Information Resources on Marine Mammals


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Cetaceans–Diving / Locomotion / Energetics / Hydrodynamics



Acevedo Gutierrez, A., D.A. Croll, and B.R. Tershy (2002). High feeding costs limit dive time in the largest whales. Journal of Experimental Biology 205(12): 1747-1753. ISSN: 0022-0949.
NAL Call Number: 442.8 B77
Descriptors: Balaenoptera musculus, Balaenoptera physalus, energy expenditure, foraging, diving, north Pacific, Mexico, Baja California, USA, California, lunge feeding, dive duration, energetic costs.

Akamatsu, T., D. Wang, K. Wang, Z. Wei, Q. Zhao, and Y. Naito (2002). Diving behaviour of freshwater finless porpoises (Neophocaena phocaenoides) in an oxbow of the Yangtze River, China. ICES Journal of Marine Science 59(2): 438-443. ISSN: 1054-3139.
Descriptors: behavior, freshwater ecology, bottom time, dive depth, diving behavior, horizontal travel distance, swimming speed, finless porpoise, Yangtze River, China.

Alexander, R.M. (2004). Hitching a lift hydrodynamically--in swimming, flying and cycling. Journal of Biology 3(2): 7. ISSN: 1475-4924.
Abstract: Swimming animals set the water around them moving, and flying animals generate air movements. Other animals traveling with them can save energy by exploiting these movements of the fluid medium; similarly, a cyclist can save energy by riding close behind another. A new study of dolphin mothers and calves exemplifies the advantages of moving in concert.
Descriptors: behavior, animal physiology, bicycling physiology, dolphins physiology, swimming physiology, mothers.

Alexander, R.M. (2002). Tendon elasticity and muscle function. Comparative Biochemistry and Physiology. A, Molecular and Integrative Physiology 133A(4): 1001-1011. ISSN: 1095-6433.
NAL Call Number: QP1.C6
Descriptors: vertebrata, literature review, biomechanics, skeletal musculature, tendons, tendon elasticity and muscel function relationships, locomotion, tendon elasticity and muscle function relationships, review.

Amano, M. and M. Yoshioka (2003). Sperm whale diving behavior monitored using a suction-cup-attached TDR tag. Marine Ecology Progress Series 258: 291-295. ISSN: 0171-8630.
NAL Call Number: QH541.5.S3M32
Descriptors: Physeter macrocephalus, activity recording, telemetry techniques, foraging, aquatic diving, north Pacific, Japan, Kumano coast, diving behavior monitoring using suction cup attached tdr tag.

Amano, M., M. Yoshioka, and K. Mori (2003). Study on diving behavior of sperm whales using suction cup attached TDR tag: an overview. Otsuchi Marine Science 28: 1-5. ISSN: 1344-8420.
Descriptors: Physeter macrocephalus, behavioral techniques, west Pacific, Japan and Ogasawara Islands, diving behavior analysis using suction cap attached time depth recorder tag, overview.

Baird, R.W., J.F. Borsani, M.B. Hanson, and P.L. Tyack (2002). Diving and night-time behavior of long-finned pilot whales in the Ligurian Sea. Marine Ecology Progress Series 237: 301-305. ISSN: 0171-8630.
NAL Call Number: QH541.5.S3M32
Descriptors: behavior, dive depths, diving, foraging, night time behavior, social behaviors, pilot whales, Ligurian Sea, deep dives, radio tags, foraging.

Baumgartner, M.F. and B.R. Mate (2003). Summertime foraging ecology of North Atlantic right whales. Marine Ecology Progress Series 264: 123-135. ISSN: 0171-8630.
NAL Call Number: QH541.5.S3M32
Descriptors: Eubalaena glacialis, food availability, feeding rate, foraging, aquatic diving, ecology, salinity, temperature, north Atlantic, Canada, lower Bay of Fundy and southwestern Scotian Shelf, summer foraging ecology and influences.

Beasley, I. and T.A. Jefferson (2002). Surface and dive times of finless porpoises in Hong Kong's coastal waters. Raffles Bulletin of Zoology Supplement 10: 125-129.
Descriptors: behavior, finless porpoise, ecology, helicopter survey, land based survey, line transect method, coastal waters, dive times, surfacing behavior, trackline detection probability, Hong Kong.

Blake, R.W. (1983). Energetics of leaping in dolphins and other aquatic animals Killer whales, Orcinus orca, baleen whales, penguins. Journal of the Marine Biological Association of the United Kingdom 63(1): 61-70. ISSN: 0025-3154.
NAL Call Number: 442.9 M331
Descriptors: dolphins, leaping, energetics, killer whales, penguins, baleen whales, aquatic animals, Orcinus orca.

Blix, A.S. and L.P. Folkow (1995). Daily energy expenditure in free living minke whales. Acta Physiologica Scandinavica 153(1): 61-6. ISSN: 0001-6772.
NAL Call Number: QP1.A2
Abstract: Six minke whales (Balaenoptera acutorostrata) were instrumented with VHF-radio transmitters and four with sonic speed-depth transmitters off the west coast of northern Norway and Svalbard and followed within view for up to 24 h. During such periods their respiratory rate was continuously recorded and their energy expenditure estimated according to Folkow & Blix (1992) at different swimming speeds and types of activity. We found that cost of swimming is remarkably low in these large animals and that their estimated daily energy expenditure on average only amounts to 80 kJ kg-1 day-1.
Descriptors: energy metabolism physiology, respiration physiology, whales physiology, eating, kinetics, Norway, sleep, time factors.

Boutilier, R.G., J.Z. Reed, and M.A. Fedak (2001). Unsteady-state gas exchange and storage in diving marine mammals: the harbor porpoise and gray seal. American Journal of Physiology. Regulatory, Integrative and Comparative Physiology 281(2): R490-4. ISSN: 0363-6119.
Abstract: Breath-by-breath measurements of end-tidal O(2) and CO(2) concentrations in harbor porpoise reveal that the respiratory gas exchange ratio (R(R); CO(2) output/O(2) uptake) of the first lung ventilation in a breathing bout after a prolonged breath-hold is always well below the animal's metabolic respiratory quotient (RQ) of 0.85. Thus the longest apneic pauses are always followed by an initial breath having a very low R(R) (0.6-0.7), which thereafter increases with each subsequent breath to values in excess of 1.2. Although the O(2) stores of the body are fully readjusted after the first three to four breaths following a prolonged apneic pause, a further three to four ventilations are always needed, not to load more O(2) but to eliminate built-up levels of CO(2). The slower readjustment of CO(2) stores relates to their greater magnitude and to the fact that they must be mobilized from comparatively large and chemically complex HCO/CO(2) stores that are built up in the blood and tissues during the breath-hold. These data, and similar measurements on gray seals (12), indicate that it is the readjustment of metabolic RQ and not O(2) stores per se that governs the amount of time an animal must spend ventilating at the surface after a dive.
Descriptors: diving physiology, porpoises physiology, pulmonary gas exchange physiology, respiration, seals, earless physiology, oxygen metabolism, time factors.

Brix, O., S.G. Condo, A. Bardgard, B. Tavazzi, and B. Giardina (1990). Temperature modulation of oxygen transport in a diving mammal (Balaenoptera acutorostrata). Biochemical Journal 271(2): 509-13. ISSN: 0264-6021.
NAL Call Number: QP501.B64
Abstract: The functional properties of haemoglobin from the Lesser Rorqual whale (Balaenoptera acutorostrata) have been characterized as a function of the heterotropic effector concentrations and temperature. The results obtained suggest the existence of sophisticated modulation mechanisms based on the interplay of organic phosphates, carbon dioxide, lactate and temperature. These, together with the very small apparent heat of oxygenation (delta H) of oxygen binding, have been physiologically interpreted on the basis of the specific metabolic needs of this diving mammal.
Descriptors: diving, hemoglobins metabolism, oxygen blood, whales blood, 2,3 diphosphoglycerate, biological transport, carbon dioxide pharmacology, diphosphoglyceric acids pharmacology, hydrogen ion concentration, lactates pharmacology, lactic acid, temperature, thermodynamics.

Brix, O., M. Ekker, S.G. Condo, R. Scatena, M.E. Clementi, and B. Giardina (1990). Lactate does facilitate oxygen unloading from the hemoglobin of the whale, Balaenoptera acutorostrata, after diving. Arctic Medical Research 49(1): 39-42. ISSN: 0782-226X.
Abstract: The present study reports on a specific effect of lactate on the oxygen binding properties of the hemoglobin from the whale, Balaenoptera acutorostrata. In fact 0.1 mM lactate may increase the amount of oxygen unloaded to the tissues as much as 30%. Under these conditions the Bohr shift, of the magnitude of about -1, does not alter the oxygen affinity, but plays an important role in the isohydric transport of carbon dioxide.
Descriptors: Cetacea blood, diving, hemoglobins metabolism, lactates blood, oxygen blood, whales blood, lactic acid.

Cozzi, B., P. Bagnoli, F. Acocella, and M.L. Costantino (2005). Structure and biomechanical properties of the trachea of the striped dolphin Stenella coeruleoalba: evidence for evolutionary adaptations to diving. Anatomical Record 284A(1): 500-510. ISSN: 0003-276X.
Descriptors: Stenella coeruleoalba, biomechanics, biomechanical properties, trachea, structure and biomechanical properties, evidence for evolutionary adaptations to diving, evolutionary adaptation, aquatic diving.

Croll, D.A., A. Acevedo Gutierrez, B.R. Tershy, and J. Urban Ramirez (2001). The diving behavior of blue and fin whales: is dive duration shorter than expected based on oxygen stores? Comparative Biochemistry and Physiology. A, Molecular and Integrative Physiology 129(4): 797-809. ISSN: 1095-6433.
NAL Call Number: QP1.C6
Abstract: Many diving seabirds and marine mammals have been found to regularly exceed their theoretical aerobic dive limit (TADL). No animals have been found to dive for durations that are consistently shorter than their TADL. We attached time-depth recorders to 7 blue whales and 15 fin whales (family Balaenopteridae). The diving behavior of both species was similar, and we distinguished between foraging and traveling dives. Foraging dives in both species were deeper, longer in duration and distinguished by a series of vertical excursions where lunge feeding presumably occurred. Foraging blue whales lunged 2.4 (+/-1.13) times per dive, with a maximum of six times and average vertical excursion of 30.2 (+/-10.04) m. Foraging fin whales lunged 1.7 (+/-0.88) times per dive, with a maximum of eight times and average vertical excursion of 21.2 (+/-4.35) m. The maximum rate of ascent of lunges was higher than the maximum rate of descent in both species, indicating that feeding lunges occurred on ascent. Foraging dives were deeper and longer than non-feeding dives in both species. On average, blue whales dived to 140.0 (+/-46.01) m and 7.8 (+/-1.89) min when foraging, and 67.6 (+/-51.46) m and 4.9 (+/-2.53) min when not foraging. Fin whales dived to 97.9 (+/-32.59) m and 6.3 (+/-1.53) min when foraging and to 59.3 (+/-29.67) m and 4.2 (+/-1.67) min when not foraging. The longest dives recorded for both species, 14.7 min for blue whales and 16.9 min for fin whales, were considerably shorter than the TADL of 31.2 and 28.6 min, respectively. An allometric comparison of seven families diving to an average depth of 80-150 m showed a significant relationship between body mass and dive duration once Balaenopteridae whales, with a mean dive duration of 6.8 min, were excluded from the analysis. Thus, the short dive durations of blue whales and fin whales cannot be explained by the shallow distribution of their prey. We propose instead that short duration diving in large whales results from either: (1) dispersal behavior of prey; or (2) a high energetic cost of foraging.
Descriptors: diving physiology, feeding behavior physiology, oxygen metabolism, whales physiology, body mass index, statistics, time factors.

