 348

22 Phylum Nematoda: Form, Function, and Classification 349

Historical Aspects 349 Form and Function 350 Body Wall 350 Musculature 352 Pseudocoel and Hydrostatic Skeleton 353 Nervous System 355 Digestive System and Acquisition

of Nutrients 359 Secretory-Excretory System 362 Reproduction 363 Development 367 Eggshell Formation 367 Embryogenesis 368 Embryonic Metabolism 369 Hatching 369 Growth and Ecdysis 370 Metabolism 371 Energy Metabolism 371 Synthetic Metabolism 372 Classification of Phylum Nematoda 373

Learning Outcomes 376

References 376

Additional Readings 376

23 Nematodes: Trichinellida and Dioctophymatida, Enoplean Parasites 377

Order Trichinellida 377 Family Trichuridae 377 Family Capillariidae 380 Family Anatrichosomatidae 381 Family Trichinellidae 381 Order Dioctophymatida 388 Family Dioctophymatidae 388

Learning Outcomes 390

References 390

Additional Readings 390

24 Nematodes: Tylenchina, a Functionally Diverse Clade 391

Family Steinernematidae 391 Family Rhabdiasidae 392 Family Strongyloididae 393 Strongyloides Species 393

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Learning Outcomes 396

References 396

Additional Readings 396

25 Nematodes: Rhabditomorpha, Bursate Roundworms 397

Family Ancylostomatidae 397 Family Strongylidae 405 Family Syngamidae 406 Family Trichostrongylidae 406 Family Dictyocaulidae 408 Other Trichostrongyles 408 Metastrongyles 408 Family Angiostrongylidae 408

Learning Outcomes 410

References 410

Additional Readings 410

26 Nematodes: Ascaridomorpha, Intestinal Large Roundworms 411

Superfamily Ascaridoidea 411 Family Ascarididae 411 Family Anisakidae 420 Superfamily Heterakoidea 421 Family Ascaridiidae 421 Family Heterakidae 422

Learning Outcomes 423

References 423

Additional Readings 423

27 Nematodes: Oxyuridomorpha, Pinworms 425

Family Oxyuridae 425 Rodent Pinworms 428

Learning Outcomes 429

References 429

Additional Readings 429

28 Nematodes: Gnathostomatomorpha and Spiruromorpha, a Potpourri 431

Gnathostomatomorpha Family Gnathostomatidae 431

Spiruromorpha 433 Family Acuariidae 434 Family Physalopteridae 434 Family Tetrameridae 435 Family Gongylonematidae 436 Family Spirocercidae 437 Family Thelaziidae 438