Table of Contents

CHAPTER 1 Introduction to Venomous Systematics. P.M. BRIGNOLI A. Introduction 1 I. What are Arthropoda? 1 II. The Main Divisions of the "Type" 1 III. The 2 1. Scorpionida 2 2. Uropygi or Thelyphonida 2 3. Pseudoscorpionida 3 4. Opiliones 3 5. Acarina 3 6. Araneae 3 IV. The Crustacea 4 V. The "Myriapoda" 4 1. Chilopoda 4 2. Diplopoda 4 VI. The Hexapoda or Insecta 5 1. Blattodea and Dermaptera 5 2. Rhynchota and Anoplura 6 3. Aphaniptera 6 4. Coleoptera 6 5. Hymenoptera 6 6. Diptera 6 7. Lepidoptera 6 VII. Some General Advices 7 1. How to Identify an Arthropod 7 2. How to Conserve an Arthropod 7 3. What to Expect from the Bibliography 7 References 8

CHAPTER 2 of Crustacea and Merostomata. Y. HASHIMOTO and S. KONOSU. With 10 Figures A. Introduction 13 B. Suspected of Being Poisonous 14

http://d-nb.info/780105869 X Table of Contents

C. of 15 I. Crabs Containing 15 II. Toxicity of pictor 19 D. Biology of Poisonous Xanthid Crabs 19 I. aeneus 19 1. Description 20 2. Color in Life 20 3. Habitat and Distribution 20 4. Feeding Habits and Spawning Season 20 II. Platypodia granulosa 20 1. Description 21 2. Color in Life 22 • 3. Habitat and Distribution 22 III. floridus 22 1. Description 22 2. Color in Life 22 3. Habitat and Distribution 24 IV. 24 1. Description 24 2. Coloration 24 3. Distribution 24 E. Chemistry of in Crabs 24 I. Purification of Z. aeneus 25 II. Chemical Properties of Z. aeneus Toxin 26 III. Chemical Properties of Toxins from Other Crabs 26 F. Pharmacology of Toxins in Crabs 27 I. Z. aeneus Toxin 27 1. Cases and Symptoms 27 2. Signs in Test 27 3. Effects on Nerve Excitation 28 II. The Toxin of L. pictor 32 III. Toxin in Other Crabs 33 G. Coconut Poisoning 33 I. Biology of Coconut Crab 34 1. Description 34 2. Color when Alive, Habitat, and Distribution 34 II. Toxicity of the Coconut Crab 34 H. Horseshoe-Crab Poisoning 35 I. Biology of Horseshoe Crabs 35 II. Toxicity of Horseshoe Crabs 36 III. Clinical Characteristics 37 I. Prevention and Treatment of Crab Poisoning 37 References 37 Table of Contents XI

CHAPTER 3 Defensive Secretions of Millipeds. T. EISNER, D. ALSOP, K. HICKS, and J. MEINWALD. With 13 Figures Introduction 41 A. Distribution, Structure, and Mode of Operation of the Glands 42 B. Chemistry of the Secretions 51 C. Biochemistry of the Secretions 55 D. Defensive Effectiveness of the Glands 58 E. Implications to Humans 65 F. Other Defenses of Millipeds 66 References 67

CHAPTER 4 Secretions of Centipedes. A. MINELLI. With 2 Figures A. The Centipedes 73 B. Glands and Secretions 75 I. Defensive and Offensive Weapons of Centipedes 75 II. Topography of the Exocrine Glands 75 III. Histology of the Glands 76 1. The Forcipular Gland 76 2. Ventral and Coxal Glands of the Geophilomorphs 77 IV. Secretions 78 1. Sticky Secretions 78 2. Luminescent Secretions 79 3. Smelling Secretions 79 4. of the Forcipular Glands 80 a) Effects of Bite on Humans 80 b) Effects of Bite on Laboratory Animals 81 c) Chemistry of the Venom 82 References 82

CHAPTER 5 Secretions of Opilionids, Whip and Pseudoscorpions. T. EISNER, D. ALSOP, and J. MEINWALD. With 7 Figures A. Order Opiliones 87 B. Order Uropygi 94 C. Order Pseudoscorpiones 97 References 97 XII Table of Contents

