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Table of Contents
PREFACE AND ACKNOWLEDGEMENTS
SECTION I IN THE BEGINNING
CHAPTER 1. WADING IN INTRODUCTION TO THE FISH-BIRD
General Discussion of Penguins and Substance of the Book
An Impressive Number of Penguin Species: Evolution of Their Unique Capabilities
Penguin Species Radiation and the Ontogeny of Their Watery World
Penguin Evolution: Body Size and Climate
Penguin Evolution: Radiation into Vacant Niches
Penguin Evolution: Body Size and Crossing the Sea-Land Boundary
CHAPTER 2. LAND AHOY! A TIRESOME BUSINESS
Crossing the Land-Ocean Interface is Affected by Body Size
Why and How Often do Penguins Come Ashore?
Tying Land-life to the At-sea Life of Fish-Birds: Foraging and Breeding
Success Vary with Prey Availability
Molt Necessary, Brief Respite from the Sea
SECTION II PENGUIN MARINE HAUNTS AND FOOD HABITS
CHAPTER 3.FISH-BIRDS AT HOME IN THEIR OCEAN HABITATS
Oceanographic Fronts and Water Masses Important to Penguins: General Discussion
Penguins Require High Productivity Water Masses
Large Scale: Oceanographic Boundaries and At-sea Distributions of Penguins
Southern Boundary of the Antarctic Circumpolar Current
Emperor and King penguins
Adlie and Chinstrap penguins
Antarctic Polar Front and Subantarctic Front
Gentoo and Yellow-eyed penguins
Macaroni and Royal penguins
Northern and Southern Rockhopper, Fiordland, Snares penguins
Subtropical Front and Continental Boundary Currents
Galpagos, Humboldt, Magellanic, African penguins
Little penguins
Meso- and Small-Scale Ocean Processes Facilitating Penguin Exploits
Island wakes
Headland wakes
Shelves and banks
Submarine canyons
Shelfbreak fronts
Marginal ice zones
Thermo-/haloclines
CHAPTER 4. SEA FOOD THE FISH-BIRD MENU
General Considerations
Diet Quality: Survival in Cold Water
Energy density of prey
Prey size may or may not differ by penguin size
Prey availability
Diet Comparison among Penguin Species
Polar/subpolar, mesopelagic penguins
Subpolar, demersal/benthic, continental-shelf penguins
Temperate, upper water column, continental-insular shelf penguins
Polar, upper water column, continental shelf/slope penguins
Subpolar, upper water column, continental slope/pelagic penguins
Polar, upper water column, continental slope/pelagic penguins
CHAPTER 5. ECOLOGICAL CONSEQUENCES OF DIET COMPOSITION
Intraspecific Competition among Penguins
Foraging Range is Key: General Discussion
Penguin Species Central-Place Foraging Range Patterns
Sex Differences in Foraging
Interspecific Competition Involving Penguins
Penguins Eat A lot!
Competition between Penguin Species
Competition between penguins and other seabirds
Competition between penguins and marine mammals
Competition between penguins and industrial fisheries
SECTION III THE HARDWARE OF A FISH-BIRD
CHAPTER 6. THE SLIPPERY SHAPE, HOT AIR AND THE POWERHOUSE HOW FISH-BIRDS SWIM
Water Hard Taskmaster
The Four Forces Relevant to Penguins
Vertical Forces Weight and Upthrust
Buoyancy: How Much Air Do Penguins Hold?
