The Hawaiian Spinner Dolphin

Twenty years in the making by a distinguished dolphin expert and his associates, The Hawaiian Spinner Dolphin is the first comprehensive scientific natural history of a dolphin species ever written. From their research camp at Kealakeakua Bay in Hawaii, these scientists followed a population of wild spinner dolphins by radiotracking their movements and, with the use of a windowed underwater vessel, observing the details of their underwater social life.

The authors begin with a description of the spinner dolphin species, its morphology and systematics, and then examine the ocean environment, the organization of dolphin populations, and the way this school-based society of mammals uses shorelines for rest and instruction of the young. The dolphins' reproductive cycle, their vision, vocalization, hearing, breathing, and feeding, and the integration of the school are carefully analyzed. The authors conclude with a comprehensive evolutionary analysis of this marine cultural system, with its behavioral flexibility and high levels of cooperation.

This absorbing book is the richest source available of new scientific insights about the lives of wild dophins and how their societies evolved at sea.

Kenneth S. Norris is Emeritus Professor of Biology at the University of California, Santa Cruz, co-editor of Dolphin Societies (California, 1990), and author of the award-winning Dolphin Days (1991). Bernd Würsig is Professor of Marine Mammalogy at Texas A & M University, where Melany Würsig is a research associate. Randall S. Wells is a conservation biologist with the Chicago Zoological Society. Shannon M. Brownlee is Senior Editor for Science at U.S. News and World Report. Christine Johnson teaches in the Department of Cognition at the University of California, San Diego. Jody Solow is a doctoral candidate in geography at the University of Cambridge. Jenny Wardrip is a freelance illustrator and a lecturer at the University of California, Santa Cruz.

FIGURES
INTRODUCTION
Kenneth S. Norris

CHAPTER 1 THE SPINNER DOLPHIN 
Kenneth S. Norris, Bernd Wiirsig, and Randall S. Wells
WORLDWIDE DISTRIBUTION 
HISTORY OF SPINNER DOLPHIN STUDY 
ENVIRONMENT OF THE SPINNER DOLPHIN IN THE EASTERN TROPICAL PACIFIC 
OCEANIC FAUNAL RADIATION 
ADAPTATION TO LOCAL RESOURCES 
VARIATION 
ECOTYPES 
POPULATION PATTERNS 
LOCAL OCEANOGRAPHY 
PATTERN 
PHYSICAL VARIATION 
BEHAVIORAL STUDIES 

CHAPTER 2 THE ISLAND HABITAT 
Randall S. Wells and Kenneth S. Norris
THE ISLAND OF HAWAII 
OCEANOGRAPHIC CORRELATES 
STRUCTURAL HYDROGRAPHY OF KEALAKE'AKUA BAY 
BOTTOM TOPOGRAPHY AND DOLPHIN ABUNDANCE 
AERIAL SURVEYS: LOCAL AND SEASONAL PATTERNS 
SIGHTING EFFICIENCY 
IS THERE AN ISLAND OF HAWAII SPINNER POPULATION? 
Circumisland Movements 
Rest Coves 
The Spinner Population: Open or Closed? 

CHAPTER 3 OBSERVING DOLPHINS UNDERWATER 
Kenneth S. Norris and Randall S. Wells
METHODS OF OBSERVING DOLPHINS 
UNDERWATER VIEWING VEHICLES 
APPROACHING DOLPHINS 
LISTENING 
THE MOBILE OBSERVATION CHAMBER 
THE MAKA ALA 
THE SMYG TITTAR'N 

CHAPTER 4 A SPINNER DOLPHIN'S DAY 
Bernd Wursig, Randall S. Wells, Kenneth S. Norris, and Melany Wursig
METHODS OF OBSERVATION 
AN OUTLINE OF THE DAILY CYCLE 
BAY RESIDENCE PATTERNS 
DESCENT INTO REST 
REST 
AWAKENING 
ZIG-ZAG SWIMMING 
SPREAD FORMATION 
NIGHTTIME PATTERNS 
THE QUESTION OF SCHOOL FLUIDITY 