Curren, K., N. Bose, and J. Lien (1994). Swimming kinematics of a harbor porpoise (Phocoena phocoena) and an atlantic white-sided dolphin (Lagenorhynchus acutus). Marine Mammal Science 10(4): 485-492. ISSN: 0824-0469.
NAL Call Number: QL713.2.M372
Descriptors: behavior, mathematical biology, computational biology, morphology, systematics and taxonomy, behavior, comparative morphology, mathematical relationship.

Dearolf, J.L., W.A. McLellan, R.M. Dillaman, D. Frierson Jr., and D.A. Pabst (2000). Precocial development of axial locomotor muscle in bottlenose dolphins (Tursiops truncatus). Journal of Morphology 244(3): 203-15. ISSN: 0362-2525.
NAL Call Number: 444.8 J826
Descriptors: dolphins growth and development, muscle development, muscle, skeletal growth and development, newborn animals, body weight, dolphins anatomy and histology, immunohistochemistry, locomotion physiology, muscle, skeletal anatomy and histology.

Drouot, V., A. Gannier, and J.C. Goold (2004). Diving and feeding behaviour of sperm whales (Physeter macrocephalus) in the northwestern Mediterranean Sea. Aquatic Mammals 30(3): 419-426. ISSN: 0167-5427.
Abstract: Sperm whale (Physeter macrocephalus) is one of the common cetacean species inhabiting the western Mediterranean Sea. The aim of this study was to describe the dive cycle of sperm whales in this region and gain insight into their foraging activity. Dedicated summer field seasons were conducted from 2001-2003. Visual and acoustic measures were undertaken and their relationships analyzed. The measures included surface/dive periods, blow rate, distance travelled, timing of clicks & creak activity, and inter-pulse interval measurements of sperm whale clicks. The whales exhibited dive cycle parameters consistent with those measured in other parts of the world: approximately 45 min dive duration, 9 min surface period (i.e., inter-dive interval), with 5 blows/min, and 1.3 nmi horizontal displacement between dives. An average of 25 creaks per dive were registered. Whale body size appeared to be significantly related with both the number of creaks per dive and the dive time at which the first creak occurred, suggesting that larger whales may increase their prey intake and use deeper water layers than smaller whales. The timing of the first creak and the last click of the dive (around 6 min after fluke-up, and just before the surfacing, respectively) suggest a foraging depth of between 500 and 800 m, based on known descent and ascent rates.
Descriptors: Physeter macrocephalus, echolocation, creak and click phenology, foraging, aquatic diving, foraging relationships, activity patterns, dive cycle, vertical distribution, depth, foraging depth, Mediterranean Sea, France, provence, dive cycle and foraging relationships.

Edel, R.K. and H.E. Winn (1978). Observations on underwater locomotion and flipper movement of the humpback whale Megaptera novaeangliae. Marine Biology (Berlin) 48(3): 279-287. ISSN: 0025-3162.
NAL Call Number: QH91.A1M35
Descriptors: humpback whale, underwater locomotion, flipper movement, observations.
Language of Text: English summary.

Fish, F.E. (2000). Biomechanics and energetics in aquatic and semiaquatic mammals: platypus to whale. Physiological and Biochemical Zoology 73(6): 683-98. ISSN: 1522-2152.
NAL Call Number: QL1.P52
Abstract: A variety of mammalian lineages have secondarily invaded the water. To locomote and thermoregulate in the aqueous medium, mammals developed a range of morphological, physiological, and behavioral adaptations. A distinct difference in the suite of adaptations, which affects energetics, is apparent between semiaquatic and fully aquatic mammals. Semiaquatic mammals swim by paddling, which is inefficient compared to the use of oscillating hydrofoils of aquatic mammals. Semiaquatic mammals swim at the water surface and experience a greater resistive force augmented by wave drag than submerged aquatic mammals. A dense, nonwettable fur insulates semiaquatic mammals, whereas aquatic mammals use a layer of blubber. The fur, while providing insulation and positive buoyancy, incurs a high energy demand for maintenance and limits diving depth. Blubber contours the body to reduce drag, is an energy reserve, and suffers no loss in buoyancy with depth. Despite the high energetic costs of a semiaquatic existence, these animals represent modern analogs of evolutionary intermediates between ancestral terrestrial mammals and their fully aquatic descendants. It is these intermediate animals that indicate which potential selection factors and mechanical constraints may have directed the evolution of more derived aquatic forms.
Descriptors: energy metabolism physiology, locomotion physiology, mammals physiology, biomechanics, body temperature regulation physiology, platypus physiology, swimming physiology, whales physiology.

Fish, F.E. (1998). Comparative kinematics and hydrodynamics of odontocete cetaceans: morphological and ecological correlates with swimming performance. Journal of Experimental Biology 201(20): 2867-77. ISSN: 0022-0949.
NAL Call Number: 442.8 B77
Abstract: Propulsive morphology and swimming performance were compared for the odontocete cetaceans Delphinapterus leucas, Orcinus orca, Pseudorca crassidens and Tursiops truncatus. Morphological differences were apparent among the whales. The general body contour and low-aspect-ratio caudal flukes of D. leucas indicated that this species was a low-performance swimmer compared with the other species. Propulsive motions were video-taped as animals swam steadily in large pools. Video tapes were analyzed digitally using a computerized motion-analysis system. Animals swam at relative velocities ranging from 0.4 to 2.4 body lengths s-1. The stroke amplitude of the flukes decreased linearly with velocity for D. leucas, but amplitude remained constant for the other species. Tail-beat frequencies were directly related to relative swimming velocity, whereas the pitch angle of the flukes was inversely related to relative swimming velocity. Unsteady lifting-wing theory was used with regression equations based on kinematics to calculate thrust power output, drag coefficients and propulsive efficiency. Compared with other species, O. orca generated the largest thrust power (36.3 kW) and had the lowest drag coefficient (0.0026), whereas T. truncatus displayed the largest mass-specific thrust power (23.7 W kg-1) and P. crassidens had the highest efficiency (0.9). D. leucas did not swim as rapidly as the other species and had a comparatively higher minimum drag coefficient (0.01), lower mass-specific thrust power (5.2 W kg-1) and lower maximum efficiency (0.84). Minimum drag coefficients were associated with high swimming speeds, and maximum efficiencies corresponded with velocities in the range of typical cruising speeds. The results indicate that the kinematics of the propulsive flukes and hydrodynamics are associated with the swimming behaviors and morphological designs exhibited by the whales in this study, although additional factors will influence morphology.
Descriptors: Cetacea physiology, swimming physiology, biomechanics.

Fish, F.E. and J.M. Battle (1995). Hydrodynamic design of the humpback whale flipper. Journal of Morphology 225(1): 51-60. ISSN: 0362-2525.
NAL Call Number: 444.8 J826
Abstract: The humpback whale (Megaptera novaeangliae) is reported to use its elongate pectoral flippers during swimming maneuvers. The morphology of the flipper from a 9.02-m whale was evaluated with regard to this hydrodynamic function. The flipper had a wing-like, high aspect ratio planform. Rounded tubercles were regularly interspersed along the flipper's leading edge. The flipper was cut into 71 2.5-cm cross-sections and photographed. Except for sections near the distal tip, flipper sections were symmetrical with no camber. Flipper sections had a blunt, rounded leading edge and a highly tapered trailing edge. Placement of the maximum thickness placement for each cross-section varied from 49% of chord at the tip to 19% at mid-span. Section thickness ratio averaged 0.23 with a range of 0.20-0.28. The humpback whale flipper had a cross-sectional design typical of manufactured aerodynamic foils for lift generation. The morphology and placement of leading edge tubercles suggest that they function as enhanced lift devices to control flow over the flipper and maintain lift at high angles of attack. The morphology of the humpback whale flipper suggests that it is adapted for high maneuverability associated with the whale's unique feeding behavior.
Descriptors: extremities anatomy and histology, locomotion, whales anatomy and histology, adaptation, anatomic biological, models.

Fish, F.E., J.E. Peacock, and J.J. Rohr (2003). Stabilization mechanism in swimming odontocete cetaceans by phased movements. Marine Mammal Science 19(3): 515-528. ISSN: 0824-0469.
NAL Call Number: QL713.2.M372
Descriptors: movement and support, swimming speed, vertical oscillations, cetaceans, propulsive movements, stability, energy expenditure.

Fish, F.E. (2004). Evolution of swimming mechanics in cetaceans. Journal of Morphology 260(3): 291. ISSN: 0362-2525.
NAL Call Number: 444.8 J826
Descriptors: evolution and adaptation, aquatic environment, swimming mechanics, diving, maneuverability, propulsive performance.
Notes: Meeting Information: Seventh International Congress of Vertebrate Morphology, Boca Raton, FL, USA, 2004.

Fish, F.E. (2002). Balancing requirements for stability and maneuverability in cetaceans. Integrative and Comparative Biology 42(1): 85-93.
NAL Call Number: QL1.I67
Descriptors: morphology, body design variability, body flexibility, dynamic stabilization, maneuverability, morphological design, stability, turning performance, balancing requirements, cetaceans, control surfaces.

Geertsen, B.M., J. Teilmann, R. Kastelein, H.N.J. Vlemmix, and L.A. Miller (2004). Behaviour and physiological effects of transmitter attachments on a captive harbour porpoise (Phocoena phocoena). Journal of Cetacean Research and Management 6(2): 139-146. ISSN: 1561-0713.
Abstract: A captive harbour porpoise (Phocoena phocoena) was monitored for 80 consecutive days, 10 days before attachment of a satellite dive recorder and a VHF-radio tag, 30 days during attachment and 40 days after removal of the transmitters. Dive data recorded by the satellite transmitter was collected during the attachment. Daily food intake was measured and each week the porpoise was taken out of the water for a physical examination. Behavioural observations logged on the handheld computer showed an immediate effect of the tagging in time spent resting at the surface (logging), which was four to six times higher on the day of attachment. Digital video recordings showed a significant increase in the mean duration of rolls at the surface immediately after attachment. The mean duration of dives was shorter before attachment than both after the tagging and after removal of the transmitters. Furthermore the frequency of surfacings farthest away from where the porpoise was taken out of the pool for tagging, was highest the first five days following the tagging. Dive data from the satellite tag showed a semidiurnal diving pattern, with increased mean dive depth in the first 24 hours after attachment. The heart rate was fairly constant during the tagging. but the mean heart rate increased significantly from 161 beats per minute (bpm) to 180 bpm after the first hole in the dorsal fin was made. The body weight of the porpoise increased up to the time of tagging (16 May 2000), after which it decreased until six days prior to release (28 July 2000) this was probably due to the seasonal trend in blubber thickness of harbour porpoises rather than an effect from the tagging. After one month of attachment, a reaction occurred around the frontal pinhole and the transmitters were removed. This reaction was probably due to drag from two tags and seaweed attached to the tags during the last part of the attachment period. After the tags were removed epithelia closed the pinholes after two days.
Descriptors: Phocoena phocoena, tracking techniques, satellite dive recorder and vhf radio tag, behavioral and physiological effects, weight, body weight, feeding rate, heart beat, swimming, aquatic diving, behavior.