CHAPTER 6 Review of the Families, with Notes on the Lesser-Known Poisonous Forms. S. BETTINI and P.M. BRIGNOLI A. Orthognatha = Theraphosomorphae = Mygalomorphae 103 B. Labidognatha = 108 I. Haplogynae 108 II. Entelegynae = Trionychae 110 1. Araneoidea 110 a) Araneidae = Argiopidae 110 b) Linyphiidae Ill c) Ill 2. Lycosoidea Ill a) Agelenidae 112 b) Argyronetidae 112 c) Pisauridae 112 d) Lycosidae 112 e) Oxyopidae 114 III. Entelegynae = Dionychae 114 1. Gnaphosidae ( = Drassodidae = Drassidae) 114 2. Clubionidae 114 3. Ctenidae 116 4. Eusparassidae ( = Heteropodidae) 116 5. Thomisidae 116 6. Salticidae 117 IV. Cribellatae 117 References 118

CHAPTER 7 Venoms of Dipluridae. M.R. GRAY and S.K. SUTHERLAND

Introduction 121

A. Systematics and Distribution. M.R. GRAY. With 1 Figure 121

B. Biology and Venoms. S.K. SUTHERLAND. With 10 Figures 126 I. Introduction 126 II. Notes on Atrax robustus 127 III. Source of Venom for Experimentation 129 1. Maintenance of in the Laboratory 129 2. Collection of Venom and Venom Yields 130 IV. Structure of the Venom Glands in Atrax robustus 130 V. Toxicity of Venom in Various 132 1. Atrax robustus Venom 132 2. Venom of Other Species of A trax 133 Table of Contents XIII

VI. Chemistry of Sydney Funnel-Web Spider (Atrax robustus) Venom . 134 VII. Pharmacopathologic Studies with Atrax robustus Venom 137 1. Intact Animal Experiments 137 2. Studies Upon Isolated Preparations 139 3. Investigation of Antidotes 141 VIII. Immunologic Studies on Atrax Venom 142 IX. Human by Atrax robustus 143 1. General 143 2. Signs and Symptoms of Envenomation 144 3. Treatment of Victims 145 4. Prognosis 146 X. Discussion 146 References 147

CHAPTER 8 Venoms of Theridiidae, Genus

A. Systematics, Distribution and Biology of Species; Chemistry, Pharmacology and Mode of Action of Venom. S. BETTINI and M. MAROLI. With 8 Figures I. Systematics and Distribution of Species 149 II. Biology 151 1. Habitat 151 2. Habits 152 3. Life Cycle 152 4. Act 154 III. Venomous Apparatus 154 IV. Venom 156 1. Methods for Obtaining the Venom 156 2. Chemistry 157 3. Separation of Venom Components 158 V. Toxicity of Venom in Different Animal Species 159 1. Whole Venom 159 2. Venom Components 160 VI. Pharmacology and Mode of Action on Different Zoological Groups 162 1. Invertebrates 166 a) Crustacea 166 b) Insecta 167 c) Mollusca 170 2. 170 a) Pisces 170 b) Amphibia 170 c) Aves 172 d) Mammalia 172 3. Toxicity in Cells Cultivated in vitro 177 4. Effect of Toxin on Lipid Bilayer Membranes 177 XIV Table of Contents

5. Distribution of Venom in Envenomed Animals 177 6. Protective Activity of Venom Against Botulin Toxin 178 VII. Toxicity of Eggs and Tissue Extracts 178 References 179

B. Epidemiology of Envenomation, Symptomatology, Pathology and Treatment. Z. MARETIC. With 3 Figures I. Epidemiology 185 1. Geographical Distribution 185 a) Europe 185 b) Asia 186 c) Africa 186 d) America 186 e) 187 2. Factors Influencing Number of Spiders and Frequency of Contacts with Man. Epidemics of 187 3. Other Epidemiological Data 191 II. Human Symptomatology 192 III. Differential Diagnosis 199 IV. Clinical Course, Duration of Hospitalization, Convalescence, and Complications 200 1. Course 200 2. Convalescence 200 3. Complications 201 V. Prognosis 201 VI. Post-Mortem Findings 201 VII. Aspecific Therapy 202 VIII. Specific Therapy 204 1. Preparation of Sera 204 2. Cross Activity of Sera Obtained with Antigens from Different Species of Spiders 206 3. Time and Dose of Serum Administration 206 IX. Prevention of Latrodectism 207 References 207

CHAPTER 9 Venoms of Theridiidae, Genus . Z. MARETIC. With 2 Figures A. Introduction 213 B. Venomous Spiders of the Genus Steatoda 213 I. Steatoda paykulliana 213 1. Distribution, Biology, Morphology, and Venomous Apparatus . 213 2. Effects of the Toxin 214 II. Steatoda grossa 216 References 216 Table of Contents XV

