Table of Contents

Birds and Humans: A Historical Perspective Birds as Food H-3 Use of Skins and Feathers H-4 Birds in Literature, Culture, and Religion H-6 Art H-6 Religion H-9 Folklore H-10 Literature H-11 Music and Dance H-13 Music and Dance of Indigenous Cultures H-13 Music and Dance of Western Cultures H-18 The Evolution of North American Ornithology H-19 The Early Years: From Aristotle to the 17th Century H-19 The 18th Century H-23 The 19th Century H-25 The American Ornithologists' Union and the U.S. Biological Survey H-32 The First Audubon Movement H-33 The Second Audubon Movement H-34 The 20th Century and the Expanding Role of the Bird Watcher H-35 The Development of the Field Guide H-36 Academic Training in Ornithology H-37 Bird Conservation, Bird Watching and the Age of Technology H-39 Suggested Readings H-42 Cornell Laboratory of Ornithology Chapter 1 Introduction: The World of Birds Ornithological Terms 1-3 The Form of a Bird 1-3 Bill 1-6 Head and Neck 1-7 Trunk 1-9 Wings 1-9 Tail 1-12 Hind Limbs 1-12 Diversity in Bird Form 1-15 The Bill 1-16 The Wings 1-18 The Tail 1-19 The Feet 1-20 Feathering 1-23 Internal Anatomy 1-24 Diversity in Bird Movement 1-24 Movement on Land 1-24 Movement in Water 1-28 Naming and Classification of Birds 1-32 History 1-32 Methods Used to Classify Birds 1-40 Binomial Nomenclature and Classification System 1-45 The Species 1-53 The Formation of Species 1-55 Orders and Families of World Birds 1-61 Orders and Families of North American Birds 1-64 How Naming and Classification Can Help You 1-64 The Use of Common Names 1-64 Evolution of Birds and Avian Flight 1-66 Bird Distribution 1-66 Distribution of Land Birds 1-69 Palearctic Region 1-69 Nearctic Region 1-71 Neotropical Region 1-72 Afrotropical Region 1-81 Oriental Region 1-88 Australasian Region 1-91 Island Distribution 1-97 Distribution of Marine Birds 1-98 Northern Marine Region 1-99 Southern Marine Region 1-100 Tropical Marine Region 1-101 Plankton and Bird Distribution 1-101 The Importance of Biodiversity 1-106 Appendix A: Orders and Families of World Birds 1-107 Appendix B: Orders and Families of North American Birds 1-111 Appendix C: Geological Time Scale 1-113 Sidebars Sidebar 1: Which Way is Up? 1-4 Sidebar 2: The Evolution of an Idea: Darwin's Theory 1-34 Sidebar 3: Latin and Greek Roots of Biological Terms 1-48 CHAPTER 2: A Guide to Bird Watching How to Identify Birds 2-4 Shape 2-5 Postures and Flight Patterns 2-5 Behaviors 2-7 Size 2-7 Comparing Body Features 2-9 Field Marks 2-10 Head 2-10 Bill Shape and Color 2-11 Wings 2-12 Tail 2-12 Legs 2-13 Colors and Plumage Patterns 2-13 Songs 2-14 Habitat 2-18 Range and Abundance 2-19 Time of Year 2-19 Sorting Out Birds 2-23 Closing the Distance 2-23 Sitting Quietly 2-23 Pishing and Squeaking 2-26 Mobbing 2-26 Playback Songs 2-28 Bird Blinds 2-29 Viewing Birds 2-29 Using Binoculars 2-29 Pointing Out Birds to Others 2-32 Selecting Binoculars 2-34 Magnification Power 2-34 Light-gathering Capacity 2-35 Field of View 2-37 Resolution 2-37 Alignment 2-38 Binocular Designs 2-38 Mini Binoculars 2-39 Binoculars for Eyeglass Wearers 2-40 How to Shop for Binoculars 2-40 How to Clean Binoculars 2-41 Protecting Binoculars 2-41 Selecting a Spotting Scope and Tripod 2-42 How to Shop for a Spotting Scope 2-44 Recording Observations 2-44 