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Survival of bowhead whales, Balaena mysticetus, estimated from 1981-1998 photoidentification data.

Annual survival probability of bowhead whales, Balaena mysticetus, was estimated using both Bayesian and maximum likelihood implementations of Cormack and Jolly-Seber (JS) models for capture-recapture estimation in open populations and reduced-parameter generalizations of these models. Aerial photographs of naturally marked bowheads collected between 1981 and 1998 provided the data. The marked whales first photographed in a particular year provided the initial 'capture' and 'release' of those marked whales and photographs in subsequent years the 'recaptures'. The Cormack model, often called the Cormack-Jolly-Seber (CJS) model, and the program MARK were used to identify the model with a single survival and time-varying capture probabilities as the most appropriate for these data. When survival was constrained to be one or less, the maximum likelihood estimate computed by MARK was one, invalidating confidence interval computations based on the asymptotic standard error or profile likelihood. A Bayesian Markov chain Monte Carlo (MCMC) implementation of the model was used to produce a posterior distribution for annual survival. The corresponding reduced-parameter JS model was also fit via MCMC because it is the more appropriate of the two models for these photoidentification data. Because the CJS model ignores much of the information on capture probabilities provided by the data, its results are less precise and more sensitive to the prior distributions used than results from the JS model. With priors for annual survival and capture probabilities uniform from 0 to 1, the posterior mean for bowhead survival rate from the JS model is 0.984, and 95% of the posterior probability lies between 0.948 and 1. This high estimated survival rate is consistent with other bowhead life history data.

Animals↗

Chimeric structure of omp2 of Brucella from Pacific common minke whales (Balaenoptera acutorostrata).

In the Pacific common minke whale (Balaenoptera acutorostrata ), a new variant of Brucella has been detected using the polymerase chain reaction. Detailed analysis of the porin protein genes, omp2a and omp2b from the whale Brucella showed that these two genes have some motifs in common with Atlantic marine strains in the 5'-terminal one-third region. On the other hand, the nucleotide sequences in the 3'-terminal two-thirds region of the two genes were almost identical to the respective genes of terrestrial strains. Thus, Pacific whale Brucella omp2 genes are chimeras between marine and terrestrial strains.

Animals↗

Estimates of relatedness in groups of humpback whales (Megaptera novaeangliae) on two wintering grounds of the Southern Hemisphere.

Group formation in humpback whales has been described in relation to different components of the migratory cycle, yet it is debated whether such groups represent real social bonding or ephemeral aggregations. Cooperative behaviours are exhibited during feeding activities, and it has been suggested that males may cooperate during competition for mates. Since most cooperative behaviours are expected to originate among kin, genetic relatedness represents a critical variable in the understanding of any social phenomenon, especially when cooperation cannot be confirmed unequivocally. Using an approach combining multi-locus microsatellite genotyping and several genetic relatedness estimators, we analyzed whale associations for two different wintering grounds in the Southern Hemisphere. The analyses included 648 whales sampled from 292 groups off the coast of Gabon and Northeast Madagascar, and screened for eleven microsatellite loci. Through simulations, we assessed the performance of three pairwise relatedness estimators. The individuals were molecularly sexed and their associations were investigated in the context of sex and group type. No significant association among relatives was found with the exception of mother-offspring pairs, supporting previous indications of extended maternal care. The analysis from the Gabon population also suggests that related males may avoid each other during competitive activities. Our results demonstrate that if cooperative behaviours occur on wintering grounds they are not favoured by kin selection.

Animals↗

Isolation and characterization of fin whale prolactin.

A highly purified prolactin preparation has been obtained from fin whale pituitaries by extraction with acid acetone. salt precipitation, isoelectric fractionation, and exclusion chromatography on Sephadex G-100 and ion-exchange chromatography on DEAE-CELLULOSE. Fin whale prolactin was isolated in a yield of 250 mg/kg wet weight tissue. It was found to have a molecular weight (SDS disc gel electrophoresis) of 23,600 daltons and an alpha-helix content (circular dichroism) of 50%. The amino acid composition and circular dichroism spectra were very similar to those of porcine prolactin. The partial amino acid sequence has been determined by the method of fluorescein-isothiocyanate. Fin whale prolactin was found to be 80% as potent as ovine prolactin with regard to pigeon crop-sac assay.

Amino Acid Sequence↗

Zones of impact around icebreakers affecting beluga whales in the Beaufort Sea.