Gillett, M.A., C.L. Efremoff, and E.B. Giffin (1994). Vertebral variation and locomotor pattern in odontocete cetaceans. Journal of Vertebrate Paleontology 14(3 Suppl.): 27A. ISSN: 0272-4634.
Descriptors: evolution and adaptation, morphology, paleobiology, skeletal system, movement and support, systematics and taxonomy, evolution, forelimbs, fossils, meeting abstract, swimming, fossil.
Notes: Meeting Information: Fifty-fourth Annual Meeting of the Society of Vertebrate Paleontology, Seattle, Washington, USA, 1994.

Halsey, L. (2002). Ultimate divers. Biologist (London) 49(4): 161-4. ISSN: 0006-3347.
NAL Call Number: 442.9 IN7
Abstract: An extraordinary variety of mammals, birds, amphibians and reptiles are adapted to life in the oceans. Many of them spend their entire lives there without being able to breathe underwater. But just how do they exploit this hostile environment?
Descriptors: adaptation, physiological, birds physiology, diving physiology, respiration, whales physiology, evolution, oceans and seas.
Notes: Comment In: Biologist (London). 2002 Oct;49(5):192; author reply 192.

Heide Jorgensen, M.P., N. Hammeken, R. Dietz, J. Orr, and P.R. Richard (2001). Surfacing times and dive rates for narwhals (Monodon monoceros) and belugas (Delphinapterus leucas). Arctic 54(3): 284-298. ISSN: 0004-0843.
Abstract: Time spent at and near the sea surface was measured for 25 narwhals, Monodon monoceros, and 39 belugas or white whales, Delphinapterus leucas, in West Greenland and Canada from 1993 through 1999, using satellite-linked data recorders. Narwhals spent less time at the surface than belugas did, and the surfacing time of belugas also varied between localities. No clear differences in surfacing time were associated with the time of day, but belugas tended to make more dives during the night than during the day. Despite large variability in surfacing behaviour among individual whales, time spent at the surface by both species declined from August through November. The few data collected from narwhals from November to February indicate that surfacing times remained low during this period although more than 25% of each 6 h period was spent at the surface. Whales made between 2 and 20 dives per hour, and narwhals made significantly fewer dives than did belugas, for which number of dives varied with locality. The number of dives deeper than 8 m declined substantially during the autumn for belugas and narwhals that were moving offshore. When travelling, the whales apparently made fewer dives than at other times.
Descriptors: Delphinapterus leucas, Monodon monoceros, aquatic diving, dive depths, seasonal activity, circadian activity, Arctic Ocean, Canada and Greenland, west, diving depths, rates and surfacing times, daily and seasonal patterns.

Heide, J.M.P., D. Bloch, E. Stefansson, B. Mikkelsen, L.H. Ofstad, and R. Dietz (2002). Diving behaviour of long-finned pilot whales Globicephala melas around the Faroe Islands. Wildlife Biology 8(4): 307-313. ISSN: 0909-6396.
NAL Call Number: SK351.W663
Descriptors: behavior, marine ecology, satellite tracking, applied and field techniques, body size, continental slopes, diving behavior, habitat use, niche utilization, offshore areas, swimming speed, water columns, water depth, long finned pilot whales, Faroe Islands.

Hind, A.T. and W.S. Gurney (1997). The metabolic cost of swimming in marine homeotherms. Journal of Experimental Biology 200(3): 531-42. ISSN: 0022-0949.
NAL Call Number: 442.8 B77
Abstract: This paper describes a model of the metabolic cost of swimming in pinnipeds and its application to other marine homeotherms. The model takes account of both hydrodynamic and thermal processes. The thermal component incorporates both free and forced convection and takes account of the effect of hair on free convection. Using data from the literature to evaluate all but two of the parameters, we apply the model to metabolic rate data on phocid seals, otariids (sea lions), penguins and minke whales. We show that the model is able to reproduce two unusual features of the data; namely, a very rapid increase in metabolic rate at low velocities and an overall rise in metabolic rate with velocity which is slower than the rise in hydrodynamic drag force. The work shows the metabolic costs of propulsion and thermoregulation in a swimming homeotherm to be interlinked and suggests differing costs of propulsion for different modes of swimming. This is potentially of ecological significance since the swimming speed that minimises the cost of transport for an animal will change with changes in water temperature.
Descriptors: models, biological, swimming physiology, behavior, animal, energy metabolism, mammals.

Houser, D.S., R. Howard, and S. Ridgway (2001). Can diving-induced tissue nitrogen supersaturation increase the chance of acoustically driven bubble growth in marine mammals? Journal of Theoretical Biology 213(2): 183-95. ISSN: 0022-5193.
NAL Call Number: 442.8 J8223
Abstract: The potential for acoustically mediated causes of stranding in cetaceans (whales and dolphins) is of increasing concern given recent stranding events associated with anthropogenic acoustic activity. We examine a potentially debilitating non-auditory mechanism called rectified diffusion. Rectified diffusion causes gas bubble growth, which in an insonified animal may produce emboli, tissue separation and high, localized pressure in nervous tissue. Using the results of a dolphin dive study and a model of rectified diffusion for low-frequency exposure, we demonstrate that the diving behavior of cetaceans prior to an intense acoustic exposure may increase the chance of rectified diffusion. Specifically, deep diving and slow ascent/descent speed contributes to increased gas-tissue saturation, a condition that amplifies the likelihood of rectified diffusion. The depth of lung collapse limits nitrogen uptake per dive and the surface interval duration influences the amount of nitrogen washout from tissues between dives. Model results suggest that low-frequency rectified diffusion models need to be advanced, that the diving behavior of marine mammals of concern needs to be investigated to identify at-risk animals, and that more intensive studies of gas dynamics within diving marine mammals should be undertaken.
Descriptors: Cetacea physiology, diving physiology, lung metabolism, nitrogen metabolism, sound adverse effects, diffusion, models, biological.

Ianov, V.G. (2001). Vliianie dopolnitel'nykh nagruzok na kinematiku i gidrodinamiku del'finov. [Effect of additional loading on kinematics and hydrodynamics of dolphins]. Biofizika 46(3): 563-72. ISSN: 0006-3029.
Abstract: The results of comparative measurements of hydrodynamic resistance of a solid model of dolphin and of additional load imitating an autonomous apparatus placed on the dolphin body are presented. The loading included cylinders (with rounded anterior and posterior ends) and spheres. They were fixed on the dorsal fin of the dolphin model in such a way that the interrelationship between the load and the model by a water stream minimal. In several experiments, a thick ring (dr = 10 mm) was fixed in the region of the dolphin respiratory opening to reach the turbulence of the boundary layer. The dolphin model with loads was towed in a speed range between 1.0 and 4.5 m/s. It was shown that the hydrodynamic resistance of additional loads is comparable with the resistance of the solid dolphin model. For the entire spectrum of loading in the preset range of towing speeds, the coefficient of hydrodynamic resistance of loading (according to the area of the wet body surface) was from 10 to 145% of the coefficient of the dolphin model resistance. It was assumed that the results of the measurements of kinematics and hydrodynamic characteristics of a freely-swimming dolphin can substantially differ from the results obtained on a dolphin carrying devices (at restricted power of the animal). Similar problems can also arise if the animals will carry devices with a higher hydrodynamic resistance.
Descriptors: dolphins physiology, swimming physiology, biomechanics, models, biological, water.
Language of Text: Russian.

Kingsley, M.C.S., S. Gosselin, and G.A. Sleno (2001). Movements and dive behaviour of belugas in northern Quebec. Arctic 54(3): 262-275. ISSN: 0004-0843.
Abstract: Three adult and three juvenile belugas were fitted with satellite-linked radio tags in eastern Hudson Bay in mid-August 1993, and one adult was tagged in mid-October 1995 in extreme northeastern Hudson Bay. The tags transmitted data on dive behaviour, and the receiving satellite calculated positions by Doppler-shift triangulation. The belugas tagged in summer in eastern Hudson Bay made no directed or long-distance movements while the tags were attached. Their range did not include the Belcher Islands, and belugas observed in aerial surveys of those islands do not appear to belong to the eastern coastal stock. The single beluga tagged in northern Quebec in October moved into the deep water of western Hudson Strait and travelled east along the southern coast of Hudson Strait, slowing up on reaching shallower water off Salluit and near Charles Island. This whale was still off Salluit when the tag stopped transmitting. All the tagged belugas dived intensively while the tags were attached, although there were individual differences, some belugas diving noticeably less than others. Dive behaviour varied over time, with periods lasting several days of concentrated diving interspersed with periods of less intense diving. 'Intense' periods entailed diving for up to 80% of the time. All belugas, even the one that was in deep water in Hudson Strait, showed dive depth characteristics that were consistent with diving usually to the bottom. However, all belugas always - even in deep water - made dives that usually lasted less than 10 min and very seldom lasted more than 12 min. Belugas tagged as pairs of adults and young showed striking correlations of dive behaviour. The data obtained indicate that it would be appropriate to correct aerial surveys by adding 85% to aerial counts.
Descriptors: Delphinapterus leucas, aquatic diving, behavior, distribution within habitat, north Atlantic, Canada, Quebec, Hudson Bay and Hudson Strait, distribution within habitat and diving behavior.

Klima, M. (1992). Schwimmbewegungen und Auftauchmodus bei Walen und Ichthyosaurieren. 1. Anatomische Grundlagen der Schwimmbewegungen. [Motions of swimming and emerge of whales and ichthyosaurians. 1. Anatomic demands of swimming movements]. Natur Und Museum 122(1): 1-17. ISSN: 0028-1301.
NAL Call Number: 509 F856
Descriptors: whales, dolphins, lizards, animal morphology, locomotion, movement, fins, spine, muscles, abdomen, animal morphology, body regions, bones, Cetacea, mammals, movement, musculoskeletal system, physiological functions, reptiles.
Language of Text: German summary.

Klima, M. (1992). Schwimmbewegungen und Auftauchmodus bei Walen und Ichtyosaurieren. 2. Vergleich das Auftauchmodus. [Motions of swimming and emerge of whales and ichthyosaurians. 2. Comparison of the emergence modus]. Natur Und Museum 122(3): 73-89. ISSN: 0028-1301.
NAL Call Number: 509 F856
Descriptors: whales, dolphins, lizards, animal morphology, locomotion, movement, respiratory system, nose, animal morphology, Cetacea, mammals, movement, physiological functions, reptiles, respiratory system.
Language of Text: German summary.

Kooyman, G.L. (1972). Deep diving behaviour and effects of pressure in reptiles, birds, and mammals. Symposia of the Society for Experimental Biology 26: 295-311. ISSN: 0081-1386.
Descriptors: birds physiology, diving, mammals physiology, pressure, reptiles physiology, body weight, Cetacea, dolphins, hydrostatic pressure, nitrogen blood, Pinnipedia, turtles, water.

Kozlov, L.P. and O.D. Nikishova (1973). Bioenergetic and hydrodynamic aspects of dolphin swimming. Visnyk Akademiyi Nauk Ukrayins'Koyi RSR (2): 42-53.
NAL Call Number: TRANSL 20094
Descriptors: dolphin swimming, bioenergetic, hyrodynamic.
Notes: JPRS: 59056; CSO: 1850-S. Translated by the Joint Publications Research Service, 1973.