Checklists 2-46 Journals 2-47 Reporting Rare Birds 2-54 Listing Birds 2-54 Counting Birds 2-55 Cornell Laboratory of Ornithology Conclusion 2-57 The Birder's Essential Resources 2-58 Field Guides 2-58 General References 2-59 Textbooks 2-60 Popular Bird Magazines 2-60 Scientific Journals 2-61 Leisure Reading 2-62 Audio Guides 2-63 Web Sites 2-63 Sidebars Sidebar 1: Attracting Birds to Your Yard 2-24 Sidebar 2: How to Calibrate Binoculars For Your Eyes 2-30 Sidebar 3: Sketching Birds in the Field 2-50 CHAPTER 3: Form and Function: The External Bird Feather Tracts 3-2 Feather Form and Function 3-3 Feather Structure 3-3 Types of Feathers 3-4 Contour Feathers 3-5 Down Feathers 3-16 Semiplumes 3-17 Filoplumes 3-17 Powder Downs 3-18 Care of Feathers 3-18 Preening 3-18 Oiling 3-20 Head-Scratching 3-20 Bathing 3-21 Sunning 3-22 Anting 3-22 Ectoparasites 3-23 Development of Feathers 3-26 Molts and Plumages 3-28 Annual Molt and Wear Cycles 3-29 Subadult and Definitive Plumages 3-30 Plumage Naming Systems 3-33 The Progression of a Molt 3-34 Nonfeathered Areas 3-39 Eyes 3-39 Bill 3-39 Legs and Feet 3-43 Other Unfeathered Areas 3-46 Colors 3-48 Pigments 3-50 Abnormalities and Variations in Pigment Colors 3-52 Structural Colors 3-54 Functions of Color and Color Patterns 3-60 Cryptic Coloration and Patterns 3-60 Blending In 3-61 Disruptive Coloration 3-62 Countershading 3-63 Behaviors that Aid Concealment 3-63 Conspicuous Markings and Predation 3-64 Reduction of Glare for Foraging 3-64 The Role of Color and Pattern in Social Behavior 3-65 Species Recognition 3-66 Age Recognition 3-66 Sex Recognition 3-67 Individual Recognition 3-67 Flock Attraction 3-68 Sexual Selection 3-68 Sidebars Sidebar 1: Feather Detective 3-6 Sidebar 2: Feather Facts 3-11 Sidebar 3: Iridescence 3-56 CHAPTER 4: What's Inside: Anatomy and Physiology The Skeletal System 4-3 Axial Skeleton 4-10 Skull 4-10 Hyoid Apparatus 4-13 Vertebral Column 4-14 Appendicular Skeleton 4-18 Pectoral Girdle 4-19 Bones of the Wing 4-21 Sternum 4-23 Pelvic Girdle 4-24 Bones of the Hind Limb 4-24 The Muscular System 4-26 Skeletal Muscle 4-26 Smooth Muscle 4-28 Cardiac Muscle 4-31 The Nervous System 4-31 The Neuron 4-32 Sensory and Motor Neurons 4-33 Central Nervous System 4-35 Brain 4-36 Spinal Cord 4-38 Peripheral Nervous System 4-40 Cranial Nerves 4-40 Spinal Nerves 4-42 Autonomic Nervous System 4-43 The Senses 4-45 Vision 4-45 The Structure of the Eye 4-46 How Birds See 4-50 The Ear and Hearing 4-54 Structure and Function of the Ear 4-56 Hearing Ability 4-61 Olfaction 4-62 Taste 4-65 Skin Senses 4-69 The Endocrine System 4-69 Pituitary Gland 4-72 Thyroid Glands 4-74 Parathyroid and Ultimobranchial Glands 4-74 Adrenal Glands 4-74 Gonads 4-75 Pancreas 4-75 The Circulatory System 4-76 The Heart 4-77 Heart Valves 4-78 Blood Supply to the Heart Tissue 4-79 Conducting System of the Heart 4-79 Location of the Heart 4-80 Blood Vessels 4-81 Capillaries 4-81 Arterial System 4-82 Venous System 4-84 Blood 4-86 Lymphatic System 4-88 The Respiratory System 4-89 Nostrils and Nasal Cavities 4-90 Pharynx 4-91 Larynx 4-91 Trachea 4-92 Syrinx 4-93 Lungs and Air Tubes 4-98 Air Sacs 4-100 Breathing and Gas Exchange 4-100 The Digestive System 4-103 Oral Cavity 4-103 Bill 4-103 Tongue 4-111 Salivary Glands and Saliva 4-111 Pharynx 4-112 Alimentary