The Bowhead Whale: Balaena Mysticetus: Biology and Human Interactions

Front Cover
The Bowhead Whale
Copyright Page
Dedication
Contents
List of contributors
Preface
Acknowledgments
I. Basic biology
	1 Higher level phylogeny of baleen whales
		The phylogenetic branching history of the bowhead whale
		Challenges for estimation of divergence times in Mysticeti
		Conclusions
		Acknowledgments
		References
	2 Fossil record
		Introduction
		Balaenid origins and the Miocene gap
		Late Neogene diversification and the emergence of bowheads
		Acknowledgements
		References
	3 The stocks of bowheads
		Introduction
		Genetics of bowhead whales
			Mitochondrial DNA
			Microsatellites
			Whole genomes and single nucleotide polymorphisms
		Bowhead stocks
			Bering-Chukchi-Beaufort Seas stock
			East Greenland-Svalbard-Barents Sea stock
			Okhotsk Sea stock
			East Canada-West Greenland stock
		Historical demography and evolutionary history
		Acknowledgments
		References
	4 Distribution and behavior of Bering-Chukchi-Beaufort bowhead whales as inferred by telemetry
		Introduction
		Description of the tagged sample of Bering-Chukchi-Beaufort bowhead whales
		Seasonal distribution of tagged Bering-Chukchi-Beaufort bowhead whales
			Spring migration
			Summer range
			Autumn migration
			Winter range
		Dive behavior
		Proximate mechanisms driving distribution
			Cape Bathurst
			Tuktoyaktuk shelf
			Eastern Alaskan Beaufort Sea
			Point Barrow
			Chukotka coast
			Anadyr Strait and Gulf of Anadyr
			Role of sea ice
		Recent changes in distribution
		Limitations of satellite telemetry
		Research needs
		Acknowledgments
		References
	5 Distribution, migrations, and ecology of the Atlantic and the Okhotsk Sea Populations
		Introduction
		The East Canada-West Greenland population
		The East Greenland-Svalbard-Barents Sea population
		The Okhotsk Sea population
		Diving activity
		Comparison of diet among stocks
		Discussion
		Acknowledgments
		References
	6 Abundance
		Introduction
		The Bering-Chukchi-Beaufort Seas stock
		The East Canada-West Greenland stock
		The Okhotsk Sea stock
		The East Greenland-Svalbard-Barents Sea stock
		References
	7 Life history, growth, and form
		Introduction
		Growth and form
			General description
			Bowhead growth phases
			Age: birth to year 1 (neonate or aġvaaq)
			Age: 1–2 years (ingutuq)
			Age: ~2–6 years (qairiliq)
			Age: ~6–25 years (sexually immature subadults)
			Age: ~26 to 200+ years (sexually mature adults)
		Specific morphological characteristics
			Baleen
			Blubber
			Body length/girth relationships
			Pectoral fin
			Flukes
		Body mass
			Blue whale-sized bowheads?
