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The Biology of Gila Monsters and Beaded Lizards by Daniel D. Beck with contributions from Brent E. Martin and Charles H. Lowe and color photographs by Thomas A. Wiewandt Contents: 1. Monsters in Our Midst: History, Humans, and Heloderma (9,300 words) Introduction Whatís in a Name Heloderma Bites and the Western Frontier Heloderma in Indigenous Folklore Heloderma and Hollywood Heloderma in Medicine Folklore and Science How We Know What We Know Some Early Field Studies Techniques and New Directions 2. Evolution, Distribution and Systematics (10,300 words) Contemporary Helodermatid Lizards Heloderma horridum Wiegmann, 1829 Heloderma horridum exasperatum Heloderma horridum horridum Heloderma horridum alvarezi Heloderma horridum charlesbogertii Heloderma suspectum Cope 1869 Heloderma suspectum suspectum Heloderma suspectum cinctum What limits the distribution of Heloderma? The Fossil Record The Cretaceous Fossils, 100 ñ 65 million years ago Primaderma nessovi Nydam, 2000 Palaeosaniwa canadensis Gilmore, 1928 Estesia mongoliensis, Norell et al., 1992 Gobiderma pulchrum Borsuk-Bialynicka, 1984 Paraderma bogerti Estes, 1964 The Tertiary Fossils, 40 ñ 5 million years ago Eurheloderma gallicum Hoffstetter, 1957 Conferee Eurheloderma Bartels, 1983 Lowesaurus mathewi Gilmore Heloderma texana Stevens 1977 Recent Fossils Phylogeny Relationships with Varanus Relationships with Snakes Interpretations of the Fossil Record and Evolutionary Scenarios Evolution of a Sedentary Lifestyle Evolution of venom 3. The Venom System and Envenomation (9,400 words) Historical Overview Venom Delivery System Effects of the Venom Chemical Makeup of the Venom Hyaluronidase Serotonin Phospholipase A2 Nerve Growth Factor Helothermine The Kallikrein-Like Toxins Gilatoxin Horridum toxin Unnamed lethal toxin Helodermatine The Bioactive Peptides The Helospectins Helodermin Exendin-3 and Exendin-4 Gilatide Ecological/Evolutionary Role of Helodermatid Venoms Human Envenomation by Heloderma: Overview of Bite Case Histories Treatment of Bites 4. Physiological Ecology (8,400 words) Hot Dogs and Gila Monsters Thermal Biology Thermal tolerance Activity temperatures Basking behavior Body temperatures when at rest Energetics Metabolic Rates Temperature effects on metabolic rates Maximal metabolic rates Cost of locomotion Assimilation efficiency Energy Budgets Fat storage, drinking, and water loss rates Water Loss 5. Habitat Use and Activity Patterns (11,400 words) Intro: What defines habitat? Habitat types Mexican Beaded Lizards Tropical Dry Forest Other habitat types for H. horridum Gila Monster Habitats Sinaloan Thornscrub Southwestern Desertscrub Sonoran Desert habitats Arizona Upland Lower Colorado River Valley Central Gulf Coast and Plains of Sonora Semidesert Grassland Mojave Desert Habitats Chihuahuan Desertscrub Habitats Woodland Microhabitat Use Home range and dispersion of shelters The role of shelters Shelter fidelity Activity patterns Activity Bouts and Sprint-Speed Gila Monster Activity: Seasonal Patterns A Gila Monster in My Sleeping Bag Activity Bouts Timing of Activity Amount of Activity Conclusions 6. Population Ecology (with B. E. Martin and C.H. Lowe) (5,000 words) Intro Body size Size classes and population structure - Arizona, New Mexico, Utah, Jalisco Sex ratios Growth rates - laboratory - field Longevity Adult survivorship and population dynamics Predation Population density Are Helodermatids Rare? Summary of life history traits 7. Diet, Feeding, and Foraging Ecology (6,500 words) Red Tongue: Black Tongue (poem) Food habits Heloderma horridum Heloderma suspectum Gut and Fecal Analyses Field Observations Juvenile diet Behavioral Interactions Between Gila monsters and Their Prey Variation in Diet Feeding Behavior How prey are eaten Feeding with a spoon and a fork Foraging Ecology How prey are located Field observations of foraging behaviors Foraging ìstrategiesî 8. Reproduction, Behavior and Heloderma in Captivity (9,400 words) Sexing Heloderma and Sexual Dimorphism Reproductive cycles Male Cycle Female Cycle Frequency of Reproduction Reproductive behaviors Finding Mates Combat Heloderma suspectum Heloderma horridum Courtship and Mating Eggs and Oviposition The Puzzle of the Incubation Schedule Hatching and Hatchlings Heloderma in Captivity 9. Conservation (12,200 words) Introduction Threats to Helodermatid Lizards Habitat Loss and Alteration Fragmentation and Degradation Invasive Plants and Fire Habitat Threats and Human Population Growth Heloderma horridum and Tropical Dry Forest Direct Human-Caused Mortality Road Kills Pets Commercial Trade Poaching and Illegal Trade Current Conservation and Legal Status National and International Status State Legal Status Arizona California New Mexico Nevada Utah Vulnerability Conservation Options and Strategies The tortoise and the Gila monster: a case study Conservation Strategies Habitat protection and restoration Managing Reserves Buffer Zones Corridors Restoration Displaced Heloderma and Relocation Captive Propagation and Repatriation Living With Heloderma in Our Midst 10. Future Directions (2,500 words) Taxonomy, Evolution, and Distribution The Venom System Physiological Ecology Habitat Use and Activity Population Biology Food Habits Reproduction Conservation Literature Cited (12,500 words) Total word count = 97,000 words
Library of Congress Subject Headings for this publication:
Helodermatidae.