Canal 4-113 Esophagus 4-113 Stomach 4-118 Small Intestine 4-120 Colic Ceca 4-123 Large Intestine 4-123 Cloaca 4-123 Liver 4-124 The Urogenital System 4-124 Urinary System 4-125 Genital System 4-127 Male Genitals 4-127 Female Genitals 4-128 Copulation and Fertilization 4-133 Sex Determination 4-133 Handbook of Bird Biology Hormones and Secondary Sex Characters 4-137 Factors Bringing Birds into Breeding Condition 4-140 Metabolism 4-144 Body Temperature 4-146 Countercurrent Heat-Exchange Systems 4-148 Cooling 4-152 Torpor 4-153 Heart Size and Heart Rate 4-154 Respiratory Rate 4-156 Water and Salt Regulation 4-157 Life Span and Senescence 4-158 Major Anatomical Differences between Birds and Mammals 4-161 Skeleton 4-161 Muscles 4-161 Nervous System 4-161 Ear 4-162 Eye 4-162 Circulatory System 4-162 Respiratory System 4-162 Digestive System 4-163 Urogenital System 4-163 Suggested Readings 4-163 Sidebars Sidebar 1: The Amazing World of Avian ESP 4-66 Sidebar 2: Bird Song: From Oboe and Trombone to Orator and Soprano 4-94 CHAPTER 5: Birds on the Move: Flight and Migration The Flight Syndrome 5-2 Functions of the Flight Muscles 5-7 How Do Birds Fly? 5-8 Forces Acting on a Bird in Flight 5-9 Gravity 5-10 Lift 5-10 Drag 5-16 Thrust 5-16 Function of the Tail 5-21 Landing 5-21 Hovering 5-26 Complex Control of Flight 5-30 Wing Loading 5-31 Turbulence 5-33 Variations in Wing Shape and Flight Style 5-36 Elliptical Wings 5-37 High-speed Wings 5-38 Slotted High-lift Wings 5-39 High-Aspect-Ratio Wings 5-42 Some Flight Facts and Figures 5-45 Air Speed 5-45 Cornell Laboratory of Ornithology Wingbeat Frequency 5-45 Flocking and Flying in Formation 5-45 Loss of Flight 5-48 Migration 5-51 Patterns of Migration 5-52 The Origin and Evolution of Migration 5-57 Controlling and Synchronizing the Annual Cycle 5-61 The Physiology of Migration 5-63 Daily Timing of Migration 5-65 The Altitude of Migration 5-66 Flight Speed and the Progress of Migration 5-68 Weather and Migration 5-69 Migration Routes 5-73 Site Fidelity 5-75 Orientation and Navigation 5-79 Compass Mechanisms 5-84 Sun Compass 5-84 Star Compass 5-86 Magnetic Compass 5-89 Navigational Maps 5-92 Sunset Cues 5-92 Suggested Readings 5-99 Sidebars Sidebar 1: Flapping Flight 5-22 Sidebar 2: Showdown at Delaware Bay 5-76 Sidebar 3: Polarized Light 5-94 Evolution of Birds and Avian Flight Archaeopteryx and Other Urvogels 2 The Descent of Birds 7 Flight Origins 13 Ground-Up (Cursorial) Theory 14 Trees-Down (Arboreal) Theory 18 Early Bird Flight 19 The Early Fossil Record of Birds 20 Palaeognathous Birds 21 Bird Evolution's Big Bang 25 Appendix A: Bird Evolution Theories and Early Diapsid Reptiles 27 Appendix B: Hypothesized Relationships Among Ancient and Modern Bird Groups 29 Appendix C: Index to Fossil Organisms 31 Figure Credits for Appendix C 34 CHAPTER 6: Understanding Bird Behavior Questions About Behavior 6-2 The Proximate Basis of Bird Behavior 6-4 Ethology, Ornithology, and Instincts 6-7 Learned Behavior 6-8 A Comparison of Instincts and Learning 6-14 Ultimate Causes of Bird Behavior 6-15 Territoriality, Dominance Hierarchies, and Ritualized Aggression 6-22 The Evolution of Ritualized Displays 6-30 Courtship Displays 6-34 The Use of Darwinian Evolutionary Theory 6-42 Feeding Behavior: Why do Birds Generally Restrict their Diets, Ignoring Some Edible Foods in Favor of Others? 