CHAPTER 10 Venoms of Ctenidae. S. SCHENBERG and F.A. PEREIRA LIMA. With 15 Figures A. Introduction 217 B. Morphology of the Venomous Apparatus 219 I. Chelicerae 219 1. Fangs or Claws 219 2. Basal Segment 220 3. Musculature 220 II. Venom Glands 221 1. Musculature 221 2. Histology of the Glandular Tissue 221 C. Methods of Collecting Venoms 222 1. Method of Collecting Venom by its Extraction from the Glands . 223 2. Method of Collecting Venom by Electrical Stimulation 223 D. Physiology of Biting 224 E. Crude Venom Pharmacology 224 I. Effect on Dogs 225 1. Excruciating Local Pain 225 2. Sneezing 226 3. Lacrimation and Mydriasis 226 4. Hypersalivation . 226 5. Erection 226 6. Ejaculation 228 7. Toxicity 229 8. Hypotensive Response 228 9. Tachyphylaxis 229 II. Guinea Pig Ileum Contraction 229 III. Effects on Mice 229 1. Local Pain 230 2. Hypersalivation 230 3. Erection 231 4. Ejaculation 231 5. Toxicity 231 6. Distensive Paralysis 232 IV. Effects on Other Animals 232 F. Biochemistry of the Phoneutria nigriventer Venom, and Procedures for its Fractionation 233 I. Immunologic Aspects of the Venom 233 II. Enzymic Inactivation 233 III. Physicochemical Properties of the Active Polypeptides 234 IV. Dialysis 235 V. Venom Fractionation 236 1. Ammonium Sulfate, Electrophoresis, Barium Sulfate Adsorption 236 2. Flaccid Paralysis 236 3. Hemorrhagic Effect 237 XVI Table of Contents

VI. Gel Chromatography 237 VII. Ion Exchange Columns 240 G. Envenomation in Man 242 I. Symptomatology 242 II. Frequency of Cases 243 III. Therapy of Envenomation 243 References 244

CHAPTER 11 Venoms of Scytodidae. Genus Loxosceles. H. SCHENONE and G. SUAREZ. With 6 Figures A. Distribution and Biology of Venomous Species. Chemistry, Toxicity, Phar- macology and Mode of Action of Venom 247 I. Systematics and Distribution of Species 247 II. Morphology and Biology 248 III. Anatomy and Histology of the Venomous Apparatus 249 IV. Physiology, in Particular Physiology of the Bite 250 V. Chemistry of the Venom 251 1. Methods of Obtaining the Venom 251 2. Chemical Composition of Venom Extracts 252 3. Fractionation and Characterization of Venom Proteins ..... 252 4. Enzymatic Properties of Venom 253 5. Immunogenicity of the Venom 254 VI. Toxicity of Venom and other Components of Loxosceles on Various Animal Species 254 1. In vivo Studies 254 2. In vitro Studies 254 VII. Pharmacology and Mechanism of Action 260 B. Epidemiology, Symptomatology, Pathology, Prognosis, Treatment, and Prevention of 260 I. Epidemiology of Envenomations. Geographic and Temporal Distri- bution of Cases. Other Epidemiologic Data 260 II. Human Symptomatology 262 1. Cutaneous Form of 262 2. Viscerocutaneous Form of Loxoscelism 264 III. Differential Diagnosis 266 1. Cutaneous Forms 266 2. Viscerocutaneous Form 266 IV. Clinical Course and Duration of Hospitalization 266 1. Local and Ulcer 266 2. Viscerocutaneous Involvement 266 V. Prognosis and Frequency of Lethal Cases 266 VI. Pathology 267 VII. Laboratory Findings 268 Table of Contents XVII

VIII. Treatment 269 1. Nonspecific 269 2. Specific 270 IX. Prevention 271 References 271

CHAPTER 12 The Genus Centruroides (Buthidae) and Its Venom. H.L. STAHNKE. With 2 Figures A. Species and Their Distribution 279 B. Some Aspects of Behavior 280 C. The Prevention and Control of Scorpions 282 D. Morphology and Function of the Venom Apparatus 284 E. Extracting Venom 286 F. Mammalian Physiologic Reaction to the Venom 287 I. Clinical Observations 287 1. Symptoms of Severe Envenomization (Children under 16 Years of Age) 287 2. Symptoms of Envenomization in Adults 287 II. Preclinical Observations 287 1. Pharmacologic Considerations 287 2. Pathologic Effects 288 3. Allergic Potentialities 289 G. Human Behavior and Scorpions 289 H. Venom Chemistry .289 J. Suggestions for Therapy 294 References 294

