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CONTENTS 1 INTRODUCTION TO PARASITOLOGY Relationship of Parasitology to Other Sciences 1 Parasitology and Human Welfare 2 Parasites of Domestic and Wild Animals 3 Careers in Parasitology 4 Some Basic Definitions 4 Interactions of Symbionts 5 Hosts 7 References 7 Additional References 8 Parasitology on the World Wide Web 8 2 BASIC PRINCIPLES AND CONCEPTS I: PARASITE ECOLOGY AND EVOLUTION Parasite Ecology 9 The Host as an Environment 9 Trophic Relationships 9 The Parasite's Ecological Niche 10 Parasite Populations 11 Parasite Reproduction 13 Adaptations for Transmission 14 Epidemiology 14 Mathematical Models 15 Parasite Evolution 16 Evolutionary Associations Between Parasites and Hosts 16 Parasitism and Sexual Selection 18 Parasite Life History 18 Systematics and Taxonomy of Parasites 18 References 19 Additional References 20 3 BASIC PRINCIPLES AND CONCEPTS II: IMMUNOLOGY AND PATHOLOGY Susceptibility and Resistance 21 Innate Defense Mechanisms 21 Physical and Chemical Barriers 21 Cellular Defenses: Phagocytosis 22 Acquired Immune Response in Vertebrates 22 Basis of Self and Nonself Recognition 23 Lymphocytes 24 Cytokines 25 Generation of a Humoral Response 25 The Cell-Mediated Response 26 Inflammation 26 Acquired Immune Deficiency Syndrome (AIDS) 28 Danger Hypothesis 29 Immunity in Invertebrates 29 Immunodiagnosis 30 Pathogenesis of Parasitic Infections 32 Accommodation and Tolerance in the Host-Parasite Relationship 33 References 35 Additional References 36 4 PARASITIC PROTOZOA: FORM, FUNCTION, AND CLASSIFICATION Form and Function 37 Nucleus and Cytoplasm 38 Locomotor Organelles 40 Reproduction and Life Cycles 44 Encystment 46 Feeding and Metabolism 46 Excretion and Osmoregulation 48 Endosymbionts 48 Classification of the Protozoan Phyla 48 References 54 Additional References 54 5 KINETOPLASTA: TRYPANOSOMES AND THEIR KINAA55 Forms of the Trypanosomatidae 55 Genus Trypanosoma 57 References 78 Additional References 81 6 OTHER FLAGELLATED PROTOZOA Order Retortamonadida 83 Family Retortamonadidae 83 Order Diplomonadida 84 Family Hexamitidae 84 Order Trichomonadida 89 Family Trichomonadidae 89 Family Monocercomonadidae 93 Order Opalinida 96 Family Opalinidae 96 References 97 Additional References 99 7 THE AMEBASAA101 Order Amoebida 101 Family Entamoebidae 101 Order Schizopyrenida 110 Family Vahlkamphidae 110 Family Hartmannellidae 111 Family Leptomyxiidae 113 References 113 ADDITIONAL REFERENCESAA115 8 PHYLUM APICOMPLEXA: GREGARINES, COCCIDIA, AND RELATED ORGANISMS The Apical Complex 117 Class Gregarinea 118 Order Eugregarinida 119 Class Coccidea 121 Order Adeleida 121 Order Eimerida 121 References 137 Additional References 139 9 PHYLUM APICOMPLEXA: MALARIA ORGANISMS AND PIROPLASMS Order Haemosporida 141 Genus Plasmodium 141 Genus Haemoproteus 156 Genus Leucocytozoon 157 Order Piroplasmida 158 Family Babesiidae 159 Family Theileriidae 161 References 162 Additional References 165 10 PHYLUM CILIOPHORA: CILIATED PROTISTAN PARASITES Class Spirotrichea 167 Subclass Heterotrichia, Order Clevelandellida, Family Nyctotheridae 167 Class Litostomatea 167 Subclass Trichostomatia, Order Vestibuliferida, Family Balantidiidae 167 Order Entodiniomorphida 169 Class Oligohymenophorea 169 Subclass Hymenostomatia, Order Hymenostomatida, Family Ichthyophthiriidae 169 Subclass Peritrichia, Order Mobilida, Family Trichodinidae 171 References 172 Additional References 172 11 PHYLA MICROSPORA AND MYXOZOA: PARASITES WITH POLAR FILAMENTS Phylum Microspora 173 Class Microsporea 173 