		Longevity
		Morphometric regressions
		Life history
			Life history theory
			Reproduction
			Calving areas
			Migration
			Ice navigation
			Evolution of exceptional longevity and delayed sexual maturity
		Acknowledgments
		References
	8 Prenatal development
		Introduction
		Description and comparisons
			Embryos of 8.7–9.1cm
			Fetuses of 12.8–16.6cm
			Fetuses of 27.4–40.3cm
			Fetuses of 84–175cm
			Full-term fetus
		Discussion
		References
	9 Anatomy of skull and mandible
		Introduction
		The bones of the skull
		Mandible and hyoid apparatus
		Bones of the cranial vault
		Boney orbit and position of the eye
		The boney nasal opening and nasal cavity
		Skull growth
		References
	10 Postcranial skeleton and musculature
		Introduction
		Muscles of the head and neck
		Axial skeleton and musculature
		Ribs and sternum
		Forelimb muscles and skeleton
		Hindlimb
		Conclusions
		References
	11 Hematology, serum, and urine composition
		Introduction
		Hematology
		Serum electrolytes
		Serum chemistry
		Serum chemistry and feeding status
		Immunoglobulins
		Urine analysis
		Urine electrolytes and aminograms
		Conclusions
		Acknowledgments
		References
	12 Anatomy and physiology of the gastrointestinal system
		Introduction
		Wax ester digestion
		Setting the stage—evolutionary and chemical considerations
		Anatomy of the stomach
		Gut passage times and fecal isotopes
		Proximate composition of digesta and fatty acid abundance
		Digestive efficiency
		Future considerations
		Acknowledgments
		References
	13 Female and male reproduction
		Introduction
		Reproductive tract morphology
			External genitalia
			Female reproductive tract
			Ovary
			Uterine tube
			Uterus and cervix
			Vagina, vulva, and clitoris
			Mammary gland
			Male reproductive tract
			Testis and epididymis
			Ductus deferens, ampulla, and accessory glands
			Penis
		Functional parameters of the bowhead whale reproductive cycle
			Female body length and age at sexual maturity
			Ovulation
			Conception
			Gestation and parturition
			Pregnancy rate and calving interval
			Lactation
			Male total body length and age at sexual maturity
		Acknowledgements
		References
	14 Anatomy and function of feeding
		Introduction: baleen and oral morphology
		Feeding behavior and functional ecology
		Acknowledgments
		References
	15 Cardiovascular and pulmonary systems
		Introduction
		Cardiovascular and respiratory function
		Aging and cardiovascular function
			Structural adaptations
			Molecular adaptations
		Summary
		Acknowledgments
		References
	16 Thermoregulation and energetics
		Introduction
		Body temperature
			Heat loss measurements
			Regional heterothermy
			Thick blubber and thermoregulation
		Basal and resting metabolic rates
			Resting metabolic rates and Kleiber law
			Bowhead resting metabolic via heat loss models
			Metabolic rates estimated by lung volume and respiration
			Summary of metabolic rate estimates
			Energetic models for East Canada-West Greenland bowheads
		Anatomical specializations
		Other energetic considerations
			Relative organ size, muscle, and blubber proportions
			Duration of feeding season and winter feeding
			Comparisons with North Atlantic right whales
			Energetics of locomotion
			Growth and reproduction costs
			Cost of disturbance
			Fasting endurance of bowhead whales
		Summary
			Suggestions for further research
		Acknowledgments
		References
	17 Brain
		Introduction
		Description and comparisons
			Shape and size of the bowhead whale brain
			Overview and surface morphology of the bowhead whale brain
			Gyrencephalic index and corpus callosum size
			Olfaction
			Hippocampus
			Cerebral cortex cytoarchitecture
		Discussion
		Acknowledgments
		References
	18 Sensory systems
		Introduction
		Olfaction and gustation
		Vision and magnetosense
		Audition
		Balance
		Mechanosense
		Discussion
		References
	19 Endocrinology and blubber physiology
		Introduction
		Blubber structure and physiology
			Leptin: the blubber hormone
		Circulating hormones
			Reproductive hormones
			Thyroid hormones
			Adrenal glucocorticoid hormones
		Quantifying hormones in alternative sample types
			Blubber steroid hormones