CHAPTER 13 Venoms of Buthinae A. Systematics and Biology of Buthinae. A. SHULOV and G. LEVY 309 References 312

B. Symptomatology and Treatment of Buthinae Stings. P. EFRATI 312

I. Pathophysiology of the Envenomation 314 II. Treatment 316 References 316 XVIII Table of Contents

C. Chemistry and Pharmacology of Buthinae Venoms. E. ZLOTKIN, F. MIRANDA and H. ROCHAT. With 11 Figures I. Introduction 317 II. Chemistry 318 1. Isolation and Purification 319 2. Composition and Structure 323 3. Chemical Modifications 328 III. Pharmacology 328 1. Lethal Potency and Symptomatology . 328 2. Cardiovascular Effects 330 a) Clinical Investigations 330 b) Histopathologic Changes 330 c) Vascular Effects 332 d) Cardiac Dynamics 334 3. Respiratory Effects 336 4. Muscular Effects 337 a) Direct Effects on Skeletal Muscles 338 b) Effects Through Neuromuscular Junctions 338 c) Effects on Smooth Muscles 340 5. Action on Axonal Membranes 341 6. Action on 345 IV. Action Mechanisms 351 1. Autopharmacologic Intoxication 351 2. Is There a Direct Excitatory Effect on Muscles? 353 3. Interactions with Sodium, Calcium, and TTX 354 4. The Pharmacologic Diversity of Scorpion Toxins 356 References 362

CHAPTER 14 Venoms of Tityinae

A. Systematics, Distribution, Biology, Venomous Apparatus, etc. of Tityinae; Venom Collection, Toxicity, Human Accidents and Treatment of Stings. W. BUCHERL. With 3 Figures 1. The Venomous Species and Their Geographical Distribution . .371 2. Description of the Most Venomous Species 374 3. Frequency of the Most Venomous Species 375 4. Food and Life Habits 375 5. Venom Glands 376 6. Quantities and Storage of the Venom 377 7. Toxicity and Human Accidents 377 8. Treatment of Scorpion Envenomation 378 References 378 Table of Contents XIX

B. Chemical and Pharmacologic Aspects of Tityinae Venoms. C.R. DINIZ. With 8 Figures Introduction 379 I. Chemical Properties of Tityinae Venoms 380 1. Purification of Toxic Components from Tityinae Venoms. . . .380 II. Pharmacology of Tityus Venom Intoxication 383 1. Release of by Tityus Venom 383 a) 383 b) Epinephrine and 386 III. General Effects 391 IV. Conclusion 391 References 391

CHAPTER 15 Chactoid Venoms M. GOYFFON and J. KOVOOR. With 6 Figures I. Introduction 395 II. Morphology of the Telson in Chactoids 396 III. Anatomy and Development of the Venom Glands 397 IV. Comparative Histology and Histochemistry of the Venom Glands in Adults 398 V. Chemical Composition of Chactoid Venoms 402 VI. Pharmacology of Chactoid Venoms 406 VII. Experimental Toxicity 408 VIII. Toxicity for Humans 410 IX. Treatment and Prophylaxis of Scorpion Poisoning 412 References 413

CHAPTER 16 . M.F. MURNAGHAN and F.J. O'ROURKE. With 3 Figures A. Introduction 419 B. Ticks 420 I. The Biology of Ixodid Ticks 420 II. Classification of Ticks 422 III. Feeding in Ixodid Ticks 424 IV. Virulence 425 C. Hosts 426 I. Humans 426 1. Distribution 426 2. Seasonal Incidence 429 3. Age 429 4. Sex 429 XX Table of Contents

II. Livestock 430 1. Natural Occurrence 430 2. Experimentally Induced 430 III. Wildlife 431 IV. Pets 432 1. Natural Occurrence 432 2. Experimentally Induced 433 V. Laboratory Animals 435 D. Symptomology 436 I. Humans 436 II. Animals 438 E. Mechanism of the Paralysis 439 I. Physiology and Pharmacology of Paralysis due to Dermacentor ander- soni 439 1. Neuromuscular Transmission 439 2. Nerve Conduction 444 3. Central Nervous Transmission 447 4. Miscellaneous Agents 448 II. Experimental Physiological Findings with Other Ticks Species . . . 449 1. Ixodes holocyclus 449 2. Argas (Persicargas) persicus 449 III. Identification of the Presence and Removal of the Toxin 449 IV. Immunity 453 V. Autopsy Findings 454 F. Summary 455 References 458