Phylum Myxozoa 175 Family Myxosomatidae 175 References 180 Additional References 181 12 THE MESOZOA: PIONEERS OR DEGENERATES? Class Rhombozoa 183 Order Dicyemida 183 Order Heterocyemida 183 Class Orthonectida 184 Morphology and Biology 184 Phylogenetic Position 186 Physiology and Host-Parasite Relationships 186 Classification of Phylum Mesozoa 187 References 188 Additional References 188 13 INTRODUCTION TO THE PHYLUM PLATYHELMINTHES Platyhelminth Systematics 190 Classification of Phylum Platyhelminthes 192 Turbellarians 193 Acoels 193 Rhabditophorans 194 Temnocephalideans 195 Alloeocoels 196 Tricladids 196 Polycladids 196 References 196 Additional References 197 14 TREMATODA: ASPIDOBOTHREA Form and Function 199 Body Form 199 Tegument 199 Digestive System 199 Osmoregulatory System 199 Nervous System 200 Reproductive Systems 200 Development 202 Aspidogaster conchicola 204 Stichocotyle nephropsis 204 Phylogenetic Considerations 204 Classification of the Aspidobothrea 205 References 205 ADDITIONAL REFERENCES 15 TREMATODA: FORM, FUNCTION, AND CLASSIFICATION OF DIGENEANS Form and Function 207 Body Form 207 Tegument 208 Muscular System 211 Nervous System 211 Excretion and Osmoregulation 213 Acquisition of Nutrients and Digestion 214 Reproductive Systems 215 Development 217 Embryogenesis 217 Larval and Juvenile Development 217 Development in the Definitive Host 223 Trematode Transitions 223 Summary of Life Cycle 224 Metabolism 224 Energy Metabolism 224 Synthetic Metabolism 227 Biochemistry of the Tegument 227 Phylogeny of Digenetic Trematodes 228 Classification of Subclass Digenea 228 References 230 Additional References 233 16 DIGENEANS: STRIGEIFORMES Superfamily Strigeoidea 235 Family Diplostomidae 235 Family Strigeidae 236 Superfamily Schistosomatoidea 237 Family Schistosomatidae: Schistosoma Species and Schistosomiasis 238 References 250 Additional References 252 17 DIGENEANS: ECHINOSTOMATIFORMES AND PARAMPHISTOMIFORMES Superfamily Echinostomatoidea 253 Family Echinostomatidae 253 Echinostomatids as Models in Experimental Parasitology 255 Family Fasciolidae 256 Other Fasciolid Trematodes 259 Superfamily Paramphistomoidea 261 Family Paramphistomidae 261 Family Diplodiscidae 261 Family Gastrodiscidae 262 REFERENCESAA262 Additional References 263 18 DIGENEANS: PLAGIORCHIFORMES AND OPISTHORCHIFORMES Order Plagiorchiformes 265 Suborder Plagiorchiata 265 Family Dicrocoeliidae 265 Suborder Troglotrematata 268 Family Troglotrematidae 268 Order Opisthorchiformes 274 Family Opisthorchiidae 275 Family Heterophyidae 279 References 280 Additional References 281 19 MONOGENEA Form and Function 283 Body Form 283 Tegument 285 Muscular and Nervous Systems 286 Osmoregulatory System 287 Acquisition of Nutrients 288 Male Reproductive System 289 Female Reproductive System 290 Development 291 Oncomiracidium 291 Subclass Polyonchoinea 292 Subclass Polystomatoinea 293 Subclass Oligonchoinea 294 Phylogeny 295 Classification of Class Monogenoidea 295 References 296 Additional References 297 20 CESTOIDEA: FORM, FUNCTION, AND CLASSIFICATION OF THE TAPEWORMS Form and Function 299 Strobila 299 Scolex 300 Tegument 304 Calcareous Corpuscles 306 Muscular System 306 Nervous System 307 Excretion and Osmoregulation 307 Reproductive Systems 308 Development 312 Larval and Juvenile Development 313 Development in the Definitive Host 315 Metabolism 317 Acquisition of Nutrients 317 Energy Metabolism 318 Synthetic Metabolism 320 Hormonal Effects of Metabolites 320 Classification of Class Cestoidea 321 References 323 Additional References 326 21 TAPEWORMS