Lafortuna, C.L., M. Jahoda, A. Azzellino, F. Saibene, and A. Colombini (2003). Locomotor behaviours and respiratory pattern of the Mediterranean fin whale (Balaenoptera physalus). European Journal of Applied Physiology 90(3-4): 387-95. ISSN: 1439-6319.
Abstract: Twenty-four Mediterranean fin whales were tracked in open sea with a method based on the assessment of the animal differential position in respect of the observer's absolute position aboard a vessel, with the concomitant recording of the respiratory activity. Short distance video recording was also performed in two whales, permitting the simultaneous determination of single breath expiratory (TE) and inspiratory (TI) durations. In the 24 whales swimming at an average velocity of 1.39 (0.47) m.s(-1) [mean (SD), range: 0.62-2.44 m.s(-1)], 2068 breaths organized in 477 respiratory cycles were observed. Each cycle entailed a prolonged apnoea dive phase [225 (91) s, Tdive) followed by a period near the surface [62 (28) s, surfacing], during which a series of breaths [4.6 (1.8)] was performed at short intervals. On the basis of track length and swimming velocity, two groups of animals were devised differing for convolution of the course (p<0.001), extension of ranging territory (p<0.01) and horizontal swimming velocity (p<0.05), which may represent two distinct behaviours. A possibly general mechanism of control of breathing in cetaceans was found, consistent with a model of constant tidal volume and variable respiratory frequency. Coherently with this model, TE was independent of TI or Tdive, in line with a passive expiration, while TI appeared to be negatively correlated with Tdive (p<0.05), otherwise suggesting, similarly with terrestrial mammals, a significant role of hypercapnic stimulation. The estimated O2 consumption of about 150 l.min(-1) is in line with the general allometric regression for mammals and corresponds to an energetic expenditure of 85-95 kJ.kg(-1).day(-1).
Descriptors: locomotion physiology, respiratory mechanics physiology, whales physiology, behavior, animal physiology, diving physiology, energy metabolism physiology, exhalation physiology, inhalation physiology, kinetics, Mediterranean Sea, oxygen consumption physiology, pulmonary ventilation physiology, radar, swimming physiology, tidal volume physiology.

Lockyer, C. (1981). Estimates of growth and energy budget for the sperm whale, Physeter catodon. In: J. Gordon-Clark (Editor), Mammals in the Seas. General Papers and Large Cetaceans. FAO Advisory Committee on Marine Resources Research, Working Party on Marine Mammals, FAO Fisheries Series, Vol. 3, FAO: Rome (Italy), p. 489-504. ISBN: 92-5-100513-3.
NAL Call Number: QL713.2.F66
Descriptors: sperm whales, nutrient balance, growth, energy budget, Physeter catodon.
Language of Text: English, Spanish, and French summaries.

Lockyer, C. (1981). Growth and energy budgets of large baleen whales from the southern hemisphere. In: J. Gordon-Clark (Editor), Mammals in the Seas. General Papers and Large Cetaceans. FAO Advisory Committee on Marine Resources Research, Working Party on Marine Mammals, FAO Fisheries Series, Vol. 3, FAO: Rome (Italy), p. 379-487. ISBN: 92-5-100513-3.
NAL Call Number: QL713.2.F66
Descriptors: whales, nutrition physiology, growth, energy budgets, baleen whales, FAO.
Language of Text: English, Spanish, and French summaries.

Lockyer, C. (1976). Growth and energy budgets of large baleen whales from the southern hemisphere. Fisheries Dept. Advisory Committee on Marine Resources Research. Scientific Consultation on Marine Mammals, August 31, 1976, Bergen, Norway, FAO: Rome, Italy, Vol. No. 34403, 180 p.
Descriptors: growth, energy budgets, large baleen whales, southern hemisphere.
Language of Text: English summary.

Lockyer, C., G. Desportes, K. Hansen, S. Labberte, and S. Siebert (2003). Monitoring growth and energy utilisation of the harbour porpoise (Phocoena phocoena) in human care. North Atlantic Marine Mammal Commission (NAMMCO) Scientific Publications 5: 107-120. ISSN: 1560-2206.
Abstract: Two harbour porpoises of an estimated age of 1-2 yrs were held in captivity from April 1997 and were still alive in April 2002, after rescue from pound nets set in inner Danish waters. They are presently housed in an outdoor penned-off area of Kerteminde fjord. Their growth (total body length, girth, body weight and blubber thickness) and daily dietary intake (weight of fish, dietary composition and energy value) have been monitored since capture. The general activity of the animals was regularly monitored, including two 24-hour long observation periods. Initial body weights were 37.5 kg for Eigil (male) and 40.5 kg for Freja (female). Both porpoises lost 4 to 5 kg in the first few days because of their initial refusal to feed from the hand. Then body weight increased steadily reaching a peak of 44.75 kg for Eigil and 51.6 kg for Freja in early February 1998. A fluctuation in body weight with peaks of 44 to 45 kg for the male and 51to 56 kg for the female in winter followed by lows of 41 to 44 kg and 47 to 48 kg respectively in summer, established a clear pattern of seasonal fluctuation, mirrored by girth and blubber thickness variation. Length increased steadily from 130.5 cm to 139cm in Eigil, and from 127.5 cm to 150 cm in Freja. Food intake also fluctuated seasonally, and increases in food intake preceded weight gains. Daily food consumption in Eigil and Freja represented about 7 to 9.5% of body weight. The growth of the animals resembles that of wild porpoises in the region. The sudden initial weight losses suggested that the energy reserves of the animals may only be short-term. The large weight increase in the winter months with colder water, correlating with the increase in girth and blubber thickness, suggest that energy reserves and blubber fat may be important for insulation. During the two 24-hour observations, the animals spent most of their time cruising around, although slow swimming and logging at the surface increased at night. Breathing rates were lower in the early morning hours, consistent with diminished activity. Both animalsÆ movements were influenced by external activities at poolside.
Descriptors: Phocoena phocoena, weight, energy expenditure, energy utilization, feeding rate, food intake, growth, food intake and energy utilization relations, Baltic Sea, Denmark, Kerteminde Fjord, growth and energy utilization, captive study.

Long Jr., J.H., D.A. Pabst, W.R. Shepherd, and W.A. McLellan (1997). Locomotor design of dolphin vertebral columns: bending mechanics and morphology of Delphinus delphis. Journal of Experimental Biology 200(1): 65-81. ISSN: 0022-0949.
NAL Call Number: 442.8 B77
Abstract: The primary skeletal structure used by dolphins to generate the dorsoventral bending characteristic of cetacean swimming is the vertebral column. In the vertebral column of the saddleback dolphin Delphinus delphis, we characterize the static and dynamic mechanical properties of the intervertebral joints, describe regional variation and dorsoventral asymmetries in mechanical performance, and investigate how the mechanical properties are correlated with vertebral morphologies. Using a bending machine that applies an external load (N m) to a single intervertebral segment, we measured the resulting angular deformation (rad) of the segment in both dorsal extension and ventral flexion. Intervertebral segments from the thoracic, lumbar and caudal regions of the vertebral column were tested from five individuals. Using quasi-static bending tests, we measured the initial (low-strain) bending stiffness (N m rad-1) as a function of segment position, direction of bending (extension and flexion) and sequential cutting of intervertebral ligaments. We found that initial bending stiffness was significantly greater in the lumbar region than in adjacent thoracic and caudal regions, and all joints were stiffer in extension than is predicted (r2 = 0.554) by the length and width of the intervertebral disc and the length of the cranial vertebral body in the segment. Stiffness in flexion is predicted (r2 = 0.400) by the width of the nucleus pulposus, the length of the caudal vertebral body in the segment and the height of the transverse processes from the ventral surface of the vertebral body. We also performed dynamic bending tests on intervertebral segments from the lumbo-caudal joint and the joint between caudal vertebrae 7 and 8. Dynamic bending stiffness (N m rad-1) increases with increasing bending amplitude and is independent of bending frequency. Damping coefficient (kg m2 rad-2 s-1) decreases with increasing bending amplitude and frequency. Resilience (% energy return) increases from approximately 20% at low bending amplitudes (+/-0.6 degree) to approximately 50% at high bending amplitudes (+/-2.9 degrees). Based on these findings, the dolphin's vertebral column has the mechanical capacity to help control the body's locomotor reconfigurations, to store elastic energy and to dampen oscillations.
Descriptors: dolphins physiology, locomotion, spine physiology, biomechanics, joints physiology, ligaments physiology, muscles physiology.

Madar, S. and J.G.M. Thewissen (2003). The locomotor repertoire of pakicetid cetaceans: combined evidence of gross morphology and skeletal ultrastructure. Journal of Vertebrate Paleontology 23(3, Suppl.): 74A. ISSN: 0272-4634.
Descriptors: evolution and adaptation, paleobiology, skeletal system, movement and support, amphibious behavior, buoyancy control, cursorial behavior, evolutionary changes, gross morphology, locomotor repertoires, aquatic, postcranial remains, skeletal ultrastructure.
Notes: Meeting Information: Sixty-Third Annual Meeting of the Society of Vertebrate Paleontology, St. Paul, MN, USA, 2003.

Maresh, J.L., F.E. Fish, D.P. Nowacek, S.M. Nowacek, and R.S. Wells (2004). High performance turning capabilities during foraging by bottlenose dolphins (Tursiops truncatus). Marine Mammal Science 20(3): 498-509. ISSN: 0824-0469.
NAL Call Number: QL713.2.M372
Abstract: Large predators should have difficulty catching small prey because small animals demonstrate greater maneuverability and agility compared to large animals. The ability of a predator to capture small prey indicates locomotor strategies to compensate for inequities in maneuverability. Bottlenose dolphins (Tursiops truncatus) in Sarasota Bay, Florida feed on fish at least one order of magnitude smaller than themselves. To examine the locomotor strategies involved in prey capture, the foraging movements of these dolphins were videotaped from overhead using a remotely-controlled camera suspended from a helium-filled aerostat, which was tethered to an observation vessel. Dolphins were observed to rapidly maneuver during chases of fish in open water or around patches of rooted vegetation. Video analysis of the chase sequences indicated that the dolphins could move the rostrum through small radius turns with a mean value of 0.20 body lengths and with a minimum value of 0.08 body lengths. Mean rate of turn was 561.6degree/sec with a maximum rate measured at 1,372.0degree/sec. High turning rates with small turning radii were primarily the result of maneuvers in which the dolphin rolled 90degree and rapidly flexed its body ventrally. The ability of dolphins to change body orientation in multiple rotational axes provides a mechanism to reduce turning radius and increase turning rate to catch small, elusive prey.
Descriptors: biogeography, population studies, marine ecology, ecology, environmental sciences, video analysis, applied and field techniques, predator prey situations, turning rates.

Martin, A.R., P. Hall, and P.R. Richard (2001). Dive behaviour of belugas (Delphinapterus leucas) in the shallow waters of western Hudson Bay. Arctic 54(3): 276-283. ISSN: 0004-0843.
Abstract: Beluga diving was studied in western Hudson Bay, where the preferred habitat of this whale in summer is shallow coastal waters. No relationship was found between the duration of a dive and the surface interval either preceding or following it. During rare periods of intense diving, 77% of an average dive cycle was spent below the water surface. Only in waters deeper than 25 m did any whale spend more than half its time below the surface zone; in rivers, less than 15% of the time was spent at depths of 4 m or more. The frequency of long dives increased with water depth. Maximum dive duration was 15.6 min. Dives to depths of 25 m or less were of variable duration and time-depth profile; most dives exceeding 50 m had a square profile and reached the seabed. The easterly migration in early September took the whales into water deeper than 100 m for the first time in several months, and they dived frequently to the bottom during this period. No clear difference in dive capability was found between western Hudson Bay belugas and those that inhabit deeper waters farther north. The preference of this population for shallow water in summer is not dictated by an inability to dive to greater depths; belugas can utilize benthic resources throughout Hudson Bay.
Descriptors: Delphinapterus leucas, aquatic diving, behavior, migration, autumn, habitat preference, north Atlantic, Canada, Hudson Bay, west, diving behavior, habitat preference and autumn migration.