CHAPTER 17 Toxins of Blattaria. L.M. ROTH and D.W. ALSOP. With 24 Figures A. Systematics 465 B. Biology 466 C. Allergy Caused by Cockroaches 467 1. Contactant Allergens 467 2. Allergens 468 3. Injectant Allergens 469 4. Ingestant Allergens 469 D. Defensive Glands 470 1. Type I Glands 471 2. Type II Glands 472 3. Type III Glands 478 4. Type IV Glands 481 5. Type V Glands 482 References 484 Table of Contents XXI

CHAPTER 18 Venoms of Rhyncota (Hemiptera). J. WEATHERSTON and J.E. PERCY A. Introduction 489 I. Aquatic Heteroptera 490 II. Terrestrial Heteroptera 490 B. Morphology and Histology of the Scent Glands 491 I. Adult Insects 491 II. Larvae 494 C. Chemistry of the Secretions 495 I. Structure of Secretion Components 495 1. Types of Compounds 496 2. Toxins of Aquatic Heteroptera 497 3. Toxins Isolated from Terrestrial Heteroptera 497 4. Toxins Isolated from Homoptera 497 II. Extraction Methods 502 III. Methods of Purification and Structural Elucidation 502 1. Gas Liquid Chromatography 504 2. Thin-Layer Chromatography 505 3. Spectroscopic Methods 505 D. Function of Hemipteran Secretions 505 E. Other Glands and Secretions 506 References 506

CHAPTER 19 Venoms of Coleoptera. J. WEATHERSTON and J.E. PERCY A. Introduction 511 B. Morphology and Histology of the Scent Glands 512 I. Pygidial Gland 512 1. Reservoir 512 2. Secretory Cells 517 3. Fine Structure of Cells 518 4. Accessory Gland 519 II. Thoracic Glands 519 III. Other Glands 519 C. Chemistry of the Secretions 520 I. Structure of Secretion Components 520 1. Types of Compounds 520 2. Toxins of the Alleculidae 521 3. Toxins of the Cantharidae 521 4. Toxins of the Carabidae 521 5. Toxins of the Cerambycidae 526 6. Toxins of the Chrysomelidae 527 XXII Table of Contents

7. Toxins of the Coccinellidae 527 8. Toxins of the Dytiscidae 527 9. Toxins of the Gyrinidae 530 10. Toxins of the Meloidae 533 11. Toxins of the Silphidae 534 12. Toxins of the Staphylinidae 534 13. Toxins of the Tenebrionidae 535 II. Extraction Methods 542 III. Methods of Purification and Structural Elucidation 543 1. Gas Liquid Chromatography 543 2. Thin-Layer and Paper Chromatography 544 3. Spectroscopic Methods 545 IV. Biosynthesis 545 D. Biological and Physiological Effects of the Toxins 547 References 548

CHAPTER 20 Venoms of Lepidoptera. A. DELGADO QUIROZ. With 19 Figures A. Introduction 555 B. Erucism 556 I. Description 556 II. Pathology 558 III. Symptomatology 560 1. Pathogenesis 560 a) Foreign-Body Syndrome 560 b) Toxic Syndrome 562 c) Infectious Syndrome 563 d) Allergic Syndromes 563 2. Evolution 563 3. Immunity 564 IV. Diagnosis 564 1. Super-Family Papilionoidea (DYAR, 1902) 565 a) Morphidae (KOLLAR, 1850) 565 b) Nymphalidae (SWAINSON, 1872) 565 2. Super-Family Bombycoidea (DYAR, 1902) 565 a) Lasiocampidae (HARRIS, 1841) 567 b) Thaumetopoeidae ( = Notodontidae) 568 c) Saturniidae ( = Hemileucidae) (WALKER, 1855) 568 3. Super-Family Noctuoidea (MOSHER, 1916) 570 a) Arctiidae (STEPHENS, 1829) 570 b) Lymantriidae ( = Liparidae) (HAMPSON, 1892) 570 c) Noctuidae (STEPHENS, 1829) 571 4. Super-Family Zygaenoidea (GRAVENHORST, 1843) 571 a) Cochlidiidae ( = Eucleidae) (DYAR, 1898) 571 b) Megalopigydae (BERG, 1882) 571 Table of Contents XXIII