CHAPTER 5 AERIAL BEHAVIOR 
Kenneth S. Norris, Bernd Wursig, and Randall S. Wells
CATEGORIES OF AERIAL BEHAVIOR 
NOSE-OUTS AND FLUKE-OUTS 
Nose-Outs 
Fluke-Outs 
SLAPS 
Tail Slaps 
Head Slaps and Back Slaps 
LEAPS 
Arcuate Leaps 
Salmon Leaps 
Tail-Over-Head Leaps 
SPINS 
PATTERNS OF OCCURRENCE 
FUNCTIONS OF AERIAL PATTERNS 
Remora Dislodgement 
Sound and Bubble Production 
OTHER CORRELATES OF AERIAL BEHAVIOR 
CONCLUSIONS 

CHAPTER 6 POPULATION STRUCTURE 
Bernd Wiirsig, Randall S. Wells, Melany Wiirsig, and Kenneth S. Norris
SCARS AND MARKS ANALYSIS 
OCCUPANCY OF REST COVES 
INDIVIDUAL IDENTIFICATION 
ISLAND USE PATTERNS 
ASSOCIATIONS 
MOVEMENTS 
ACCESSION RATE 
POPULATION PATTERNS AND SEGREGATION 
A POPULATION MODEL 
COMPARISONS TO OTHER POPULATIONS 
ABUNDANCE
PERSPECTIVES

CHAPTER 7 THE VISUAL DOMAIN
Kenneth S. Norris, Randall S. Wells, and Christine M. Johnson
DIURNAL LIGHT CYCLE 
LIGHT SCATTERING 
FLICKER 
EFFECTS OF LIGHT INTENSITY ON SKIN 
REFRACTION 
THE DOLPHIN VISUAL SYSTEM 
VISUAL PIGMENTS 
VISUAL SIGNS, SIGNALS, AND VISUAL FIELDS 
Visual Fields 
Whole-Body Signals 
The Pectoral Fin and Associated Patterns 
Other Body Patterns 
The Rostrum 
Locomotory Signs 
Sexual Pattern 

CHAPTER 8 THE ACOUSTIC DOMAIN 
Shannon M. Bnrwnlee and Kenneth S. Norris
THE ACOUSTIC ENVIRONMENT
Source Levels and Beam Characteristics
Distortion of the Sound Path, Transmission Losses, and Bubble Eflects 
Target Strength 
Ocean Noise 
Detection Thresholds 
SOUND EMISSION PATTERNS 
Data Gathering 
Classes of Sounds 
Sounds and Behavior 
INTERPRETATIONS OF ACOUSTIC SIGNALS 
Whistles: The Phatic System 
Classification of Message Types 
Signature Whistles 
Social Facilitation 
Burst-Pulse Signals 

CHAPTER 9 PATTERNS OF REPRODUCTION 
Randall S. Wells and Kenneth S. Norris
PATTERNS OF CETACEAN REPRODUCTION 
Gestation and Development 
The Male Cycle 
The Mating System 
Mate Selection 
The Female Cycle 
Reproductive Seasonality 
DETERMINANTS OF REPRODUCTIVE SYSTEMS 
HORMONAL AND BEHAVIORAL CORRELATES OF REPRODUCTION 
HORMONAL CYCLES OF HAWAIIAN SPINNERS 
The Male Spinner 
The Female Spinner 
Behavioral Correlates 

CHAPTER 10 LOCOMOTION 
Kenneth S. Norris and Christine M. Johnson
CROSSOVER SPEED 
PROPULSIVE PATTERN 
AXIS OF LOCOMOTION 

CHAPTER 11 BREATHING AT SEA 
Kenneth S. Norris and Christine M. Johnson
BREATHING WHILE SWIMMING 
BREATH HOLDING 
SEPARATING FOOD AND AIR 
PHONATION 
PRESSURE EFFECTS 
LUNG COLLAPSE 
BREATHING AT SEA 
Exhalation 
Inhalation 
Locating a Breathing Place 

CHAPTER 12 FOOD AND FEEDING 
Bernd Wursig, Randall S. Wells, and Kenneth S. Norris
MECHANICS OF FOOD CAPTURE AND HANDLING 
The Cage Jaw 
The Piston Tongue 
Ingestion 
OWNERSHIP OF PREY 
DIET AND FEEDING DEPTH 
RESOURCE PARTITIONING 
THE TUNA-DOLPHIN BOND 
THE FOOD OF HAWAIIAN SPINNER DOLPHINS 
DIVING PATTERNS 