Order Pseudophyllidea 327 Family Diphyllobothriidae 327 Other Pseudophyllideans Found in Humans 331 Sparganosis 331 Order Caryophyllidea 332 Order Spathebothriidea 332 Order Cyclophyllidea 333 Family Taeniidae 333 Family Hymenolepididae 342 Family Davaineidae 343 Family Dilepididae 345 Family Anoplocephalidae 345 Family Mesocestoididae 346 Family Dioecocestidae 347 Order Proteocephalata 347 Order Tetraphyllidea 348 Order Trypanorhyncha 349 Subcohort Amphilinidea 350 Cohort Gyrocotylidea 351 References 351 Additional References 354 22 PHYLUM NEMATODA: FORM, FUNCTION, AND CLASSIFICATION Historical Aspects 355 Form and Function 355 Body Wall 356 Musculature 359 Pseudocoel and Hydrostatic Skeleton 359 Nervous System 361 Digestive System and Acquisition of Nutrients 365 Secretory-Excretory System 368 Reproduction 369 Development 373 Eggshell Formation 373 Embryogenesis 374 Embryonic Metabolism 374 Hatching 375 Growth and Ecdysis 375 Metabolism 377 Energy Metabolism 377 Synthetic Metabolism 378 Classification of Phylum Nematoda 379 References 381 Additional References 384 23 Nematodes: Trichurida and Dioctophymatida, Enoplean Parasites 385 Order Trichurida 385 Family Trichuridae 385 Family Capillariidae 387 Family Anatrichosomatidae 388 Family Trichinellidae 388 Order Dioctophymatida 394 Family Dioctophymatidae 394 References 396 Additional References 398 24 NEMATODES: RHABDITIDA, PIONEERING PARASITES Families Steinernematidae and Heterorhabditidae 399 Family Rhabdiasidae 399 Family Strongyloididae 400 References 403 Additional References 403 25 NEMATODES: STRONGYLIDA, BURSATE RHABDITIANS Superfamily Ancylostomatoidea 405 Family Ancylostomidae 405 Superfamily Strongyloidea 412 Family Strongylidae 412 Family Syngamidae 412 Superfamily Trichostrongyloidea 412 Family Trichostrongylidae 412 Family Dictyocaulidae 414 Other Trichostrongyloids 414 Superfamily Metastrongyoidea 414 Family Angiostrongylidae 415 Other Metastrongyloids 415 References 416 Additional References 417 26 NEMATODES: ASCARIDIDA, INTESTINAL LARGE ROUNDWORMS Superfamily Ascaridoidea 419 Family Ascarididae 419 Family Anisakidae 427 Superfamily Heterakoidea 428 Family Ascaridiidae 428 Family Heterakidae 428 References 429 Additional References 431 27 NEMATODES: OXYURIDA, THE PINWORMS Family Oxyuridae 433 References 436 Additional References 437 28 NEMATODES: SPIRURIDA, A POTPOURRI Family Acuariidae 439 Family Gnathostomatidae 440 Gnathostoma spinigerum 440 Family Physalopteridae 441 Physaloptera Species 441 Family Tetrameridae 443 Family Gongylonematidae 443 Family Spirocercidae 444 Spirocerca lupi 444 Family Thelaziidae 445 References 446 Additional References 446 29 NEMATODES: FILAROIDEA, THE FILARIAL WORMS Family Onchocercidae 447 Wuchereria bancrofti 447 Brugia malayi 451 Onchocerca volvulus 452 Loa loa 456 Other Filariids Found in Humans 457 Dirofilaria immitis 457 References 459 Additional References 460 30 NEMATODES: CAMALLANINA, THE GUINEA WORMS AND OTHERS Family Camallanidae 461 Family Philometridae 461 Family Dracunculidae 462 Dracunculus medinensis 462 References 467 Additional References 467 31 PHYLUM ACANTHOCEPHALA: THORNY-HEADED WORMS Form and Function 469 General Body Structure 469 Body Wall 469 Reproductive System 472 Excretory System 474 Nervous System 475 Acquisition and Use of Nutrients 475 Uptake 475 Metabolism 476 Development and Life Cycles 476 Class Eoacanthocephala 476 Class Palaeacanthocephala 477 Class Archiacanthocephala 478 Effects of the Parasite on Its Host 478 Acanthocephala in Humans 479 Phylogenetic Relationships 480 