The Aquatic World of Penguins : Biology of Fish-Birds

Penguin Transitions
Preface
Acknowledgments
Contents
Part I: In the Beginning
Chapter 1: Wading In: Introduction to Fish-Birds
An Impressive Number of Penguin Species: Evolution of Unique Capabilities
Penguin Species Radiation and the Ontogeny of their Watery World
Penguin Evolution: Body Size and Climate
Penguin Evolution: Radiation into Vacant Niches
Conclusions
References
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: A Necessary, Brief Respite from the Sea
References
Part II: Penguin Marine Haunts and Food Habits
Chapter 3: Fish-Birds at Home in Their Oceanic Habitats
Oceanographic Fronts and Water Masses Are Important to Penguins: General Discussion
Penguins Require High-Productivity Water Masses
Seasonal At-Sea Movement of Penguins: Is It Migration or Dispersion?
Large-Scale Edge: The At-Sea Distributions of Penguins and Oceanographic Boundaries
The Southern Boundary of the Antarctic Circumpolar Current and the Large-Scale Sea–Ice Edge
The Antarctic Polar Front
The Subantarctic and Subtropical Fronts
Eastern Boundary Currents
Meso- and Small-Scale Factors: Regional and Local Ocean Processes that Facilitate the At-Sea Foraging of Penguins
Island Wakes
Headland Wakes, Eddies, and Upwelling Plumes
River Outflow Plumes
Shelves and Banks
Submarine Canyons
Shelf-Break Fronts
Marginal Ice Zones
Thermoclines/Haloclines
References
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
Penguin Diet Composition: A Comparison
Polar/Subpolar, Mesopelagic Foraging Penguins
Emperor Penguins
King Penguins
Subpolar, Full Water Column/Demersal/Benthic, Continental-Shelf Foraging Penguins
Yellow-Eyed Penguins
Gentoo Penguins
Temperate, Upper Water Column, Continental-/Insular-Shelf Foraging Penguins
Spheniscus Penguins
Australian and New Zealand Little Penguins
Polar, Upper Water Column, Continental-Shelf/Continental-Slope Foraging Penguins
Subpolar, Upper Water Column, Continental-Slope/Pelagic Foraging Penguins
Polar, Upper Water Column, Continental-Slope/Pelagic, Foraging Penguins
Chinstrap Penguins
Macaroni Penguins
References
Chapter 5: Ecological Consequences of Diet Composition
Intraspecific Competition Among Penguins
Foraging Range Is Key: General Considerations
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
References
Part III: The Hardware of a Fish-Bird
Chapter 6: The Slippery Shape, Hot Air, and the Powerhouse: How Fish-Birds Swim
Water: A Hard Taskmaster
The Four Forces Relevant to Penguins
Vertical Forces: Weight and Upthrust
Buoyancy: How Much Air Do Penguins Hold?
Buoyancy and Bergmann’s Rule Revisited
Horizontal Forces: Drag
The Interplay Between Upthrust and Drag in Gliding Penguins
The Drag Devil Lies 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, Cost of Transport, and Beyond
References
Chapter 7: Hot Penguins: Cold Water
Resting or 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”
References
Chapter 8: Fish-Birds: The Inside Story
Surface Issues: Uptake of Oxygen
Oxygen Management Underwater
Role of the Air Spaces
Gas Exchange to Body Tissues
The Aerobic Dive Limit (ADL) 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
References
Part IV: The Software of Fish-Birds
Chapter 9: Embracing the Depths: The Fish-Bird Dive
Submergence
The Time Underwater: Basic Dive Descriptors
Dive Profiles
Dive Distance–Depth Profiles
Dive Aspect Ratios
Horizontal Dive Directionality/Tortuosity
Basic Dive Types
T-Dives (for Traveling)
V-Dives (Prey Assessment)
P (Parabolic)-Dives (Prospecting with No Prey Capture)
PO-Dives (Parabolic Dives with a Circular Trajectory)
U-Dives (Depth-Directed Prospecting)
W-Dives/Up-Dives (U-Dives with Prey Pursuit)
Depth Duration Effects Over Multiple Dives
References
Chapter 10: Decisions, Decisions, and More 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 Foraging Strategies
References
Chapter 11: The Final Seconds: How Fish-Birds Catch Prey
Prey Acquisition: A Departure from the Norm
Performance Metrics for Prey Capture in Penguins
Capture of Solitary Prey
Changing Buoyancy with Depth Affects Prey Capture Strategies
Prey Pursuit Against Interfaces
Capture of Aggregating/Schooling Prey
Crustaceans
Fish
Non-Corralling Feeding Behavior
Clarity on Limitations of Penguin Underwater Vision
References
Part 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 Penguins
Penguins Avoiding Seals
Fur Seals as Predators
Sea Lions as Predators
Killer Whales as Predators
Large Fish as Predators
References
Chapter 13: Penguins Coping with 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
References
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
References
Part VI: Our Final Thoughts
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
References
Appendices
Appendix Table A1
Appendix Table A2
Appendix Table A3
Appendix Table A4
Glossary
References
References for Appendix Table A1
References for Appendix Table A3
References for Appendix Table A4