CHAPTER 13 SCHOOLS AND SCHOOLING 
Kenneth S. Norris and Christine M. Johnson
FUNCTIONING OF SCHOOLS 
SCHOOL FORMATIONS 
Echelon Formation 
Sensory Windows 
SENSORY INTEGRATION SYSTEM 
Sensory Summation 
Sensory Involvement 
SCHOOL PATTERNS 
THE QUESTION OF SCHOOL LEADERSHIP 
RELATIONSHIP OF VISION TO DOLPHIN SCHOOLING 
SCHOOL TIGHTENING AND SCHOOLING DISTANCE 
SCHOOL BEHAVIOR IN RELATION TO ATTACK 
ANTICIPATION 
CONFUSION 

CHAPTER 14 SOCIAL BEHAVIOR 
Christine M. Johnson and Kenneth S. Norris
SCHOOL WARINESS AND HABITUATION 
EPISODIC OR BOUT BEHAVIOR 
Aerial Behavior Bouts 
Caressing Bouts 
Formation Swimming 
Echolocation Bouts 
DO DOLPHINS ECHOLOCATE ONE ANOTHER? 
PARTNER SHARING AND EXCHANGE 
MALE COALITIONS 
MATING 
BIRTH, NURTURE, AND JUVENILE GROUPS 
CALF AND JUVENILE BEHAVIOR 
PLAY 
AGGRESSION AND DEFENSE 
Open Mouth Contact 
Threat Posture 
Aggressive Encounters 
Convergence Between Shark and Dolphin
Behavior/Structure 
USES OF INTENSE SOUND 

CHAPTER 15 PREDATORS, PARASITES, AND MULTISPECIES AGGREGATIONS 
Kenneth S. Norris
PREDATION IN SHALLOW VERSUS OPEN SEAS 
Predation in the Open Sea 
Evidence from Hawaiian Spinners 
MULTISPECIES AGGREGATIONS 
DOLPHIN PREDATORS 
THE COOKIE CUTTER SHARK 
PARASITES 

CHAPTER 16 COMPARATIVE VIEW OF CETACEAN
SOCIAL ECOLOGY, CULTURE, AND EVOLUTION 
Kenneth S. Norris
INTRODUCTION 
Defining Some Terms 
Dolphin Variation 
Rhythmic or Oscillatory Behavior 
SELF-COMPENSATORY SYSTEMS 
ARE SOCIOECOLOGIES EVOLUTIONARY ENTITIES? 
THE ORIGIN OF CULTURES 
COMPARISON OF MARINE AND TERRESTRIAL MAMMALS 
ADAPTIVE RADIATION OF WARM WATER CETACEANS 
SPINNER DOLPHIN SOCIETY 
THREE-DIMENSIONAL SOCIETIES 
ARTHROPOD SOCIETIES 
TOWARD A MODEL FOR SOCIOECOLOGICAL AND CULTURAL EVOLUTION 
SYSTEMIC NATURE OF THE PHENOTYPE AND THE ORIGIN OF COOPERATION 
DEFINING THE PHENOTYPE 
The Extended Phenotype 
THE SOCIOECOLOGY: AN EXAMPLE OF AN EPIGENETIC SYSTEM 
OSCILLATION OF BEHAVIOR STATES 
Genetic Control of Oscillation 
The Coexistence of Organizational Levels 
ADAPTIVE RADIATION 
THE CULTURAL SYSTEM 
CULTURAL ASPECTS OF SEXUAL ROLES 
THE FITNESS OF CULTURES 
Play and Juvenility 
Adolescence in Primates and Dolphins 
COST ACCOUNTING 
THE EMERGING SYSTEMIC VIEW OF EVOLUTION 
CONCLUDING THOUGHTS ABOUT SPINNER DOLPHINS 

OVERVIEW 
APPENDIX A: THOSE WHO HELPED 
APPENDIX B: MATERIALS AND METHODS 
LITERATURE CITED 
INDEX