6-43 Antipredator Behavior: Why do Some Birds Mob Predators? 6-50 Nest Spacing: Why do Some Birds Nest in Large Colonies? 6-58 Reproductive Behavior: Why Are There Different Kinds of Avian Mating Systems? 6-68 Reproductive Behavior: Resource-defense and Female-defense Polygyny 6-73 Reproductive Behavior: Lek Polygyny 6-75 Reproductive Behavior: Polyandry 6-77 Mate Choice: Extrapair Copulations in Birds 6-79 Mate Choice: Why do Some Birds Display Elaborate Ornaments? 6-81 Mate Choice: Why Cooperate in Courtship Displays? 6-85 Parental Behavior: Why do Some Birds Ignore Lethal Aggression Among their Nestlings? 6-87 Parental Behavior: Why are there "Helpers at the Nest" that Care for Someone Else's Offspring? 6-88 How to Study Bird Behavior Yourself 6-91 Suggested Readings 6-98 Sidebars Sidebar 1: Bird Brains 6-16 Sidebar 2: Play 6-19 Sidebar 3: Defense Behavior 6-52 Sidebar 4: Living in Groups 6-60 Sidebar 5: Length of the Pair Bond 6-72 Sidebar 6: Bird Families as Models for Understanding Ourselves 6-92 CHAPTER 7: Vocal Behavior What is Sound? 7-3 Seeing Sounds: Sonagrams and Oscillograms 7-4 Use of Tape or CD with Chapter Text 7-6 Understanding Complex Songs 7-9 Vocal Repertoires 7-10 The Problem of Meaning 7-11 Song 7-14 The Structure and Function of Sounds 7-19 Vocal Development 7-23 Vocal Development in Songbirds 7-25 Vocal Development in Nonsongbirds 7-30 Songbird Diversity 7-34 Control of Song 7-37 Variation in Space and Time 7-41 Cornell Laboratory of Ornithology Species Differences 7-42 Individual Variation 7-42 Song Dialects 7-53 Geographic Variation in Suboscine Vocalizations 7-57 The Diversity of Geographic Patterns in Songbirds 7-58 Song Change Over Time 7-63 Dialects Over Broad Regions 7-64 The Functions of Song 7-66 Dawn Chorus 7-75 Duetting 7-78 Mimicry 7-81 Flight Songs 7-84 Song Repertoires 7-85 Suggested Readings 7-91 Appendix A: Descriptions of Tape/CD Tracks 7-93 Sidebars Sidebar 1: Winnows, Snaps, and Spring Thunder--Nonvocal Sounds 7-15 Sidebar 2: Listen Up!.7-44 Sidebar 3: Pushing the Limits: New Computer Techniques for Studying Bird Song 7-48 Sidebar 4: Do Birds Think? 7-67 Sidebar 5: "Call Notes" and Their Functions 7-72 Sidebar 6: Listening on Your Own 7-88 CHAPTER 8: Nests, Eggs, and Young: Breeding Biology of Birds Survival 8-3 The Timing of Breeding 8-10 Breeding Territories 8-13 Functions of Breeding Territories 8-14 Nests and Nest Building 8-15 Functions of Nests 8-18 Diversity of Nest Sites 8-20 Seasonal Changes in Nest Sites 8-22 Nest Site Selection 8-23 Diversity of Nests 8-24 The Evolution of Nest Construction 8-44 Nest Lining 8-47 Nest-building Behavior 8-50 Sex Roles in Nest Building 8-56 Duration of Nest Building 8-57 Nest Appropriation and Reuse 8-57 Eggs 8-59 Egg Structure 8-59 Egg Size 8-70 Egg Shape 8-72 Egg Surface Texture 8-74 Egg Color 8-75 Egg Laying 8-77 Clutch Size 8-78 Patterns in Clutch Size Variation 8-78 The Evolution of Clutch Size 8-79 Egg and Clutch Replacement 8-90 Number of Broods per Season 8-91 Incubation 8-93 Incubation Patch 8-94 Incubation Period 8-96 Start of Incubation 8-97 Role of the Sexes 8-99 Patterns of Attentiveness 8-99 Behavior During Incubation 8-101 Changes in Incubation Behavior 8-103 Feeding the Mate 8-103 Development of Young 8-104 