The Inadequate Environment

I: The Inadequate Environment.- 1. The Environment of All Organisms Is Inadequate.- 1.1 Natural Selection Is a Negative Process.- 1.2 Populations Press Against Limits of a Minimum Resource.- 1.3 What Essential Resource Is Most Likely to Be Limiting?.- 1.3.1 Nitrogen the Most Limiting Chemical.- 1.3.2 Nitrogen Limiting Plants.- 1.3.3 Nitrogen Limiting Animals.- 1.3.4 Energy Not Limiting.- 1.4 Competition a Consequence Not a Cause.- 1.4.1 Intra-specific Competition.- 1.4.2 Inter-specific Competition and "Competitive Exclusion".- 1.5 Self-regulation Does Not Exist.- 2. Plants as Food for Herbivores.- 2.1 Why Is the World Green?.- 2.2 How Might Plants Be an Inadequate Source of Food?.- 2.3 How and When Might Nutrients in Plants Be Too Dilute?.- 2.4 When Is a Minimum Supply of Nitrogen Critically Important?.- 2.5 How Might Herbivores Counter the Plants' Evolved Strategies?.- 2.6 How General Is Dilution of Nitrogen in Herbivore Food, and What Adaptations Have Evolved to Counter it?.- II: Herbivores in an Inadequate Environment.- 3. Insects.- 3.1 Flush and Senesence Feeders.- 3.1.1 Two Australian Psyllids on Eucalyptus.- 3.1.2 Two African Scale Insects on Californian Ice Plants.- 3.1.3 Two Aphids on Scots Pine.- 3.1.4 The Green Spruce Aphid in Scotland.- 3.1.5 Aphids on Sycamore, Apple, Wheat, and Alfalfa.- 3.1.6 Scale Insects on Euphorbia and Euonymus.- 3.1.7 A Leafhopper with Alternate Generations on Brambles and Oak.- 3.1.8 Two Species of Caterpillars Eating Oak Leaves.- 3.1.9 Two Species of Sawflies Mining in Birch Leaves.- 3.1.10 A Chewer and a Sucker on Poplar Leaves.- 3.2 Leaf-miners.- 3.2.1 The Switch from Flush to Senescence Feeding.- 3.2.2 Leaf-miners Which Induce "Green Islands" in Leaves.- 3.2.3 A New Zealand Weevil Mining in Fallen Beech Leaves.- 3.2.4 The American Holly Leaf-miner.- 3.3 Gall Makers.- 3.3.1 Physiological Galls.- 3.3.2 Nutritional Benefits of Galling.- 3.3.3 Adaptive Nature of Galls Debated Anew.- 3.3.4 Double-dipping: Prolonged Growth plus Hastened Senescence.- 3.3.5 Selection for High Nitrogen and Survival of the Young.- 3.3.6 Selection of Growing Tissues for Proliferation of Galls.- 3.4 Chewing Insects.- 3.4.1 Creaming-off as a Tactic to Increase Access to Nitrogen: White Butterflies on Crucifers.- 3.4.2 Early Instars Need More Nitrogen: Gypsy Moth on Artificial Diet.- 3.4.3 Illustrations from the Life Cycles of Economically Unimportant Butterflies.- 3.4.4 Further Examples from Forest Defoliators.- 3.4.5 Pests of Crops also Reveal the Need for Nitrogen.- 3.4.6 Examples from Biological Control of Weeds.- 3.4.7 Locusts and Grasshoppers.- 3.5 Sap-Sucking Insects.- 3.5.1 Aphids.- 3.5.2 Psyllids.- 3.5.3 Scale Insects.- 3.5.4 Planthoppers, Leafhoppers, and a Mirid.- 3.5.5 Xylem-feeders.- 3.6 Fruit Flies.- 3.7 Wood-eating Insects.- 3.7.1 The Key Role of Fungi: Increasing Nitrogen in Wood.- 3.7.2 Termites and Woodroaches: Gut Fauna, Coprophagy, and Recycled Nitrogen.- 3.7.3 Furniture and Longhorn Beetles.- 3.7.4 Woodwasps.- 3.7.5 Borers Which Do Not Ingest Wood.- 4. Crustaceans.- 4.1 Microcrustaceans.- 4.1.1 Distribution and Abundance of Food Limited by Nitrogen.- 4.1.2 Nitrogen Content of Food also Important.- 4.1.3 Microcrustaceans Feed Selectively for Nitrogen.- 4.1.4 Coprorhexy.- 4.2 Macrocrustaceans: Land Crabs, Lobsters, and Shrimps.- 4.3 Terrestrial Crustaceans - The Isopods.- 4.3.1 The Case of a Common Woodlouse.- 4.3.2 The Role of Microorganisms and Coprophagy.- 5. Molluscs.- 5.1 Some Examples of Freshwater Snails.- 5.2 Marine Limpets and the Flow of Nitrogen Through the Food Chain.- 5.3 Death of the Young, Selective Feeding, and Animal Protein in the Diet.- 5.4 Detritus Feeders Feed Selectively, and Depend upon Microorganisms and Coprophagy.- 5.5 Terrestrial Snails Live with the Same Constraints.- 5.6 Cannibalism by Young Snails Illustrates Shortage of Nitrogen.- 5.7 Teredo Shipworms Depend upon Microorganisms Which Fix Atmospheric Nitrogen.- 6. Mammals.- 6.1 Large Mammals.- 6.1.1 Feral Donkeys in Australia.- 6.1.2 Red Deer in Scotland.