Mate, B.R., B.A. Lagerquist, M. Winsor, J. Geraci, and J.H. Prescott (2005). Movements and dive habits of a satellite-monitored longfinned pilot whale (Globicephala melas) in the northwest Atlantic. Marine Mammal Science 21(1): 136-144. ISSN: 0824-0469.
NAL Call Number: QL713.2.M372
Descriptors: Globicephala melas, aquatic diving, north Atlantic, USA, movements and dive habits, seasonal patterns.

Mate, B.R. and J. Urban Ramirez (2003). A note on the route and speed of a gray whale on its northern migration from Mexico to central California, tracked by satellite-monitored radio tag. Journal of Cetacean Research and Management 5(2): 155-157. ISSN: 1561-0713.
Descriptors: Eschrichtius robustus, swimming, swimming speed, migration, north Pacific, Mexico to California, migratory route and speed, satellite monitored radio tagged specimen.

Miller, P.J., M.P. Johnson, P.L. Tyack, and E.A. Terray (2004). Swimming gaits, passive drag and buoyancy of diving sperm whales Physeter macrocephalus. Journal of Experimental Biology 207(11): 1953-67. ISSN: 0022-0949.
NAL Call Number: 442.8 B77
Abstract: Drag and buoyancy are two primary external forces acting on diving marine mammals. The strength of these forces modulates the energetic cost of movement and may influence swimming style (gait). Here we use a high-resolution digital tag to record depth, 3-D orientation, and sounds heard and produced by 23 deep-diving sperm whales in the Ligurian Sea and Gulf of Mexico. Periods of active thrusting versus gliding were identified through analysis of oscillations measured by a 3-axis accelerometer. Accelerations during 382 ascent glides of five whales (which made two or more steep ascents and for which we obtained a measurement of length) were strongly affected by depth and speed at Reynold's numbers of 1.4-2.8x10(7). The accelerations fit a model of drag, air buoyancy and tissue buoyancy forces with an r(2) of 99.1-99.8% for each whale. The model provided estimates (mean +/- S.D.) of the drag coefficient (0.00306+/-0.00015), air carried from the surface (26.4+/-3.9 l kg(-3) mass), and tissue density (1030+/-0.8 kg m(-3)) of these five animals. The model predicts strong positive buoyancy forces in the top 100 m of the water column, decreasing to near neutral buoyancy at 250-850 m. Mean descent speeds (1.45+/-0.19 m s(-1)) were slower than ascent speeds (1.63+/-0.22 m s(-1)), even though sperm whales stroked steadily (glides 5.3+/-6.3%) throughout descents and employed predominantly stroke-and-glide swimming (glides 37.7+/-16.4%) during ascents. Whales glided more during portions of dives when buoyancy aided their movement, and whales that glided more during ascent glided less during descent (and vice versa), supporting the hypothesis that buoyancy influences behavioural swimming decisions. One whale rested at approximately 10 m depth for more than 10 min without fluking, regulating its buoyancy by releasing air bubbles.
Descriptors: acceleration, diving, gait physiology, models, theoretical, swimming physiology, whales physiology, Atlantic Ocean, body weights and measures, Mediterranean Sea, video recording, drag, buoyancy.

Milsom, B. (2000). Breathless--by choice. Biologist (London) 47(5): 239-42. ISSN: 0006-3347.
NAL Call Number: 442.9 IN7
Abstract: During diving, hibernation and sleep, powerful inhibitory inputs act to produce breath holding and are periodically lifted to allow the expression of an intense drive to breathe. Many animals that do this also possess adaptations designed to prolong the periods of breath holding and shorten the periods of breathing.
Descriptors: birds physiology, oxygen consumption physiology, respiration, seals, earless physiology, whales physiology, diving physiology, hemoglobins physiology, hibernation physiology, sleep physiology.

Moore, P. (2004). Examining dolphin hydrodynamics provides clues to calf-loss during tuna fishing. Journal of Biology 3(2): 6. ISSN: 1475-4924.
Abstract: A combination of mathematical modeling and direct observation of the swimming behavior of dolphin mother-calf pairs has shown how the calf can gain much of the energy required for swimming if it is positioned correctly relative to the mother, a situation that may be disrupted during the chases that result from tuna-fishing practices.
Descriptors: Delphinidae, fishing and fisheries, biomechanics, hydrodynamics, swimming, drafting hydrodynamics, implications for fishery related calf loss, mortality, calf loss.

Nachtigall, W. (1981). Hydromechanics and biology. Biophysics of Structure and Mechanism 8(1-2): 1-22. ISSN: 0340-1057.
Abstract: To exemplify relations between biology and hydrodynamics the Reynolds number range and the effects of viscosity and inertia in swimming and flying organisms is discussed. Comparing water beetles and penguins it is shown, that the technical drag coefficient is an adequate means to describe flow adaptation in animals. Compared to technical systems, especially the penguins'drag coefficient is astonishingly low. Furthermore, the question, why comparatively thick bodies in penguins and dolphins show rather low drag is discussed. Distributed boundary layer damping in dolphins and secretion of special high molecular slimes in fishes help to keep flow characteristics laminar. As an example of one easily understood thrust mechanism, the drag inducing pair of rowing legs in water, beetles is morphologically and hydrodynamically analysed. Fish swimming is discussed as a locomotion principle using lift components. Thrust generation by the moving tail fin of a fish is analysed in detail. Coming back to the influence if Reynolds number, it is finally shown, how very small, bristle bearing swimming legs and wings of insects make use of viscosity effects for locomotion.
Descriptors: biomechanics, birds physiology, dolphins physiology, water, beetles, locomotion, models, biological, swimming, viscosity.

Nawojchik, R., D.J. St. Aubin, and A. Johnson (2003). Movements and dive behavior of two stranded, rehabilitated long-finned pilot whales (Globicephala melas) in the northwest Atlantic. Marine Mammal Science 19(1): 232-239. ISSN: 0824-0469.
NAL Call Number: QL713.2.M372
Descriptors: Globicephala melas, diving, north west Atlantic, USA, movements and dive behavior after stranding and rehabilitation.

Nechayeva, O.V. (1975). Dolphin Propulsion, Bionika: 7 p.
NAL Call Number: TRANSL 20955
Descriptors: dolphin, propulsion.
Notes: JPRS 63848.

Noren, S.R., V. Cuccurullo, and T.M. Williams (2004). The development of diving bradycardia in bottlenose dolphins (Tursiops truncatus). Journal of Comparative Physiology. B, Biochemical, Systemic, and Environmental Physiology 174(2): 139-47. ISSN: 0174-1578.
NAL Call Number: QP33.J681
Abstract: Bradycardia is an important component of the dive response, yet little is known about this response in immature marine mammals. To determine if diving bradycardia improves with age, cardiac patterns from trained immature and mature bottlenose dolphins ( Tursiops truncatus) were recorded during three conditions (stationary respiration, voluntary breath-hold, and shallow diving). Maximum (mean: 117+/-1 beats.min(-1)) and resting (mean: 101+/-5 beats.min(-1)) heart rate (HR) at the water surface were similar regardless of age. All dolphins lowered HR in response to apnea; mean steady state breath-hold HR was not correlated with age. However, the ability to reduce HR while diving improved with age. Minimum and mean steady state HR during diving were highest for calves. For example, 1.5-3.5-year-old calves had significantly higher mean steady state diving HR (51+/-1 beats.min(-1)) than 3.5-5.5-year-old juveniles (44+/-1 beats.min(-1)). As a result, older dolphins demonstrated greater overall reductions in HR during diving. Longitudinal studies concur; the ability to reduce HR improved as individual calves matured. Thus, although newly weaned calves as young as 1.7 years exhibit elements of cardiac control, the capacity to reduce HR while diving improves with maturation up to 3.5 years postpartum. Limited ability for bradycardia may partially explain the short dive durations observed for immature marine mammals.
Descriptors: diving physiology, dolphins physiology, age factors, bradycardia etiology, bradycardia physiopathology, dolphins growth and development, heart rate physiology, respiratory physiology.

Noren, S.R. and T.M. Williams (2000). Body size and skeletal muscle myoglobin of cetaceans: adaptations for maximizing dive duration. Comparative Biochemistry and Physiology. A, Molecular and Integrative Physiology 126(2): 181-91. ISSN: 1095-6433.
NAL Call Number: QP1.C6
Abstract: Cetaceans exhibit an exceptionally wide range of body mass that influence both the capacities for oxygen storage and utilization; the balance of these factors is important for defining dive limits. Furthermore, myoglobin content is a key oxygen store in the muscle as it is many times higher in marine mammals than terrestrial mammals. Yet little consideration has been given to the effects of myoglobin content or body mass on cetacean dive capacity. To determine the importance of myoglobin content and body mass on cetacean diving performance, we measured myoglobin content of the longissimus dorsi for ten odontocete (toothed whales) and one mysticete (baleen whales) species ranging in body mass from 70 to 80000 kg. The results showed that myoglobin content in cetaceans ranged from 1.81 to 5.78 g (100 g wet muscle)(-1). Myoglobin content and body mass were both positively and significantly correlated to maximum dive duration in odontocetes; this differed from the relationship for mysticetes. Overall, the combined effects of body mass and myoglobin content accounts for 50% of the variation in cetacean diving performance. While independent analysis of the odontocetes showed that body mass and myoglobin content accounts for 83% of the variation in odontocete dive capacity.
Descriptors: adaptation, physiological physiology, body constitution, Cetacea physiology, diving physiology, muscle, skeletal physiology, myoglobin physiology, species specificity.

Noren, S.R., T.M. Williams, D.A. Pabst, W.A. McLellan, and J.L. Dearolf (2001). The development of diving in marine endotherms: preparing the skeletal muscles of dolphins, penguins, and seals for activity during submergence. Journal of Comparative Physiology. B, Biochemical, Systemic, and Environmental Physiology 171(2): 127-34. ISSN: 0174-1578.
NAL Call Number: QP33.J681
Abstract: Myoglobin is an important oxygen store for supporting aerobic diving in endotherms, yet little is known about its role during postnatal development. Therefore, we compared the postnatal development of myoglobin in marine endotherms that develop at sea (cetaceans) to those that develop on land (penguins and pinnipeds). We measured myoglobin concentrations in the major locomotor muscles of mature and immature bottlenose dolphins (Tursiops truncatus) and king penguins (Aptenodytes patagonicus) and compared the data to previously reported values for northern elephant seals (Mirounga angustirostris). Neonatal dolphins, penguins, and seals lack the myoglobin concentrations required for prolonged dive durations, having 10%, 9%, and 31% of adult values, respectively. Myoglobin contents increased significantly during subsequent development. The increases in myoglobin content with age may correspond to increases in activity levels, thermal demands, and time spent in apnea during swimming and diving. Across these phylogenetically diverse taxa (cetaceans, penguins, and pinnipeds), the final stage of postnatal development of myoglobin occurs during the initiation of independent foraging, regardless of whether development takes place at sea or on land.
Descriptors: body temperature regulation physiology, diving physiology, muscle development, muscle, skeletal growth and development, muscle, skeletal physiology, myoglobin physiology, age factors, apnea, birds, dolphins, oxygen physiology, seals, earless, species specificity.