			Fecal hormones
				Assessing reproductive status
				Assessing stress responses
			Baleen hormones
			Earplug hormones
			Hormones in exhaled breath
		Discussion
		Acknowledgments
		References
	20 Molecular insights into anatomy and physiology
		Introduction
			Adapting the genome to extend life span
			Expanding the thermal limits for mammalian life
			Streamlining the sensory system for dim light and salty seas
			Maintaining healthy skin in icy waters
			Growing teeth and ever-growing baleen
			Unusual growth of the skeleton
			Future work
		Acknowledgments
		References
	21 Age estimation
		Introduction
		Age estimation using baleen
			Cycles in stable carbon isotope values
			Baleen length method
		Age estimation using growth layers in the tympanic bone
		Age estimation based on aspartic acid racemization
		Age estimation based on ovarian corpora
		Age estimation based on whaling artifacts
		Age estimation based on photo-recapture
			Estimating age using morphometric data
		Comparison of age methods
			Age 0 (birth)–20
			Age 20–30
			Age 30–150+
		References
	22 Acoustic behavior
		Introduction
		Bowhead whale sounds
		Sound production
		Bowhead whale calls
			Call sequences
			Counter-calling
		Bowhead whale gunshots
		Source level, calling depth, and detection distance
		Bowhead whale song
		Acoustic ecology of the bowhead whale
		References
	23 Natural and potentially disturbed behavior of bowhead whales
		Introduction
		Undisturbed activities
			Feeding behavior
			Social and sexual behavior
			Mother/calf interactions
			Play
			Predator avoidance/responses
			Surfacing, diving, respirations
		Potential and known disturbance reactions
			Traveling whales
				Traveling whales near seismic operations
				Traveling whales near drilling operations
				Traveling whales near ice management and anchor handling
			Feeding whales
				Feeding whales near seismic operations
				Feeding whales near drilling operation
			Whales during social and sexual activities
		Mother–calf reactions
			Summary
		Conclusion, syntheses, and knowledge needed
		Acknowledgments
		References
	24 Ecological variation in the western Beaufort Sea
		Introduction
		Aerial Surveys field methods
		Bowhead whale seasonal distribution in the western Beaufort Sea
		Mechanisms driving interannual variability in whale distribution, density, and seasonality
			Case study I: Nearshore feeding aggregations
			Case study II: Krill trap
			Case study III: Summer feeding aggregations in Harrison Bay
			Case study IV: Bowhead whales and sea ice
		Conclusions
		Acknowledgments
		References
II. The bowhead ecosystem
	25 Physical Oceanography
		Introduction
		Bering–Chukchi–Beaufort Seas
			Bering Sea and Shelf
			Chukchi Sea
			Beaufort Sea Shelf
		Okhotsk Sea
		Nordic Seas
		East Canada–West Greenland
		Some physical processes that aggregate bowhead prey
		Acknowledgement
		References
	26 Biological environment
		Introduction
		General characteristics
		Biogeography of bowhead habitats
			The Bering, Chukchi, and Beaufort Seas
			Sea of Okhotsk
			East Greenland–Svalbard–Barents Seas
			East Canada–West Greenland
		Formation of feeding hotspots
		Climatically driven environmental changes
		Acknowledgements
		References
	27 Bowhead whale ecology in changing high-latitude ecosystems
		Introduction
		Bowhead whale ecology in regional ecosystems
			Bering–Chukchi–Beaufort stock
			East Canada–West Greenland stock
			East Greenland–Svalbard–Barents Sea stock
			Okhotsk Sea stock
		Bowhead whale status and resilience
		References
	28 Diet and prey
		Introduction
		Diet research methods
			Body measurements (morphometrics)
			Fatty acid analyses
			Stable isotope analyses
			Blood
			Feces
			Stomach contents
		Diet and feeding in four bowhead stocks
			Okhotsk Sea stock
			East Greenland–Svalbard–Barents Sea stock
			East Canada–West Greenland stock
			Bering–Chukchi–Beaufort Seas stock
		Description of BCB stock seasonal feeding by region
			Northern Bering Sea—fall
			Northern Bering Sea—winter
			Northern Bering Sea—spring
			Chukchi Sea—spring
			Beaufort Sea—spring
			Beaufort Sea—summer
			Beaufort Sea—fall
			Chukchi Sea—fall