Classification of Phylum Acanthocephala 480 References 481 Additional References 482 32 PHYLUM PENTASTOMIDA: TONGUE WORMS Morphology 485 Reproductive Anatomy 486 Biology 486 Development 487 Life Cycles 487 Pathogenesis 489 Visceral Pentastomiasis 489 Nasopharyngeal Pentastomiasis 490 Classification of Phylum Pentastomida 490 References 490 33 PHYLUM ARTHROPODA: FORM, FUNCTION, AND CLASSIFICATION General Form and Function 493 Arthropod Metamerism 493 Exoskeleton 494 Molting 497 Early Development and Embryology 499 Postembryonic Development 499 Diapause 500 External Morphology 501 Form of the Crustacea 501 The Form of Pterygote (Winged) Insects 502 The Form of the Acari 504 Internal Structure 506 Classification of Arthropodan Taxa with Symbiotic Members 509 References 513 Additional References 514 34 Parasitic Crustaceans 515 Class Maxillopoda 515 Subclass Ostracoda 515 Subclass Copepoda 515 Subclass Branchiura 526 Subclass Cirripedia 528 Subclass Tantulocarida 531 Class Malacostraca 531 Order Amphipoda 531 Order Isopoda 532 Order Decapoda 534 References 535 Additional References 537 35 PARASITIC INSECTS: MALLOPHAGA AND ANOPLURA, THE LICE Order Mallophaga 539 Morphology 539 Biology of Some Representative Species 540 Order Anoplura 542 Morphology 542 Mode of Feeding 542 Other Anoplurans of Note 545 Lice as Vectors of Human Disease 546 Epidemic, or Louse-Borne, Typhus 546 Trench Fever 547 Relapsing Fever 547 Control of Lice 547 References 548 Additional References 549 36 PARASITIC INSECTS: HEMIPTERA, THE BUGS Mouthparts and Feeding 551 Family Cimicidae 551 Morphology 552 Biology 553 Epidemiology and Control 553 Family Reduviidae 553 Morphology 554 Biology 554 Epidemiology and Control 555 References 556 37 PARASITIC INSECTS: SIPHONAPTERA, THE FLEAS Morphology 559 Jumping Mechanism 559 Mouthparts and Mode of Feeding 560 Development 560 Host Specificity 562 Families Ceratophyllidae and Leptopsyllidae 562 Family Pulicidae 562 Family Tungidae 564 Fleas as Vectors 565 Plague 565 Murine Typhus 567 Myxomatosis 567 Other Parasites 567 Control of Fleas 568 References 568 38 PARASITIC INSECTS: DIPTERA, THE FLIES Suborder Nematocera 571 Family Psychodidae 571 Family Culicidae 572 Family Simuliidae 580 Family Ceratopogonidae 581 Suborder Brachycera 582 Infraorder Tabanomorpha 582 Infraorder Muscomorpha 584 Myiasis 587 References 592 39 PARASITIC INSECTS: STREPSIPTERA, HYMENOPTERA, AND OTHERS Orders with Few Parasitic Species 595 Order Dermaptera (Earwigs) 595 Order Neuroptera (Lacewings) 595 Order Lepidoptera (Butterflies and Moths) 595 Order Coleoptera (Beetles) 596 Order Strepsiptera (the Stylops) 596 Morphology 597 Development 597 Order Hymenoptera (Ants, Bees, and Wasps) 599 Morphology 599 Development 600 Classification and Examples 601 Biological Control 603 References 604 40 PARASITIC ARACHNIDS: SUBCLASS ACARI, THE TICKS AND MITES Classification of the Arachnida and Acari 607 Order Ixodida: The Ticks 608 Biology 608 Family Ixodidae 609 Family Argasidae 614 Immunity of Ticks 616 Order Mesostigmata 616 Family Laelaptidae 616 Family Halarachnidae 616 Family Dermanyssidae 617 Family Rhinonyssidae 619 Order Prostigmata 619 Family Cheyletidae 619 Family Pyemotidae 620 Family Psorergatidae 620 Family Demodicidae 620 Family Trombiculidae 621 Order Oribatida 622 Order Astigmata 622 Family Psoroptidae 623 Family Sarcoptidae 623 Family Knemidokoptidae 624 Family Pyroglyphidae 625 Bee Mites 625 References 625 Additional References 627 Glossary 629 Index 647
Library of Congress Subject Headings for this publication: Parasitology