Noren, S.R. (2004). Buffering capacity of the locomotor muscle in cetaceans: correlates with postpartum development, dive duration, and swim performance. Marine Mammal Science 20(4): 808-822. ISSN: 0824-0469.
NAL Call Number: QL713.2.M372
Abstract: Skeletal muscles of marine mammals must support the metabolic demands of exercise during periods of reduced blood flow associated with the dive response. Enhanced muscle buffering could support anaerobic metabolic processes during apnea, yet this has not been fully investigated in cetaceans. To assess the importance of this adaptation in the diving and swimming performance of cetaceans, muscle buffering capacity due to non-bicarbonate buffers was measured in the longissimus dorsi of ten species of odontocete and one mysticete. Immature specimens from a subset of these species were studied to assess developmental trends. Fetal and neonatal cetaceans have low buffering capacities (range: 34.8-53.9 slykes) that are within the range measured for terrestrial mammals. A lengthy developmental period, independent of muscle myoglobin postnatal development, is required before adult levels are attained. Adult cetacean buffering capacities (range: 63.7-94.5 slykes) are among the highest values recorded for mammals. Cetacean species that demonstrate extremely long dive durations or high burst speed swimming tend to have greater buffering capacities. However, the wide range of body size across cetaceans may complicate these trends. Enhanced muscle buffering capacity may enable small-bodied species to extend breath-hold beyond short aerobic dive limits for foraging or predator evasion when necessary.
Descriptors: Cetacea, biochemistry, proteins, myoglobin, skeletal musculature, locomotor muscles buffering capacity, postnatal development and swim, dive performance correlation, development, evolutionary adaptation, swimming, aquatic diving.

Noren, S.R., G. Lacave, R.S. Wells, and T.M. Williams (2002). The development of blood oxygen stores in bottlenose dolphins (Tursiops truncatus): implications for diving capacity. Journal of Zoology (London) 258(1): 105-113. ISSN: 0952-8369.
Descriptors: Tursiops truncatus, age, foraging, blood, hematological characters, respiratory gas transport, blood oxygen reserves, postnatal development and implications for diving capacity and foraging behavior, young development, diving, areobic dive limits.

Nowacek, D.P., M.P. Johnson, P.L. Tyack, K.A. Shorter, W.A. McLellan, and D.A. Pabst (2001). Buoyant balaenids: the ups and downs of buoyancy in right whales. Proceedings of the Royal Society of London. Series B. Biological Sciences 268(1478): 1811-6. ISSN: 0962-8452.
Abstract: A variety of marine mammal species have been shown to conserve energy by using negative buoyancy to power prolonged descent glides during dives. A new non-invasive tag attached to North Atlantic right whales recorded swim stroke from changes in pitch angle derived from a three-axis accelerometer. These results show that right whales are positively buoyant near the surface, a finding that has significant implications for both energetics and management. Some of the most powerful fluke strokes observed in tagged right whales occur as they counteract this buoyancy as they start a dive. By contrast, right whales use positive buoyancy to power glides during ascent. Right whales appear to use their positive buoyancy for more efficient swimming and diving. However, this buoyancy may pose added risks of vessel collision. Such collisions are the primary source of anthropogenic mortality for North Atlantic right whales, whose population is critically endangered and declining. Buoyancy may impede diving responses to oncoming vessels and right whales may have a reduced ability to manoeuvre during free ascents. These risk factors can inform efforts to avoid collisions.
Descriptors: swimming, whales physiology, behavior, animal, right whales, buoyant, balaenids, dives.

Oliver, J.S. and R.G. Kvitek (1984). Side-scan sonar records and diver observations of the gray whale (Eschrichtius robustus) feeding grounds. Biological Bulletin 167(1): 264-269. ISSN: 0006-3185.
NAL Call Number: 442.8 B52
Descriptors: grey whale, feeding grounds, sonar scan, observations.

Otani, S., Y. Naito, A. Kato, and A. Kawamura (2001). Oxygen consumption and swim speed of the harbor porpoise Phocoena phocoena. Fisheries Science (Tokyo) 67(5): 894-898. ISSN: 0919-9268.
NAL Call Number: SH1.F8195
Abstract: The rate of oxygen consumption and energetic cost of locomotion of captive harbor porpoises Phocoena phocoena were studied in Hokkaido, Japan, using respirometry together with externally attached devices (a data logger) that recorded swim speed and dive depth. Harbor porpoises swam freely in a tank at a swim speed ranging 0.5-4.2 m/s. Swimming during most dives was relatively slow with mean swim speed being 0.76-0.91 m/s. Oxygen consumption rate of the swimming harbor porpoise increased with swim speed according to a cubed function. The minimum cost of transport during underwater swimming in the harbor porpoise was 2.39-2.43 J/kg per m at an average swim speed of 1.3-1.5 m/s. However, porpoises usually swam slower to conserve energy and dived aerobically. This explains why harbor porpoises can dive repeatedly and continuously without extended rest at the sea surface.
Descriptors: Phocoena, oxygen consumption, locomotion, velocity., gas exchange, movement, physiological functions.

Otani, S. (2002). Studies of diving behavior of cetaceans. Aquabiology (Tokyo) 24(1): 58-62; 138. ISSN: 0285-4376.
NAL Call Number: QH90.A1K35
Descriptors: Cetacea, diving, review.

Otani, S., Y. Naito, A. Kato, and A. Kawamura (2000). Diving behavior and swimming speed of a free-ranging harbor porpoise, Phocoena phocoena. Marine Mammal Science 16(4): 811-814. ISSN: 0824-0469.
NAL Call Number: QL713.2.M372
Descriptors: Phocoena phocoena, swimming, speed, diving, dive depth, duration and ascent, descent angles and speeds, west Pacific, Japan, Hokkaido, Usujiri, diving behavior, swimming speed.

Pabst, D.A. (1996). Morphology of the subdermal connective tissue sheath of dolphins: a new fibre-wound, thin-walled, pressurized cylinder model for swimming vertebrates. Journal of Zoology (London) 238(1): 35-52. ISSN: 0952-8369.
NAL Call Number: QL1.J68
Abstract: The subdermal connective tissue sheath (SDS) of dolphins is a fibre-reinforced membrane connected to other locomotor tissues, including blubber, axial muscles and tendons, and vertebral column. The complicated connections between the SDS and other locomotor tissues suggest that the SDS acts as a peripheral skeletal element for the axial locomotor muscles and as an anchor for a de novo dermal appendage, the dorsal fin. The morphology of the SDS suggests that the dolphin can be modelled as a fibre-wound, thin-walled, pressurized cylinder. Existing cylinder models predict that the SDS functions to resist torsional forces, prevent aneurysms, and limit wrinkling when the dolphin bends in locomotion. I present a new functional model that more accurately represents the morphology of the dolphin cylinder wrapped by the SDS. The new model predicts that the SDS: (1) acts as a retinaculum for the terminal tendons of the axial locomotor muscles; and (2) plays a role in maintaining the laterally flattened cross-sectional shape of the caudal peduncle. The model is based on external morphological features of dolphins shared by other steady swimming aquatic vertebrates, such as carangiform and thunniform fishes. These features, which include a streamlined body shape and narrow necking of the caudal peduncle have been identified as adaptions to reduce drag. The new model offers insight into some of the structural features of the body wall required to maintain the hydrodynamically-tuned, external morphology of steady-swimming vertebrates.
Descriptors: biochemistry and molecular biophysics, integumentary system, chemical coordination and homeostasis, models and simulations, computational biology, morphology, muscular system, movement and support, physiology, axial locomotor muscles, body wall, carangiform fish, caudal peduncle, drag reduction, retinaculum, terminal tendons, thunniform fish.

Panigada, S., M. Zanardelli, S. Canese, and M. Jahoda (1999). How deep can baleen whales dive? Marine Ecology Progress Series 187: 309-311. ISSN: 0171-8630.
NAL Call Number: QH541.5.S3M32
Descriptors: behavior, marine ecology, ecology, environmental sciences, velocity time depth recorders, equipment, dive depth, baleen whales, fin whale.

Ridgway, S.H. (1971). Buoyancy regulation in deep diving whales. Nature (London) 232(5306): 133-134. ISSN: 0028-0836.
NAL Call Number: 472 N21
Descriptors: whales, deep diving, buoyancy, sperm whale, spermaceti, regulator, cooling, sea water, density.

Rodriguez, M., A. Ayala, S. Rodriguez, F. Rosa, and M. Diaz Gonzalez (2004). Application of the Max-Lloyd quantizer for ECG compression in diving mammals. Computer Methods and Programs in Biomedicine 73(1): 13-21. ISSN: 0169-2607.
NAL Call Number: RA409.5.C62
Abstract: This article presents a practical implementation of an ECG compression algorithm using a Max-Lloyd quantizer, to optimize the low resources of an ECG acquisition and transmission system (telemetry system) for dolphins and human divers. The algorithm scheme is based on a first-order differential pulse code modulation (DPCM) and uses a Max-Lloyd quantizer to code the difference between the current and predicted samples. The use of the non-uniform quantizer instead of a uniform quantizer improves the percent root mean-square difference (PRD), thereby producing a low distortion in the reconstructed signals. Due to its low computational complexity, the compression process can be accomplished on-line during the ECG acquisition process.
Descriptors: algorithms, diving physiology, dolphins physiology, electrocardiography.

Rohr, J., M.I. Latz, S. Fallon, J.C. Nauen, and E. Hendricks (1998). Experimental approaches towards interpreting dolphin-stimulated bioluminescence. Journal of Experimental Biology 201(9): 1447-60. ISSN: 0022-0949.
NAL Call Number: 442.8 B77
Abstract: Flow-induced bioluminescence provides a unique opportunity for visualizing the flow field around a swimming dolphin. Unfortunately, previous descriptions of dolphin-stimulated bioluminescence have been largely anecdotal and often conflicting. Most references in the scientific literature report an absence of bioluminescence on the dolphin body, which has been invariably assumed to be indicative of laminar flow. However, hydrodynamicists have yet to find compelling evidence that the flow remains laminar over most of the body. The present study integrates laboratory, computational and field approaches to begin to assess the utility of using bioluminescence as a method for flow visualization by relating fundamental characteristics of the flow to the stimulation of naturally occurring luminescent plankton. Laboratory experiments using fully developed pipe flow revealed that the bioluminescent organisms identified in the field studies can be stimulated in both laminar and turbulent flow when shear stress values exceed approximately 0.1 N m-2. Computational studies of an idealized hydrodynamic representation of a dolphin (modeled as a 6:1 ellipsoid), gliding at a speed of 2 m s-1, predicted suprathreshold surface shear stress values everywhere on the model, regardless of whether the boundary layer flow was laminar or turbulent. Laboratory flow visualization of a sphere demonstrated that the intensity of bioluminescence decreased with increasing flow speed due to the thinning of the boundary layer, while flow separation caused a dramatic increase in intensity due to the significantly greater volume of stimulating flow in the wake. Intensified video recordings of dolphins gliding at speeds of approximately 2 m s-1 confirmed that brilliant displays of bioluminescence occurred on the body of the dolphin. The distribution and intensity of bioluminescence suggest that the flow remained attached over most of the body. A conspicuous lack of bioluminescence was often observed on the dolphin rostrum and melon and on the leading edge of the dorsal and pectoral fins, where the boundary layer is thought to be thinnest. To differentiate between effects related to the thickness of the stimulatory boundary layer and those due to the latency of the bioluminescence response and the upstream depletion of bioluminescence, laboratory and dolphin studies of forced separation and laminar-to-turbulent transition were conducted. The observed pattern of stimulated bioluminescence is consistent with the hypothesis that bioluminescent intensity is directly related to the thickness of the boundary layer.
Descriptors: dinoflagellida physiology, dolphins physiology, luminescent measurements, rheology, dolphins anatomy and histology, swimming, video recording.