A software model estimating zones of impact on marine mammals around man-made noise [C. Erbe and D. M. Farmer, J. Acoust. Soc. Am. 108, 1327-1331 (2000)] is applied to the case of icebreakers affecting beluga whales in the Beaufort Sea. Two types of noise emitted by the Canadian Coast Guard icebreaker Henry Larsen are analyzed: bubbler system noise and propeller cavitation noise. Effects on beluga whales are modeled both in a deep-water environment and a near-shore environment. The model estimates that the Henry Larsen is audible to beluga whales over ranges of 35-78 km, depending on location. The zone of behavioral disturbance is only slightly smaller. Masking of beluga communication signals is predicted within 14-71-km range. Temporary hearing damage can occur if a beluga stays within 1-4 km of the Henry Larsen for at least 20 min. Bubbler noise impacts over the short ranges quoted; propeller cavitation noise accounts for all the long-range effects. Serious problems can arise in heavily industrialized areas where animals are exposed to ongoing noise and where anthropogenic noise from a variety of sources adds up.

Animals↗

Depth-dependent acoustic features of diving sperm whales (Physeter macrocephalus) in the Gulf of Mexico.

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.

Acoustics↗

Whole-lung resonance in a bottlenose dolphin (Tursiops truncatus) and white whale (Delphinapterus leucas).

An acoustic backscatter technique was used to estimate in vivo whole-lung resonant frequencies in a bottlenose dolphin (Tursiops truncatus) and white whale (Delphinapterus leucas). Subjects were trained to submerge and position themselves near an underwater sound projector and a receiving hydrophone. Acoustic pressure measurements were made near the thorax while the subject was insonified with pure tones at frequencies from 16 to 100 Hz. Whole-lung resonant frequencies were estimated by comparing pressures measured near the subject's thorax to those measured from the same location without the subject present. Experimentally measured resonant frequencies for the white whale and dolphin lungs were 30 and 36 Hz, respectively. These values were significantly higher than those predicted using a free-spherical air bubble model. Experimentally measured damping ratios and quality factors at resonance were 0.20 and 2.5, respectively, for the white whale, and 0.16 and 3.1, respectively, for the dolphin.

Acoustic Stimulation↗

Blue whale (Balaenoptera musculus) sounds from the North Atlantic.

Sounds of blue whales were recorded from U.S. Navy hydrophone arrays in the North Atlantic. The most common signals were long, patterned sequences of very-low-frequency sounds in the 15-20 Hz band. Sounds within a sequence were hierarchically organized into phrases consisting of one or two different sound types. Sequences were typically composed of two-part phrases repeated every 73 s: a constant-frequency tonal "A" part lasting approximately 8 s, followed 5 s later by a frequency-modulated "B" part lasting approximately 11 s. A common sequence variant consisted only of repetitions of part A. Sequences were separated by silent periods averaging just over four minutes. Two other sound types are described: a 2-5 s tone at 9 Hz, and a 5-7 s inflected tone that swept up in frequency to ca. 70 Hz and then rapidly down to 25 Hz. The general characteristics of repeated sequences of simple combinations of long-duration, very-low-frequency sound units repeated every 1-2 min are typical of blue whale sounds recorded in other parts of the world. However, the specific frequency, duration, and repetition interval features of these North Atlantic sounds are different than those reported from other regions, lending further support to the notion that geographically separate blue whale populations have distinctive acoustic displays.

Animals↗

Assessing responses of humpback whales to North Pacific Acoustic Laboratory (NPAL) transmissions: results of 2001--2003 aerial surveys north of Kauai.

Eight aerial surveys were flown north of the Hawaiian island of Kauai during 2001 when the North Pacific Acoustic Laboratory (NPAL) source was not transmitting, and during 2002 and 2003 when it was. All surveys were performed during the period of peak residency of humpback whales (Feb-Mar). During 2002 and 2003, surveys commenced immediately upon cessation of a 24-h cycle of transmissions. Numbers and distribution of whales observed within 40 km of the NPAL source during 2001 (source off) were compared with those observed during 2002 and 2003 (source on). A total of 75 sightings was noted during 2001, as compared with 81 and 55 during 2002 and 2003, respectively. Differences in sighting rates (sightings/km) across years were not statistically significant. Assessment of distributional changes relied upon comparisons of three measures: (a) location depths; (b) distance from the NPAL source; and (c) distance offshore. None of the distributional comparisons revealed statistically significant differences across years. Several possible interpretations are examined: (a) whales have habituated to the NPAL signal; (b) insufficient statistical power exists in the present design to detect any effects; and (c) the effects are short-lived and become undetectable shortly after the cessation of transmissions.