		Future concerns
			Bering–Chukchi–Beaufort bowhead specific concerns
		Conclusions
		Acknowledgments
		References
	29 Predators and impacts of predation
		Introduction
		Evidence of predation
			Attack accounts and carcass data
			Scars
		Effects of predation
			Direct mortality
			Risk effects
		Predation and Arctic warming
		Acknowledgments
		References
	30 Diseases and parasites
		Introduction
		Infectious diseases
			Viruses and bacteria
			Parasites and commensals
				Protozoa
				Helminths: cestodes
				Helminths: trematodes
				Acanthocephala
				Nematodes
				Amphipoda
				Diatoms
		Noninfectious diseases
			Neoplasia
			Integumentary system
			Musculoskeletal system
			Cardiovascular and respiratory system
			Digestive system
			Urinary system
			Reproductive system
			Immune and endocrine system
			Special senses
		Conclusions
		Acknowledgments
		References
III. Interactions with humans
	31 Whale hunting in Indigenous Arctic cultures
		Introduction
		Bowhead whaling in the scholarly literature
		Bowhead whaling as lived experience
			Contributors
			Aghveq angyiiquq: the gift of the whale on St. Lawrence Island, Alaska
			An Aġviq foundation
			The first bowhead whale after a 20 year ban
			Arvangniarniq: bowhead whale hunt in Clyde River, Nunavut, August 2014
			Bowhead whaling in Greenland
		Discussion
		Acknowledgments
		References
	32 Current indigenous whaling
		Introduction
		Data
		East Greenland, Svalbard, Barents Seas and Okhotsk Sea stocks
		East Canada–West Greenland stock
			Greenland
			Canada
		Bering–Chukchi–Beaufort Seas stock
			Canada
			Chukotka
			Alaska
				Numbers and timing of whales harvested
				Efficiency of the hunt
				Size of harvested whales
				Sex ratios
				Sharing of the harvest
			Sustainability of the hunt
		Summary and conclusions
		Acknowledgments
		References
	33 Commercial whaling
		Introduction
		The East Canada–West Greenland stock
		The East Greenland–Svalbard–Barents Sea stock
		The Bering–Chukchi–Beaufort stock
		The Okhotsk Sea stock
		Effect on indigenous people
		Discussion
			Biological implications
		References
	34 Indigenous knowledge in research and management
		Introduction
		Modes of engaging Indigenous knowledge concerning bowhead whales
			Informal use
			Formal documentation
			Intentional application to research
			Intentional application to management
		Examples of using Indigenous knowledge and scientific knowledge together
			The bowhead whale census at Utqiaġvik, Alaska
			Estimating bowhead abundance in the eastern Canadian Arctic
			Sense of smell
			Buoyancy
			Life span
			Molting
			Satellite telemetry and complementary understanding
			Indigenous scholarship
		Discussion
		Acknowledgments
		References
	35 Effects of noise
		Introduction
		Sources of noise in bowhead whale habitats
		Ambient wind-driven noise
		Continuous industrial sounds
			Sounds from an artificial oil-production island
			Vessels and other tonal sources
		Sounds from air guns
		Summary of short-term acoustic responses to fluctuations in noise
		Potential long-term impacts and conclusions
		References
	36 Fishing gear entanglement and vessel collisions
		Introduction
		Review of fishing gear entanglement by stock
			Okhotsk Sea stock
			East Greenland, Svalbard, Barents Sea stock
			East Canada–West Greenland stock
			Bering–Chukchi–Beaufort Seas stock
		Vessel strike injuries
		Discussion and conclusions
		Acknowledgements
		References
	37 Contaminants
		Introduction
		Petroleum-related contaminants
		Essential and nonessential elements
		Persistent organic pollutants
		Conclusions
		Acknowledgement
		References
	38 Conservation and management
		Introduction
		Regulations
		East Greenland-Svalbard-Barents Sea and Okhotsk Sea stocks
		East Canada-West Greenland stock
			Greenland
			Canada
		Bering-Chukchi-Beaufort bowheads
			Traditional management
			Alaska
			Chukotka
			IWC management
			Oil and gas
		Climate change
		Sustainability of the hunt
		Monitoring
		Summary and conclusions
		Acknowledgments
		References
	39 Past, present, and future
		Bowhead whales and humans
		Epilog
		References
Index
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