Hatching 8-10_ Development at Hatching 8-106 Typical Altricial Young 8-107 Typical Precocial Young 8-117 Recognition Between Parents and Young 8-125 Caring for Young 8-130 Feeding the Young 8-131 Defending the Young 8-134 Nest Sanitation 8-136 Brood Parasites 8-139 Evolution and Adaptation Among Obligate Brood Parasites 8-143 Brood Parasite Ploys 8-143 Host Counterploys and Coevolution 8-146 Evolution and Adaptation in New World Cowbirds 8-148 Conclusion 8-152 Suggested Readings 8-152 Sidebars Sidebar 1: Neat Nesting Facts 8-16 Sidebar 2: Social Weavers 8-40 Sidebar 3: Oology: From Hobby to Science 8-80 Sidebar 4: Creches 8-126 CHAPTER 9: Individuals, Populations, and Communities: The Ecology of Birds Birds as Individuals 9-7 Habitat Selection: Choosing a Place to Live 9-8 Thermoregulation: Coping with Heat and Cold 9-13 Water: A Matter of Economy 9-18 Foraging Ecology: Meeting Energy and Nutritional Demands 9-23 How Much Food Does a Bird Need? 9-23 What Types of Food are Eaten? 9-25 Where and How to Forage 9-28 Do Birds Always Forage Optimally? 9-31 Coping with Environmental Fluctuations 9-31 Relationships with Other Individuals 9-38 Cornell Laboratory of Ornithology Types of Intraspecific Competition 9-39 Life History Strategies: Putting it All Together 9-43 Birds in Populations 9-48 Characteristics of Bird Populations 9-49 Geographic Distribution Patterns 9-49 Population Size 9-58 How Do We Determine Population Size? 9-60 What Affects Population Size? 9-62 What Regulates Population Size? 9-67 Extinction: The Death of the Last Individual in a Population 9-75 Structure of Bird Populations 9-80 Bird Communities 9-82 Characteristics of Bird Communities 9-82 Patterns of Species Richness 9-85 Effects of Latitude 9-85 Effects of Habitat Complexity and Productivity 9-92 Effects of Habitat Size 9-95 Habitat Patches as "Islands" 9-97 Patterns of Relative Abundance 9-101 Ecological Niches 9-102 Are Bird Communities Organized in Optimal Ways? 9-104 Birds as Components of Ecosystems 9-109 Ecological Distribution of Birds in the Major Terrestrial Ecosystems of North America 9-109 Tundra 9-114 Coniferous Forest 9-116 Deciduous Forest 9-117 Grassland 9-118 Southwestern Oak Woodland 9-120 Chaparral 9-120 Pinyon-Juniper Woodland 9-121 Sagebrush 9-121 Scrub Desert 9-122 Two Important Ecotones 9-122 The Role of Birds in the Food Chain 9-123 What if Birds Disappeared? 9-126 Sidebars Sidebar 1: The Winter Banquet 9-32 Sidebar 2: The House Finch Hot Zone 9-71 Sidebar 3: Ant Followers 9-88 Sidebar 4: From Blackberries to Beeches: Ecological Succession in Eastern Deciduous Forests 9-110 Sidebar 5: Sapsuckers, Swallows, Willows, Aspen, and Rot 9-128 CHAPTER 10: Bird Conservation Historical Context 10-4 Global Spread of Humans Begins the Extinction Era 10-5 Early Extinctions in North America and the Caribbean 10-6 Modern Extinctions on Mainland North America 10-9 Labrador Duck --the Mystery Extinction 10-10 Passenger Pigeon --Market Hunting at its Worst 10-11 Carolina Parakeet--Removal of a Menace 10-12 Eskimo Curlew--Three Strikes in the Wink of an Eye 10-12 Ivory-billed Woodpecker and Bachman's Warbler -- Demise of the Southeastern Forests 10-16 Brief History of Bird Conservation in the United States 10-18 Are North American Birds Disappearing? 