Rohr, J.J. and F.E. Fish (2004). Strouhal numbers and optimization of swimming by odontocete cetaceans. Journal of Experimental Biology 207(10): 1633-1642. ISSN: 0022-0949.
NAL Call Number: 442.8 B77
Abstract: Swimming efficiencies of fish and cetaceans have been related to a certain synchrony between stroke cycle frequency, peak-to-peak tail/fluke amplitude and mean swimming speed. These kinematic parameters form a non-dimensional wake parameter, referred to as a Stroulial number, which for the range between 0.20 and 0.40 has been associated with enhanced swimming efficiency for fish and cetaceans. Yet to date there has been no direct experimental substantiation of what Strouhal numbers are preferred by swimming cetaceans. To address this lack of data, a total of 248 Stroubal numbers were calculated for the captive odontocete cetaceans Tursiops truncatus, Pseudorca crassidens, Orcinus orca, Globicephala melaena, Lagenorhynchus obliquidens and Stenella frontalis. Although the average Stroulial number calculated for each species is within the accepted range, considerable scatter is found in the data both within species and among individuals. A greater proportion of Strouhal values occur between 0.20 and 0.30 (74%) than the 0.25-0.35 (55%) range predicted for maximum swimming efficiency. Within 0.05 Stroulial increments, the greatest number of Strouhal values was found between 0.225 and 0.275 (44%). Where propulsive efficiency data were available (Tursiops truncatus, Pseudorca crassidens, Orcinus orca), peak swimming efficiency corresponded to this same Stroulial range. The odontocete cetacean data show that, besides being generally limited to a range of Strouhal numbers between 0.20 and 0.40, the kinematic parameters comprising the Stroulial number provide additional constraints. Fluke-beat frequency normalized by the ratio of swimming speed to body length was generally restricted from 1 to 2, whereas peak-to-peak fluke amplitude normalized by body length occurred predominantly between 0.15 and 0.25. The results indicate that the kinematics of the propulsive flukes of odontocete cetaceans are not solely dependent on Stroulial number, and the Stroubal number range for odontocete cetaceans; occurs at slightly ([approximately]20%) lower values than previously predicted for maximum swimming efficiency.
Descriptors: odontoceti, biomechanics, swimming efficiency, swimming, efficiency, strouhal number significance.

Romanenko, E.V. (2002). Fish and Dolphin Swimming, Russian Academic Monographs, Sofia, Bulgaria, 429 p. ISBN: 9546421502.
NAL Call Number: QP310.S95 R66 2002
Descriptors: animal swimming, mathematical models, animal mechanics, hydrodynamics.

Romanenko, E.V. (1995). Assessment of traction and resistance coefficient in dolphins. Uspekhi Sovremennoi Biologii 115(1): 50-57. ISSN: 0042-1324.
NAL Call Number: 442.8 ER3
Descriptors: biochemistry and molecular biophysics, mathematical biology, computational biology, physiology, systematics and taxonomy, biophysics, mathematical analysis, movement.

Romanenko, E.V. (1995). Swimming of dolphins: experiments and modelling. Symposia of the Society for Experimental Biology 49: 21-33. ISSN: 0081-1386.
NAL Call Number: 442.9 SO15
Abstract: A critical analysis of the numerous works on dolphin swimming shows that several uncertainties led to incorrect conclusions concerning the estimates of dolphin drag coefficients. Original results on dolphin kinematics and hydrodynamics indicate the existence of a drag-reduction mechanism, caused by the formation of a negative pressure gradient along the body during active swimming. The mathematical model for this mechanism is presented.
Descriptors: dolphins physiology, swimming physiology, biophysics, mathematics, models, biological, stress, mechanical.

Romanenko, E.V. (1994). Hydrodynamics of dolphins. Uspekhi Sovremennoi Biologii 114(3): 283-303. ISSN: 0042-1324.
NAL Call Number: 442.8 ER3
Descriptors: climatology, environmental sciences, estuarine ecology, ecology, environmental sciences, mathematical biology, computational biology, models and simulations, computational biology, physiology, mathematical model, pressure distribution, swimming.

Tellone, E., M.E. Clementi, A.M. Russo, S. Ficarra, A. Lania, A. Lupi, B. Giardina and A. Galtieri (2000). Oxygen transport and diving behaviour: the haemoglobin from dolphin Tursiops truncatus. In: G. Di Prisco, B. Giardina and R.E. Weber (Editors), Hemoglobin Function in Vertebrates: Molecular Adaptation in Extreme and Temperate Environments, Springer-Verlag: Milano, Berlin, p. 77-82. ISBN: 8847001072.
NAL Call Number: QP96.5.H4485 2000
Descriptors: Tursiops truncatus, respiratory gas transport, oxygen transport, hemoglobin properties and adaptations to diving, captive study, respiratory pigments, hemoglobins, evolutionary adaptation, diving adaptations, diving, diving behavior.

Thode, A. (2004). Tracking sperm whale (Physeter macrocephalus) dive profiles using a towed passive acoustic array. Journal of the Acoustical Society of America 116(1): 245-53. ISSN: 0001-4966.
Abstract: A passive acoustic method is presented for tracking sperm whale dive profiles, using two or three hydrophones deployed as either a vertical or large-aperture towed array. The relative arrival times between the direct and surface-reflected acoustic paths are used to obtain the ranges and depths of animals with respect to the array, provided that the hydrophone depths are independently measured. Besides reducing the number of hydrophones required, exploiting surface reflections simplifies automation of the data processing. Experimental results are shown from 2002 and 2003 cruises in the Gulf of Mexico for two different towed array deployments. The 2002 deployment consisted of two short-aperture towed arrays separated by 170 m, while the 2003 deployment placed an autonomous acoustic recorder in tandem with a short-aperture towed array, and used ship noise to time-align the acoustic data. The resulting dive profiles were independently checked using single-hydrophone localizations, whenever multipath reflections from the ocean bottom could be exploited to effectively create a large-aperture vertical array. This technique may have applications for basic research and for real-time mitigation for seismic airgun surveys.
Descriptors: acoustics, diving, whales physiology, oceans and seas, sound, vocalization, animal physiology.

Thode, A., D.K. Mellinger, S. Stienessen, A. Martinez, and K. Mullin (2002). Depth-dependent acoustic features of diving sperm whales (Physeter macrocephalus) in the Gulf of Mexico. Journal of the Acoustical Society of America 112(1): 308-21. ISSN: 0001-4966.
Abstract: Three-dimensional dive trajectories of three sperm whales in the Gulf of Mexico have been obtained by measuring the relative arrival times and bearings of the animals' acoustic multipath reflections, using two elements of a towed hydrophone array deployed at an unknown depth and orientation. Within the first 6-12 min of the start of a dive, the intervals between successive "clicks" of all three whales corresponded closely with the two-way travel time of an acoustic pulse traveling vertically between the animals' position and the ocean bottom. The click spectra contained multiple peaks, including a faint band of energy originally centered near 10 kHz. As the animals descended over 500 m in depth, the center frequency of this band shifted to nearly 15 kHz, but subsequently remained near this value during the rest of the dive. This frequency shift is consistent with that expected from energy scattering from an ensemble of incompressible small-scale air-filled resonators, with diameters on the order of 4 mm. One possible candidate for such an ensemble is proposed to reside in the collapsed frontal sac of the animal. A comparison of the received levels for the bottom and direct multipath arrivals indicates that the whales' acoustic directivity must range between 10-30 dB in the 5-20-kHz region.
Descriptors: acoustics, diving, echolocation, models, biological, oceans and seas, whales.

Thomas, T.A., W.R. Koski, W.J. Richardson, and B. Wuersig (2002). Surfacing, respiration and dive cycles of bowhead whales in the eastern Alaskan Beaufort Sea during late summer and autumn as related to whale activity. OCS (Outer Continental Shelf) Study Report MMS (Minerals Management Service) 2002-012: 13.1-13.22.
Descriptors: Balaena mysticetus, feeding behavior, ventilation rate, respiratory rate, swimming, surfacing and dive cycles, social behavior, seasonal activity, feeding, travelling, social activities, migration, Arctic Ocean, USA, Alaska, Beaufort Sea, east, surfacing, respiration, dives cycles, bowhead whales.

Thomas, T.A., W.J. Richardson, W.R. Koski, and B. Wuersig (2002). Surfacing, respiration and dive cycles of bowhead whales in the Beaufort Sea: calves, subadults and adults. OCS (Outer Continental Shelf) Study Report MMS (Minerals Management Service) 2002-012: 14.1-14.19.
Descriptors: Balaena mysticetus, energy budget, respiration, development, young, subadult, adult, sdr cycle comparisons, swimming, surfacing, respiration and dive cycle, Arctic Ocean, USA, Alaska, Beaufort Sea, age, size, energetic implications.

Thomson, D.H. (2002). Energetics of bowhead whales. OCS (Outer Continental Shelf) Study Report MMS (Minerals Management Service) 2002-012: 22.1-22.40.
Descriptors: Balaena mysticetus, size, weight, metabolic rate, dietary requirements, daily and annual food requirements estimation, energy requirements, daily and annual energy requirements estimation, feeding grounds assessment, Arctic Ocean, USA, Alaska, Beaufort Sea, feeding grounds assement from daily and annual energy requirements.

Tytell, E. (2004). Humpbacks' bumpy flippers. Journal of Experimental Biology 207(21): IV. ISSN: 0022-0949.
NAL Call Number: 442.8 B77
Descriptors: Megaptera novaeangliae, forelimbs, effects on hydrodynamic performance, synopsis, biomechanics, flipper tubercles, swimming, hydrodynamic performance.

Videler, J. and P. Kamermans (1985). Differences between upstroke and downstroke in swimming dolphins. Journal of Experimental Biology 119: 265-74. ISSN: 0022-0949.
NAL Call Number: 442.8 B77
Abstract: Steady swimming movements of dolphins were recorded in a search for direct evidence of asymmetry between upstrokes and downstrokes. Kinematic swimming and gliding data from frame-by-frame analysis of cine pictures taken at constant frame rates with a camera in a fixed position are presented. We estimated the propulsive forces generated by the tail blade with a simple hydrodynamic model. Dolphins accelerate during the downstroke and decelerate during the upstroke: the net hydrodynamic force in the animal is always positive during the downstroke and negative during the upstroke. Both parts of the stroke cycle are equally long. The propulsive forces of downstrokes are on average larger than the forces of the upstrokes. Occasionally the average forces within an upstroke are greater than within a downstroke of the same sequence. Our data suggest that the drag on the body during the upstroke exceeds the drag in the course of the downstroke. The specific swimming speed or stride length of dolphins swimming at low speeds is about 0.9 body lengths per tail beat.
Descriptors: biomechanics, dolphins physiology, swimming, motion pictures, species specificity, tail physiology.

Watts, P. and F.E. Fish. (2003). Fluid dynamic and maneuverability aspects of humpback whale flippers. Annual Meeting and Exhibition of the SICB (Society for Integrative and Comparative Biology), Final Program and Abstracts, January 4, 2003-January 4, 2003, Toronto, ON, Canada, Vol. 2003, 336 p.
Descriptors: behavior, models and simulations, computational biology, morphology, movement and support, computer simulation, mathematical and computer techniques, adaptation, airfoil, attack angle, drag, lift, prey capture, hunpback whale, flippers, fluid, dynamic.