5. The Apparatus 573 a) Primitive-Type Poison Apparatus (BEYER, 1922) 575 b) Developed-Type Poison Apparatus (FOOT, 1922) 578 6. 580 a) Obtaining the Poison 580 b) Conserving the Poison 580 c) Pharmacology of the Extracts of Erucic Poisoning 583 V. Treatment 584 a) Immediate Treatment 584 b) Symptomatic and Preventive Treatment 584 c) Treatment of Foreign-Body Syndrome 585 VI. Epidemiology 585 1. Erucism on the American Continent 586 2. Erucism on the Continent of Europe 588 3. Erucism in Africa, Asia and Australia 590 4. Epidemiological Structure of Erucism 590 a) Factors Relating to the Aetiological Agent 590 b) Factors Relating to Erucic Intoxication 591 c) Factors Relating to the Environment 591 5. Epidemiological Characteristics of Erucism 592 6. Prevention of Erucism 593 a) Measures Directed Against the Primary Source 593 b) Measures Directed Against the Means of Transmitting the Poi- sonous Hairs 593 c) Immunization or Desensitization 593

C. Lepidopterism 594. I. Description 594 II. Pathology 594 III. Symtomatology 594 1. Pathogenesis 594 a) Foreign-Body Syndrome 595 b) Toxic Syndrome 595 c) Infectious Syndrome 597 d) Allergic Syndrome 597 2. Evolution 597 3. Immunity 598 IV. Diagnosis 598 1. Families with Poisonous Erucae 598 2. Zygaenidae Family 598 3. Saturnidae Family ( = Hemileucidae) (WALKER, 1855) 598 4. The Poison Apparatus 600 5. Toxicology 601 a) Extraction of the Poison 601 b) Conservation of the Poison 602 c) Pharmacology 602 XXIV Table of Contents

V. Treatment 602 a) Immediate 602 b) Symptomatic and Preventive 603 VI. Epidemiology and Prevention 603 1. South American Lepidopterism 603 a) In the Guianas and Venezuela 603 b) In Peru 603 c) In Brazil 604 d) In Argentina 604 2. Epidemiological Structure of Lepidopterism 604 a) Factors Relating to the Anthropotoxic Species of the Genus Hylesia 604 b) Factors Relating to the Source of Lepidopteran Intoxication . 605 c) Factors Relating to the Environment 606 3. Prevention of Lepidopterism 606 a) Measures Against the Primary Source of Intoxication .... 606 b) Measures Against the Means of Transmission of the Poison- Bearing Flechettes 607 References 607

CHAPTER 21 Venoms of Apidae. R. O'CONNOR and M.L. PECK. With 8 Figures Introduction 613 A. Venom Apparatus, Sting Mechanism, and Venom Collection 617 I. Venom Apparatus 617 1. Apini 617 2. Bombini 619 3. Meliponini 619 II. Venom Formation 619 1. Apini 619 2. Bombini 620 3. Meliponini 620 III. Venom Collection 620 1. Venom Sac and Gland Extraction 620 2. Electrical Excitation of Groups of Bees 621 3. Pure Venom from Individual Hymenoptera 621 B. Composition of Apis mellifera Venom 622 I. Venom Quantity and Purity 622 II. Water Content 623 III. Volatile Components 623 IV. General Composition 624 V. Enzymes 626 1. Hyaluronidase 626 2. Phospholipase A System 628 3. Other Enzyme Studies 629 Table of Contents XXV

VI. Large Peptides 630 1. General Characteristics 630 2. The Melittin Family 633 3. Apamin, a 634 4. The Mast Cell Degranulating (MCD) Peptide 635 5. Other Active Peptides 636 VII. Small Molecules 636 1. Some General Considerations 636 2. Small Peptides and Free Amino Acids 639 3. Biogenic Amines 640 4. Non-Nitrogenous Compounds 640 VIII. Compounds Shown to be Absent 640 IX. Summary of Unresolved Questions on Venom Composition .... 641 C. Physiologic Activity of Apis mellifera Venom 641 I. Direct and Indirect Activity 641 II. Toxicity Studies 645 III. Antigenic Character 646 1. Venom Hypersensitivity and Immunity 646 2. Antigenic Comparisons among Various Venoms 647 IV. The Arthritis Question 647 1. General Considerations 647 2. Possible Mechanisms 647 3. Potentially Interesting Components . 648 4. Speculations 648 V. Radioprotective Properties 648 VI. Other Physiologic Properties 649 D. Venoms of Other Apidae 651 I. Other Apini 651 II. Bombini 653 Summary 653 References 653