Buoyancy and Bergmanns Rule revisited
Horizontal Forces: Drag
The Interplay of Drag and Upthrust in Gliding Penguins
The Drag Devil is in the Detail
The Penguin Powerhouse
How Penguins Swim
The Effect of Upthrust and Body Angle on Penguin Thrust and Lift Forces
Top Speeds; Power and Upthrust
The Energy Costs of Swimming
General considerations
Specific considerations
Sensible Swim Strategies and Costs of Transport
Cruising speed and integrating speed with the cost of transport,
and beyond
CHAPTER 7. HOT PENGUINS COLD WATER
Resting and Floating Penguins
The Metabolic Rate of Floating Penguins
Patterns of Heat Loss to the Sea
Overall Body Insulation/Conductance
The Nature of Penguin Insulation
Active Penguins
Activity Produces Heat
Greater Depths Impose a Higher Heat Tax
Consuming Prey Imposes a Heat Tax
Embracing The Fish in the Fish-bird
CHAPTER 8. FISH-BIRDS THE INSIDE STORY
Diving Physiology
Surface issues Uptake of oxygen
Oxygen Management Underwater
Role of the Air Spaces
Gas Exchange to Body Tissues
The Aerobic Dive Limit and Beyond
The Importance of Size in Dive Performance
Duration
Depth
Penguins Under Pressure Beating the Squeeze and the Bends
Barotrauma
Beating the Bends
A Gut Reaction in Fish-Birds
Gastric Emptying
Rotting Food
The Eyes Have It
SECTION IV THE SOFTWARE OF FISH-BIRDS
CHAPTER 9. EMBRACING THE DEPTHS - THE PENGUIN DIVE
Submergence
The time underwater basic dive descriptors
Dive profiles
Dive distance-depth profiles
Dive aspect ratios
Horizontal dive directionality/tortuosity
The Multifunctionality of Dives
Basic dive types
T-dives for travelling
V-dives (water column assessment)
P-dives (parabolic prospecting with no prey capture)
Po-dives (parabolic dives with circular trajectory)
U-dives (depth-directed prospecting)
W-dives/Up-dives (U-dives with prey pursuit)
Depth Duration Effects Over Multiple Dives
CHAPTER 10. FISH-BIRD STRATEGIES THE SEARCH FOR FICKLE PREY
Decisions, Decisions, Decisions How Fish-Birds Search for Prey
Heading in the Right Direction
Dealing with Prey Patchiness
In-depth Considerations:
Time-based efficiency
Energy-based efficiency
Superficial Considerations: Surface Pauses and Inspired Tactics
Being Picky about Food
Fish-Birds and Smart Strategies
CHAPTER 11. THE FINAL SECONDS HOW FISH-BIRDS CAPTURE PREY
Prey Acquisition, a Departure from the Dive Norm
Performance Metrics for Prey Capture
Catching Solitary Prey
Changing buoyancy with depth affects prey capture strategies
Prey pursuit against interfaces
Exploiting Aggregated Prey
Crustaceans
Fish
Non-corralling feeding behavior
Clarity on Limitations of Penguin Vision
SECTION V PENGUINS IN A FICKLE ENVIRONMENT
CHAPTER 12. TURNING THE TABLES FISH-BIRDS ON THE MENU
Basic Law of the Sea: Big Fish Eat Little Fish
Seals as Predators
Seals hunting behavior
Penguins avoiding seals
Fur Seals as Predators
Sea Lions as Predators
Killer Whales as Predators
Sharks as Predators
CHAPTER 13. PENGUINS ADJUSTING TO A CHANGING OCEAN
Penguins Have Always Been Challenged by a Changing Ocean
Prehistoric response to a changing ocean
Possible prehistoric changes to penguins food web
The Anthropocene: How will Penguins Cope, Now Also Dealing with Humans?
Response to long-term climate change
Response to short-term ocean climate variation
Response to marine pollution
CHAPTER 14. NOT FORGETTING
The Social Side - Behavior and Communication at Sea
Penguin flock fusion/cohesion
Penguin flock fission/fragmentation
Navigation
Long range
Medium to short-range
Understanding the Daily Wash
Air flux in diving penguins, an aspect of washing
Are Auks Really Northern Penguins?
Research Tags the Flip Side for Evolutionarily-Honed Fish-Birds
SECTION VI
CHAPTER 15. PENGUINS WHY THE HYPE?
Sources of Hype
Us
Many people
Researchers
The Transition
The fascination of species
Beyond the transition
Role in Ecosystems
Biomimicry
Our Last Word.