- 6.1.3 Antelope, Giraffe, and Greater Kudu in Africa.- 6.1.4 Deer in North America.- 6.1.5 The Giant Panda in China.- 6.1.6 Domestic Stock.- 6.2 Rodents.- 6.2.1 Squirrels.- 6.2.1.1 True Squirrels.- 6.2.1.2 Chipmunks and Ground Squirrels.- 6.2.2 Rats and Mice.- 6.2.2.1 The House Mouse.- 6.2.2.2 The Australian Smokey Mouse.- 6.2.2.3 The American White-Footed Mouse.- 6.2.2.4 American Woodrats.- 6.2.3 Voles.- 6.2.4 Supplemental Feeding of Small Rodents.- 6.2.5 Rabbits and Hares.- 6.2.5.1 The European Mountain Hare.- 6.2.5.2 The European Rabbit.- 6.2.5.3 The North American Snowshoe Hare.- 6.3 Primates.- 6.3.1 Colobine Monkeys.- 6.3.2 Cercopithecid Monkeys.- 6.3.3 Howler Monkeys.- 6.3.4 The Gorilla.- 6.4 Fruit and Flower Bats.- 6.5 Marsupials.- 6.5.1 The Koala.- 6.5.2 Possums and Gliders.- 6.5.3 The Habitat of Possums and Gliders.- 6.5.4 Kangaroos and Wallabies.- 7. Birds.- 7.1 Birds Eating Green Leaves.- 7.1.1 Geese in Europe and North America.- 7.1.2 European Grouse, Ptarmigan, and Capercaillie.- 7.1.3 North American Grouse.- 7.1.4 Partridges and Pheasants.- 7.1.5 Galliforms as Hindgut Fermenters.- 7.1.6 Changes in Abundance of Lagopus Species.- 7.1.7 The Takahe.- 7.1.8 The Hoatzin.- 7.2 Birds Eating Nectar and Fruit.- 7.3 Birds Eating Seeds.- 7.3.1 Columbids.- 7.3.2 African Queleas, European Finches, and the Great Tit.- 7.3.3 Darwin's Galapagos Finches.- 7.3.4 The Australian Galah.- 8. Reptiles.- 8.1 The Giant Tortoises of Aldabra Atoll.- 8.2 The Green Turtle of the Bahamas Islands.- 8.3 The Marine and Terrestrial Iguanids of the Galapagos Islands.- 8.4 The Desert Iguanid of California.- 8.5 The Green Iguanid of Panama.- 9. Fish.- 9.1 The Carnivorous Young of Fish.- 9.2 Fish Which Eat Detritus.- 9.3 Fish Which Eat Algae.- 9.4 Gut Microbes and Coprophagy in Fish.- III: Survival in an Inadequate Environment.- 10. Strategies to Counter Shortage of Nitrogen.- 10.1 Strategy A: Synchronize the Life Cycle with Availability of Good Food.- 10.2 Strategy B: Concentrate or Prolong Availability of Nitrogen in Food.- 10.3 Strategy C: Eat More Food More Quickly, and Digest More Efficiently.- 10.4 Strategy D: Enlist the Help of Microorganisms.- 10.5 Strategy E: Supplement Plant Food with Animal Protein.- 10.6 Strategy F: Apportion and Concentrate the Limited Food to a Select Few.- 11. Territorial and Social Behaviours.- 11.1 Territorial Behaviour in Carnivores.- 11.1.1 Birds.- 11.1.2 Lizards.- 11.1.3 Insects.- 11.1.4 Spiders.- 11.2 Territorial Behaviour in Herbivores.- 11.2.1 Mammals.- 11.2.2 Birds.- 11.2.3 Insects.- 11.2.4 Fish.- 11.2.5 Molluscs.- 11.3 Surplus Young, Dispersal, and Philopatry.- 11.4 Social Structures and Dominance Hierarchies.- 12. Cannibalism.- 12.1 Cannibalism by Females Producing Young.- 12.2 Cannibalism by Growing Young.- 12.3 Cannibalism, Warfare, and Protein.- IV: Predators in an Inadequate Environment.- 13. Vertebrates.- 13.1 Lions, Lynx, and Feral Cats.- 13.2 Coyotes, Wolves, and Foxes.- 13.3 Stoats, Mice, and Seed Mast.- 13.4 Pelicans, Puffins, and Other Sea Birds.- 14. Invertebrates.- 14.1 Triclad Worms.- 14.2 Spiders and Scorpions.- 14.3 Ground Beetles, Tiger Beetles, and Ant-lions.- 14.4 Praying Mantids.- 14.5 Parasitoids, Parasites, and Diseases.- V: The Alleviation of an Inadequate Environment: Outbreaks.- 15. What is an Outbreak.- 15.1 Some Examples.- 15.2 What Causes Outbreaks?.- 15.3 The Paradox of Enrichment and "r" and "K" Strategists.- 16. The Interaction of Food, Prey, and Predators in Outbreaks.- 16.1 Bacteria and Protozoa.- 16.2 Rabbits, Foxes, Cats, and Dingoes.- 16.3 The Varying Response of Predators to Changes in Prey.- 16.4 A Natural Experiment: Guano-Algae-Limpets-Oystercatchers.- 16.5 A Thought Experiment: Hot Spots in a Box of Wadding.- 17. Cyclic Outbreaks.- 18. The Influence of Weather on the Generation of Outbreaks.- 18.1 Hot Spots Again: Outbreak Centres and Boundaries.- 18.2 Spruce Budworm Outbreaks Revisited.- 18.3 Patchy Environments and Metapopulations.- 18.4 The Role of Viral Diseases.- 18.5 The Link to Climatic Oscillations.- 18.6 Major Outbreaks Which Are Independent of the Weather.- References.