10-25 The Forested Northeast 10-29 Grasslands 10-30 Southwestern Riparian Habitats 10-36 Shorebirds 10-37 Conservation Problems: The Ecology of Extinction 10-38 Birth Rates and Death Rates 10-39 Population Increases 10-40 Direct Exploitation 10-48 Introduced Predators 10-49 Chemical Toxins 10-51 Indirect Chemical Pollution 10-56 Introduced Disease 10-58 Habitat Loss 10-59 Habitat Specialization and the "Six Forms of Rarity" 10-61 1. Widely distributed, small local populations, broad habitat tolerance 10-62 2. Widely distributed, large local populations, narrowly specialized habitat requirements 10-63 3. Widely distributed, small local populations, narrowly specialized habitat requirements 10-64 4. Small geographic range, large local populations, broad habitat tolerance 10-65 5. Small geographic range, large local populations, narrowly specialized habitat requirements 10-65 6. Small geographic range, small local populations, narrowly specialized habitat requirements 10-66 Unique Problems on Islands 10-67 Habitat Fragmentation: Mainland Habitats as Islands 10-71 Conservation Genetics 10-74 Conservation Solutions: Tools and Prescriptions for Stabilizing Populations 10-76 DNA Fingerprinting and Genetic Augmentation 10-76 Population Viability Analysis and Metapopulations 10-77 Preserve Design 10-78 Habitat Management 10-81 Ecosystem Management 10-83 Adaptive Management 10-84 Translocation 10-89 Legal Protection 10-90 Endangered Species Act 10-91 Clean Water Act, Section 404 10-93 CITES 10-93 Bringing Birds Back from the Brink 10-94 Wood Duck --Regulated Hunting and Cornell Laboratory of Ornithology Adaptive Management 10-95 Whooping Crane --Protected Habitat and Captive Rearing 10-97 Peregrine Falcon--Pesticide Regulation and "Soft Release" Reintroduction 10-100 California Condor--Wild Capture, Captive-rearing, and Study of "Surrogates" 10-102 Maui Parrotbill and Akohekohe -- Protected Habitat and Feral Mammal Control 10-103 Why Protect Birds? 10-104 Direct Benefits 10-105 Food 10-105 Clothing 10-105 Other Utilitarian Uses 10-105 Recreational Hunting 10-105 Bird Watching 10-106 Indirect Benefits 10-106 Ecological and Evolutionary Roles 10-106 Environmental Services 10-107 Biological Indicators 10-108 Genetic Information 10-109 Scientific Study 10-109 Aesthetics and Spiritual Values 10-110 What Can Each of Us Do? 10-110 Backyard Conservation 10-110 Be a Citizen Scientist 10-112 Adopt a Place 10-112 Local Vigilance and Grassroots Activism 10-113 Environmental Education 10-114 Take a Child Birding 10-114 Contribute to Conservation Organizations 10-115 Never Give Up 10-116 SIDEBARS Sidebar 1: A Summer Without Bobolinks 10-31 Sidebar 2: The Best Laid Plans: What Happens When Conservation Efforts Work Too Well? 10-44 Sidebar 3: Hawk Deaths Spur Action 10-54 Sidebar 4: Conservation Planning at Ecoregional Scales 10-86

About the Author

Cornell Laboratory of Ornithology is a nonprofit membership institution working to understand and conserve the earth's biological diversity through research, education, and citizen science focused on birds.

Reviews

If you really do want to know everything about birds, turn to the second edition of the Cornell Lab of Ornithology's Handbook of Bird Biology, which for really keen readers comes in conjunction with a home-study course in the subject. -- Stephen Moss The Guardian