SECTION I IN THE BEGINNING
CHAPTER 1. WADING IN INTRODUCTION TO THE FISH-BIRD
General Discussion of Penguins and Substance of the Book
An Impressive Number of Penguin Species: Evolution of Their Unique Capabilities
Penguin Species Radiation and the Ontogeny of Their Watery World
Penguin Evolution: Body Size and Climate
Penguin Evolution: Radiation into Vacant Niches
Penguin Evolution: Body Size and Crossing the Sea-Land Boundary
CHAPTER 2. LAND AHOY! A TIRESOME BUSINESS
Crossing the Land-Ocean Interface is Affected by Body Size
Why and How Often do Penguins Come Ashore?
Tying Land-life to the At-sea Life of Fish-Birds: Foraging and Breeding
Success Vary with Prey Availability
Molt Necessary, Brief Respite from the Sea
SECTION II PENGUIN MARINE HAUNTS AND FOOD HABITS
CHAPTER 3.FISH-BIRDS AT HOME IN THEIR OCEAN HABITATS
Oceanographic Fronts and Water Masses Important to Penguins: General Discussion
Penguins Require High Productivity Water Masses
Large Scale: Oceanographic Boundaries and At-sea Distributions of Penguins
Southern Boundary of the Antarctic Circumpolar Current
Emperor and King penguins
Adlie and Chinstrap penguins
Antarctic Polar Front and Subantarctic Front
Gentoo and Yellow-eyed penguins
Macaroni and Royal penguins
Northern and Southern Rockhopper, Fiordland, Snares penguins
Subtropical Front and Continental Boundary Currents
Galpagos, Humboldt, Magellanic, African penguins
Little penguins
Meso- and Small-Scale Ocean Processes Facilitating Penguin Exploits
Island wakes
Headland wakes
Shelves and banks
Submarine canyons
Shelfbreak fronts
Marginal ice zones
Thermo-/haloclines
CHAPTER 4. SEA FOOD THE FISH-BIRD MENU
General Considerations
Diet Quality: Survival in Cold Water
Energy density of prey
Prey size may or may not differ by penguin size
Prey availability
Diet Comparison among Penguin Species
Polar/subpolar, mesopelagic penguins
Subpolar, demersal/benthic, continental-shelf penguins
Temperate, upper water column, continental-insular shelf penguins
Polar, upper water column, continental shelf/slope penguins
Subpolar, upper water column, continental slope/pelagic penguins
Polar, upper water column, continental slope/pelagic penguins
CHAPTER 5. ECOLOGICAL CONSEQUENCES OF DIET COMPOSITION
Intraspecific Competition among Penguins
Foraging Range is Key: General Discussion
Penguin Species Central-Place Foraging Range Patterns
Sex Differences in Foraging
Interspecific Competition Involving Penguins
Penguins Eat A lot!
Competition between Penguin Species
Competition between penguins and other seabirds
Competition between penguins and marine mammals
Competition between penguins and industrial fisheries
SECTION III THE HARDWARE OF A FISH-BIRD
CHAPTER 6. THE SLIPPERY SHAPE, HOT AIR AND THE POWERHOUSE HOW FISH-BIRDS SWIM
Water Hard Taskmaster
The Four Forces Relevant to Penguins
Vertical Forces Weight and Upthrust
Buoyancy: How Much Air Do Penguins Hold?