Weihs, D. (2002). Dynamics of dolphin porpoising revisited. Integrative and Comparative Biology 42(5): 1071-1078.
NAL Call Number: QL1.I67
Descriptors: Delphinidae, energy expenditure, swimming, porpoising, dynamics, model.

Weihs, D. (2004). The hydrodynamics of dolphin drafting. Journal of Biology 3(8): 8.1-8.16. ISSN: 1475-4924.
Abstract: BACKGROUND: Drafting in cetaceans is defined as the transfer of forces between individuals without actual physical contact between them. This behavior has long been surmised to explain how young dolphin calves keep up with their rapidly moving mothers. It has recently been observed that a significant number of calves become permanently separated from their mothers during chases by tuna vessels. A study of the hydrodynamics of drafting, initiated in the hope of understanding the mechanisms causing the separation of mothers and calves during fishing-related activities, is reported here. Results: Quantitative results are shown for the forces and moments around a pair of unequally sized dolphin-like slender bodies. These include two major effects. First, the socalled Bernoulli suction, which stems from the fact that the local pressure drops in areas of high speed, results in an attractive force between mother and calf. Second is the displacement effect, in which the motion of the mother causes the water in front to move forwards and radially outwards, and water behind the body to move forwards to replace the animalÆs mass. Thus, the calf can gain a æfree rideÆ in the forward-moving areas. Utilizing these effects, the neonate can gain up to 90% of the thrust needed to move alongside the mother at speeds of up to 2.4 m/sec. A comparison with observations of eastern spinner dolphins (Stenella longirostris) is presented, showing savings of up to 60% in the thrust that calves require if they are to keep up with their mothers. Conclusions: A theoretical analysis, backed by observations of free-swimming dolphin schools, indicates that hydrodynamic interactions with mothers play an important role in enabling dolphin calves to keep up with rapidly moving adult school members.
Descriptors: delphinidae, fishing and fisheries, biomechanics, hydrodynamics, swimming, drafting hydrodynamics, implications for fishing related mother calf separation.

Werth, A.J. (2004). Models of hydrodynamic flow in the bowhead whale filter feeding apparatus. Journal of Experimental Biology 207(20): 3569-3580. ISSN: 0022-0949.
NAL Call Number: 442.8 B77
Abstract: Anatomical and behavioral analyses suggest that the filtration mechanism of bowhead and right whales (Balaenidae) is driven by hydrodynamic as well as ram hydraulic pressures. Complementary models were devised to investigate biomechanical aspects of water flow in the buccal cavity of the bowhead whale Balaena mysticetus during continuous filter feeding. A mathematical model was created to test and quantify water flow predictions with steady state hydromechanical equations; a physical model of the bowhead mouth (approximately 1115 scale) was constructed to visualize flow processes. Both models rely on morphometric data obtained from whales harvested by Inupiat Eskimos for subsistence purposes along with information on foraging ecology (locomotor velocity, gape, etc.). Results indicate that unique features of balaenid oral construction and function (e.g. subrostral gap, orolabial sulcus, curvature of baleen, extensive mandibular rotation and lingual mobility) not only permit steady, unidirectional flow of water through the mouth, but also establish Bernoulli and Venturi effects during feeding. These hydrodynamic conditions allow balaenids to improve filtering efficiency and avoid creation of an anterior compressive wave (by increasing flow velocity and thereby reducing pressure) so that they may capture elusive prey even at slow swimming speeds.
Descriptors: Balaena mysticetus, biophysical techniques, mathematical techniques, hydromechanical modelling, mouth, biomechanics, hydrodynamics, feeding behavior, filter feeding, hydrodynamic flow patterns across feeding apparatus, mathematical model, buccal region, oral cavity, water movements, hydrodynamic flow.

Westgate, A.J., A.J. Read, P. Berggren, H.N. Koopman, and D.E. Gaskin (1995). Diving behaviour of harbour porpoises, Phocoena phocoena. Canadian Journal of Fisheries and Aquatic Sciences 52(5): 1064-1073. ISSN: 0706-652X.
NAL Call Number: 442.9 C16J
Abstract: The diving behaviour of seven free-ranging harbour porpoises (Phocoena phocoena) was examined using time-depth recorders. In total, 8167 individual dives were recorded over 254 h. The longest period of data collection from an individual was 106.1 h. Mean dive depths and durations ranged from 14 +- 16 to 41 +- 32 m, and from 44 +- 37 to 103 +- 67 s, respectively. The maximum recorded dive depth and duration was 226 m and 321 s. This performance may not represent the maximum capacity of harbour porpoises but rather the maximum depth of the study area. Individual dives had similar rates of descent and ascent, which ranged from 1.1 +- 0.6 to 2.3 +- 1.4 m/s, and from 0.9 +- 0.6 to 2.1 +- 1.4 m/s, respectively. Two porpoises with monitoring periods gt 2 days demonstrated a diel pattern in their diving, making fewer, but deeper dives at night. Comparison of the diving behaviour of harbour porpoises with data on the depth of 140 porpoises entanglements in groundfish gill nets in the Bay of Fundy showed these seven porpoises made between 22 and 70% of dives to depths (range 20-130 m) where the majority of entanglements were reported.
Descriptors: behavior, biosynchronization, estuarine ecology, ecology, environmental sciences, systematics and taxonomy, wildlife management, conservation, Bay of Fundy, diel pattern, diving behavior, ground fish gill net entanglement.

Williams, T.M., R.W. Davis, L.A. Fuiman, J. Francis, B.J. Le Boeuf, M. Horning, J. Calambokidis, and D.A. Croll (2000). Sink or swim: strategies for cost-efficient diving by marine mammals. Science 288(5463): 133-6. ISSN: 0036-8075.
NAL Call Number: 470 Sci2
Abstract: Locomotor activity by diving marine mammals is accomplished while breath-holding and often exceeds predicted aerobic capacities. Video sequences of freely diving seals and whales wearing submersible cameras reveal a behavioral strategy that improves energetic efficiency in these animals. Prolonged gliding (greater than 78% descent duration) occurred during dives exceeding 80 meters in depth. Gliding was attributed to buoyancy changes with lung compression at depth. By modifying locomotor patterns to take advantage of these physical changes, Weddell seals realized a 9.2 to 59.6% reduction in diving energetic costs. This energy-conserving strategy allows marine mammals to increase aerobic dive duration and achieve remarkable depths despite limited oxygen availability when submerged.
Descriptors: diving physiology, dolphins physiology, oxygen consumption, seals, earless physiology, swimming physiology, whales physiology, energy metabolism, hydrostatic pressure, lung physiology, lung volume measurements, video recording.
Notes: Comment In: Science. 2000 Apr 7;288(5463):83-5.

Williams, T.M., W.A. Friedl, M.L. Fong, R.M. Yamada, P. Sedivy, and J.E. Haun (1992). Travel at low energetic cost by swimming and wave-riding bottlenose dolphins. Nature (London) 355(6363): 821-3. ISSN: 0028-0836.
NAL Call Number: 472 N21
Abstract: Over the past 50 years there has been much speculation about the energetic cost of swimming and wave-riding by dolphins. When aligned properly in front of the bow of moving ships in the stern wake of small boats, on wind waves, and even in the wake of larger cetaceans, the animals appear to move effortlessly through the water without the benefit of propulsive strokes by the flukes. Theoretically, body streamlining as well as other anatomical and behavioural adaptations contribute to low transport costs in these animals. The economy of movement permitted by wave-riding has been perceived as an energetic advantage for the swimming dolphin, but has been hard to prove in the absence of physiological data for exercising cetaceans. Here we determine the aerobic and anaerobic costs of swimming and wave-riding in bottlenose dolphins and find that the minimum cost of transport for swimming dolphins is 1.29 +/- 0.05 J kg-1 m-1 at a cruising speed of 2.1 m s-1. Aerobic costs are nearly twice as high for swimming seals and sea lions, and 8-12 times higher for human swimmers. Wave-riding by dolphins provides additional benefits in terms of speed. The results indicate that behavioural, physiological and morphological factors make swimming an economical form of high-speed travel for dolphins.
Descriptors: dolphins physiology, energy metabolism, locomotion, heart rate, models, biological, swimming.

Williams, T.M., J.E. Haun, and W.A. Friedl (1999). The diving physiology of bottlenose dolphins (Tursiops truncatus). I. Balancing the demands of exercise for energy conservation at depth. Journal of Experimental Biology 202(20): 2739-48. ISSN: 0022-0949.
NAL Call Number: 442.8 B77
Abstract: During diving, marine mammals must rely on the efficient utilization of a limited oxygen reserve sequestered in the lungs, blood and muscles. To determine the effects of exercise and apnea on the use of these reserves, we examined the physiological responses of adult bottlenose dolphins (Tursiops truncatus) trained to breath-hold on the water surface or to dive to submerged targets at depths between 60 and 210 m. Changes in blood lactate levels, in partial pressures of oxygen and carbon dioxide and in heart rate were assessed while the dolphins performed sedentary breath-holds. The effects of exercise on breath-hold capacity were examined by measuring heart rate and post-dive respiration rate and blood lactate concentration for dolphins diving in Kaneohe Bay, Oahu, Hawaii. Ascent and descent rates, stroke frequency and swimming patterns were monitored during the dives. The results showed that lactate concentration was 1.1+/-0.1 mmol l(-1) at rest and increased non-linearly with the duration of the sedentary breath-hold or dive. Lactate concentration was consistently higher for the diving animals at all comparable periods of apnea. Breakpoints in plots of lactate concentration and blood gas levels against breath-hold duration (P(O2), P(CO2)) for sedentary breath-holding dolphins occurred between 200 and 240 s. In comparison, the calculated aerobic dive limit for adult dolphins was 268 s. Descent and ascent rates ranged from 1.5 to 2.5 m s(-1) during 210 m dives and were often outside the predicted range for swimming at low energetic cost. Rather than constant propulsion, diving dolphins used interrupted modes of swimming, with more than 75 % of the final ascent spent gliding. Physiological and behavioral measurements from this study indicate that superimposing swimming exercise on apnea was energetically costly for the diving dolphin but was circumvented in part by modifying the mode of swimming.
Descriptors: diving physiology, dolphins physiology, energy metabolism, physical conditioning, animal physiology, aerobiosis, apnea metabolism, carbon dioxide blood, carbon dioxide metabolism, dolphins blood, dolphins metabolism, heart rate, hydrogen ion concentration, lactic acid blood, lactic acid metabolism, oceans and seas, oxygen blood, oxygen metabolism, respiration, time factors.

Williams, T.M. (2001). Intermittent swimming by mammals: a strategy for increasing energetic efficiency during diving. American Zoologist 41(2): 166-176. ISSN: 0003-1569.
NAL Call Number: 410 AM3
Descriptors: Leptonychotes weddellii, Mirounga angustirostris, Tursiops truncatus, energy budget, diving, energetic efficiency strategy.

Yanov, V.G. (2001). Effect of additional loading on the kinematics and hydrodynamics of dolphin. Biofizika 46(3): 563-572. ISSN: 0006-3029.
Descriptors: Delphinidae, mathematical techniques, biomechanics, hydrodynamics during swimming, effect of additional loading, swimming.

Yanov, V.G. (2000). Structural and functional interrelation between swimming kinematics and specific features of dolphin's locomotory apparatus. Uspekhi Sovremennoi Biologii 120(1): 88-102. ISSN: 0042-1324.
NAL Call Number: 442.8 ER3
Descriptors: Tursiops truncatus, biophysics, musculature, skeleton, swimming, modes, muscular and skeletal functions, biophysical analysis.


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