CHAPTER 22 Venoms of Sphecidae, Pompilidae, Mutillidae, and Bethylidae. W. RATHMAYER. With 8 Figures A. Introduction 661 B. Venoms of Sphecidae 661 I. General Aspects 661 1. Biological Notes 661 2. Sting Apparatus and Venom Glands 662 3. Stinging of the Prey 663 4. Venom Actions 665 5. Effectiveness of Paralysis 666 6. Specificity of Venoms 668 XXVI Table of Contents

II. Venoms of Specific Genera 672 1. Genus Philanthus 672 a) Aspects of Biology and Preying 672 b) Philanthus triangulum 673 a) Anatomy of the Sting Apparatus 673 /?) Stinging of the Bee and Action of the Venom 673 y) Mechanism of Action of the Venom 677 3) Specificity of the Venom 678 2. Genus Sceliphron 680 a) Composition of the Venom 680 C. Venoms of Pompilidae 680 I. General Aspects 680 II. Stinging and Effects of the Venom on Locomotion 681 D. Venoms of Mutillidae 682 E. Venoms of Bethylidae 682 Appendix 1: Classification of the Family Sphecidae 683 References 685

CHAPTER 23 Venoms of Vespidae. H. EDERY, J. ISHAY, S. GITTER and H. JOSHUA. With 30 Figures A. Introduction 691 B. Biology of Vespidae 691 I. and Geographical Distribution 692 1. Stenogastrinae 692 2. Polistinae 693 3. Vespinae 694 II. Social Behavior 701 1. Life Cycle 701 2. The Nest Entrance 701 3. Foraging Behavior 702 4. Enemies and Parasites 704 5. Defensive Behavior 705 6. Hornet Traps 705 7. Aggressive Behavior-Stinging 705 8. Aggressiveness of Various Species 706 C. The Venomous Apparatus 706 I. The Nonglandular Parts of the Sting Apparatus. Mechanism of Stinging 706 II. The Glandular Parts of the Sting Apparatus 708 III. Collection of the Venom 713 IV. Quantity of Venom Collected from a Single Wasp or Hornet . . .715 D. Chemistry of Vespidae Venoms 715 Table of Contents XXVII

I. Physico-Chemical Properties 715 II. Components of Vespidae Venoms 715 1. Low-Molecular-Weight Substances 716 a) Acetylcholine 716 b) Histamine 718 c) Serotonin (5-Hydroxytryptamine, 5-HT) 719 d) Dopamine, Noradrenaline, and Adrenaline 719 e) Kinins 720 2. High-Molecular-Weight Substances: Enzymes 724 a) Cholinesterase 724 b) Histidine Decarboxylase 724 c) Phospholipases 725 d) Acid, Alkaline, and Natural DNAses 725 e) Hyaluronidase 725 f) Protease 726 g) Poly- and Disaccharidases 726 3. Miscellaneous Substances 726 E. Toxicology 726 I. Systemic Toxicity of V.v 727 II. Toxicity of V.v. as Related to Ontogenesis 728 III. Pathological Findings 729 F. Pharmacology 729 I. Cardiovascular Apparatus 731 1. Blood Pressure 731 2. Hemolysis 732 3. Anticoagulant Activity 739 4. Hyperglycemia 742 5. Increase in Permeability of Microcirculation Vessels 742 6. Respiratory Apparatus 743 7. Striated Muscle 743 8. Ocular Irritation 744 9. Isolated Organ Preparations 744 10. Release of Histamine 745 11. Pharmacology of Vespidae Kinins 745 G. Antigenicity and Immunological Aspects 748 H. Clinical Aspects of Envenomation 750 I. Incidence of Sting and Dangers 750 II. Symptomatology 752 1. Local Symptoms 752 2. Generalized Symptoms 753 a) Toxic Reactions 753 b) Allergic Reactions 754 III. Pathology 754 IV. Treatment '. 755 XXVIII Table of Contents

1. First Aid 755 2. Emergency Treatment 755 3. Preventive Therapy-Desensitization 756 V. Prevention of Stinging 756 VI. Control of Vespidae 757 J. Concluding Remarks 758 References 759