Buoyancy and Bergmanns Rule revisited
Horizontal Forces: Drag
The Interplay of Drag and Upthrust in Gliding Penguins
The Drag Devil is in the Detail
The Penguin Powerhouse
How Penguins Swim
The Effect of Upthrust and Body Angle on Penguin Thrust and Lift Forces
Top Speeds; Power and Upthrust
The Energy Costs of Swimming
General considerations
Specific considerations
Sensible Swim Strategies and Costs of Transport
Cruising speed and integrating speed with the cost of transport,
and beyond
CHAPTER 7. HOT PENGUINS COLD WATER
Resting and Floating Penguins
The Metabolic Rate of Floating Penguins
Patterns of Heat Loss to the Sea
Overall Body Insulation/Conductance
The Nature of Penguin Insulation
Active Penguins
Activity Produces Heat
Greater Depths Impose a Higher Heat Tax
Consuming Prey Imposes a Heat Tax
Embracing The Fish in the Fish-bird
CHAPTER 8. FISH-BIRDS THE INSIDE STORY
Diving Physiology
Surface issues Uptake of oxygen
Oxygen Management Underwater
Role of the Air Spaces
Gas Exchange to Body Tissues
The Aerobic Dive Limit and Beyond
The Importance of Size in Dive Performance
Duration
Depth
Penguins Under Pressure Beating the Squeeze and the Bends
Barotrauma
Beating the Bends
A Gut Reaction in Fish-Birds
Gastric Emptying
Rotting Food
The Eyes Have It
SECTION IV THE SOFTWARE OF FISH-BIRDS
CHAPTER 9. EMBRACING THE DEPTHS - THE PENGUIN DIVE
Submergence
The time underwater basic dive descriptors
Dive profiles
Dive distance-depth profiles
Dive aspect ratios
Horizontal dive directionality/tortuosity
The Multifunctionality of Dives
Basic dive types
T-dives for travelling
V-dives (water column assessment)
P-dives (parabolic prospecting with no prey capture)
Po-dives (parabolic dives with circular trajectory)
U-dives (depth-directed prospecting)
W-dives/Up-dives (U-dives with prey pursuit)
Depth Duration Effects Over Multiple Dives
CHAPTER 10. FISH-BIRD STRATEGIES THE SEARCH FOR FICKLE PREY
Decisions, Decisions, Decisions How Fish-Birds Search for Prey
Heading in the Right Direction
Dealing with Prey Patchiness
In-depth Considerations:
Time-based efficiency
Energy-based efficiency
Superficial Considerations: Surface Pauses and Inspired Tactics
Being Picky about Food
Fish-Birds and Smart Strategies
CHAPTER 11. THE FINAL SECONDS HOW FISH-BIRDS CAPTURE PREY
Prey Acquisition, a Departure from the Dive Norm
Performance Metrics for Prey Capture
Catching Solitary Prey
Changing buoyancy with depth affects prey capture strategies
Prey pursuit against interfaces
Exploiting Aggregated Prey
Crustaceans
Fish
Non-corralling feeding behavior
Clarity on Limitations of Penguin Vision
SECTION V PENGUINS IN A FICKLE ENVIRONMENT
CHAPTER 12. TURNING THE TABLES FISH-BIRDS ON THE MENU
Basic Law of the Sea: Big Fish Eat Little Fish
Seals as Predators
Seals hunting behavior
Penguins avoiding seals
Fur Seals as Predators
Sea Lions as Predators
Killer Whales as Predators
Sharks as Predators
CHAPTER 13. PENGUINS ADJUSTING TO A CHANGING OCEAN
Penguins Have Always Been Challenged by a Changing Ocean
Prehistoric response to a changing ocean
Possible prehistoric changes to penguins food web
The Anthropocene: How will Penguins Cope, Now Also Dealing with Humans?
Response to long-term climate change
Response to short-term ocean climate variation
Response to marine pollution
CHAPTER 14. NOT FORGETTING
The Social Side - Behavior and Communication at Sea
Penguin flock fusion/cohesion
Penguin flock fission/fragmentation
Navigation
Long range
Medium to short-range
Understanding the Daily Wash
Air flux in diving penguins, an aspect of washing
Are Auks Really Northern Penguins?
Research Tags the Flip Side for Evolutionarily-Honed Fish-Birds
SECTION VI
CHAPTER 15. PENGUINS WHY THE HYPE?
Sources of Hype
Us
Many people
Researchers
The Transition
The fascination of species
Beyond the transition
Role in Ecosystems
Biomimicry
Our Last Word.