CHAPTER 24 Venoms of Braconidae. R.L. BEARD. With 13 Figures Introduction 773 A. Parasitism by Braconidae 774 I. Stinging and Parasitism 774 II. Paralysis of Hosts 775 III. The Preferred Species for Study 776 B. The Venom Apparatus 777 I. Morphology and Histology 777 II. Mechanism of Action 777 III. Volume of Ejected Venom 778 C. Methods of Obtaining Venom for Study 779 D. Injection and Transport of Venom 780 E. Assays of Venom Activity 781 F. Braconid Venom 783 I. Chemistry 783 II. Stability 783 III. Potency 784 IV. Production, Storage, and Use of Venom 784 1. Synthesis 785 2. Storage and Volume of Venom Used 785 3. Rate of Venom Production 786 G. Mode of Action of Venom 786 I. Effect on the Heart 786 II. Effect on the Gut 787 III. Effect on Gross Respiration 788 IV. Effect on Neuromuscular Activity 789 V. Biochemical Studies 792 H. Host Preferences and Host Sensitivity to Venom 794 J. Immune Relationships 797 References 799 Table of Contents XXIX

CHAPTER 25 Venoms and Venom Apparatuses of the Formicidae: Myrmeciinae, Ponerinae, Dorylinae, Pseudomyrmecinae, Myrmicinae and Formicinae. M.S. BLUM and H.R. HERMANN. With 65 Figures A. Introduction 801 B. Venoms and Dufour's Gland Constituents 803 I. Myrmeciinae 803 1. Venom 803 a) Histamine 803 b) Smooth Muscle Stimulant 803 c) Hyaluronidase 804 d) Direct Hemolytic Factor 804 e) Phospholipase A 804 f) Histamine-Releasing Activity 804 2. Dufour's Gland Constituents 805 II. Ponerinae 805 1. Venom 805 2. Dufour's Gland Constituents 805 III. Dorylinae 805 1. Venom 805 IV. Pseudomyrmecinae 805 1. Venom 805 2. Dufour's Gland Constituents 805 V. Myrmicinae 805 1. Venom 805 2. Dufour's Gland Constituents 809 VI. Formicinae 810 1. Venom 810 2. Dufour's Gland Constituents 811 C. Morphology of the Hymenopterous Venom Apparatus 817 D. General Formicid Venom Apparatus 817 E. Venom Apparatus Sclerites and their Musculature = General Formicid Structure 819 I. Myrmeciinae and Ponerinae 827 II. Dorylinae 828 III. Pseudomyrmecinae 830 IV. Myrmicinae 830 V. Formicinae 832 F. Abdominal Muscles That Indirectly Affect the Movement of the Venom Apparatus 835 G. Muscles of the Venom Apparatus 835 H. Evolution of the Venom Sclerites 835 I. Glands Associated with the Venom Apparatus 837 I. Myrmeciinae, Ponerinae, Dorylinae and Pseudomyrmecinae .... 837 II. Myrmicinae , 839 III. Formicinae 839 XXX Table of Contents

J. Gland Phylogeny 840 K. Envenomization 843 L. Mandibular Gland Constituents 844 I. Ponerinae 844 II. Dorylinae 845 III. Pseudomyrmecinae 846 IV. Myrmicinae 846 V. Formicinae 849 M. Morphology of the Mandibular Glands 851 N. Metapleural Gland Products 853 I. Myrmicinae 853 O. Morphology of the Metapleural Glands 854 P. Natural Products of the Formicidae = An, Overview 854 Q. The Formicid Venom Apparatus =Phyletic Implications 857 I. Relationship to Wasps 857 II. Formicinae 859 III. Pseudomyrmecinae 859 IV. Myrmicinae 860 V. Ponerinae, Dorylinae, Cerapachyinae 860 VI. Old and New World Dorylinae 860 References 860

CHAPTER 26 Venom and Venom Apparatuses of the Formicidae: Dolichoderinae and Aneuretinae. M.S. BLUM and H.R. HERMANN, JR. With 11 Figures A. Introduction and Taxonomy 871 B. Dolichoderine Natural Products and Anatomy of the Venom Apparatus . 872 I. Mandibular Gland Constituents 872 II. Poison Gland Constituents 873 III. Anatomy of the Venom Apparatus Components in the Dolichoderi- nae 874 IV. Anatomy of Venom Sclerites 875 V. Dufour's Gland Constituents 877 VI. Anatomy of Dufour's Gland 880 C. Anal Gland Constituents 880 I. Anatomy of the Anal Glands in the Dolichoderinae 884 II. Biological Activities of Dolichoderine Anal Gland Products .... 885 D. Dolichoderine Natural Products = An Overview 886 E. Phyletic Implications Based on Venom Slerites and Associated Glands. . 890 References 891 Author Index 895 Subject Index 940