Acoustics↗

Sound production by North Atlantic right whales (Eubalaena glacialis) in surface active groups.

The surface active group (SAG) is the most commonly observed surface social behavior of North Atlantic right whales. Recordings were made from 52 SAGs in the Bay of Fundy, Canada between July and September, from 1999 to 2002. The call types recorded from these groups were similar to those described previously for Southern right whales (Eubalaena australis), with six major call types being termed scream, gunshot, blow, upcall, warble, and downcall. The percentage of total calls of each call type depended on the group size and composition. The most common call type recorded was the scream call. The scream calls were produced by the focal female in a SAG. Production of other sound types can be attributed to whales other than the focal female, with gunshot and upcalls produced by males, and warble calls produced by female calves. The source levels for these sounds range from 137 to 162 dB rms re 1 ,tPa-m for tonal calls and 174 to 192 dB rms for broadband gunshot sounds.

Animal Communication↗

Click production during breathing in a sperm whale (Physeter macrocephalus).

A sperm whale (Physeter macrocephalus) was observed at the surface with above- and underwater video and synchronized underwater sound recordings. During seven instances the whale ventilated its lungs while clicking. From this observation it is inferred that click production is achieved by pressurizing air in the right nasal passage, pneumatically disconnected from the lungs and the left nasal passage, and that air flows anterior through the phonic lips into the distal air sac. The capability of breathing and clicking at the same time is unique among studied odontocetes and relates to the extreme asymmetry of the sperm whale sound-producing forehead.

Acoustics↗

Directionality of sperm whale sonar clicks and its relation to piston radiation theory.

This paper investigates the applicability to sperm whales of the theory of sound radiating from a piston. The theory is applied to a physical model and to a series of sperm whale clicks. Results show that wave forms of off-axis signals can be reproduced by convolving an on-axis signal with the spatial impulse response of a piston. The angle of a recorded click can be estimated as the angle producing the spatial impulse response that gives the best match with the observation when convolved with the on-axis wave form. It is concluded that piston theory applies to sperm whale sonar click emission.

Animals↗

Acoustic detection of North Atlantic right whale contact calls using the generalized likelihood ratio test.

This paper addresses the problem of passive acoustic detection of contact calls produced by the highly endangered North Atlantic right whale Eubalaena glacialis. The proposed solution is based on using a generalized likelihood ratio test detector of polynomial-phase signals with unknown amplitude and polynomial coefficients observed in the presence of locally stationary Gaussian noise. The closed form representation for a minimal sufficient statistic is derived and a realizable detection scheme is developed. The receiver operation characteristic curves are calculated using empirical data recordings containing known right whale calls. The curves demonstrate that the proposed detector provides superior detection performance as compared with other known detection techniques for northern right whale contact calls.

Algorithms↗

Simulating the effect of high-intensity sound on cetaceans: modeling approach and a case study for Cuvier's beaked whale (Ziphius cavirostris).

A finite element model is formulated to study the steady-state vibration response of the anatomy of a whale (Cetacea) submerged in seawater. The anatomy was reconstructed from a combination of two-dimensional (2D) computed tomography (CT) scan images, identification of Hounsfield units with tissue types, and mapping of mechanical properties. A partial differential equation model describes the motion of the tissues within a Lagrangean framework. The computational model was applied to the study of the response of the tissues within the head of a neonate Cuvier's beaked whale Ziphius cavirostris. The characteristics of the sound stimulus was a continuous wave excitation at 3500 Hz and 180 dB re: 1 microPa received level, incident as a plane wave. We model the beaked whale tissues embedded within a volume of seawater. To account for the finite dimensions of the computational volume, we increased the damping for viscous shear stresses within the water volume, in an attempt to reduce the contribution of waves reflected from the boundaries of the computational box. The mechanical response of the tissues was simulated including: strain amplitude; dissipated power; and pressure. The tissues are not likely to suffer direct mechanical or thermal damage, within the range of parameters tested.

Acoustics↗

Tracking fin whale calls offshore the Galicia Margin, North East Atlantic Ocean.

Data recorded during a temporary deployment of ocean bottom seismometers (OBSs) are used in this study to monitor the presence of fin whales around the array. In the summer of 2003, ten OBSs were placed 250 km from the NW coast of Iberia in the Galicia Margin, NE Atlantic Ocean for a period of one month. The recorded data set provided a large variety of signals, including fin whale vocalizations identified by their specific acoustic signature. The use of a dense array of seafloor receivers allowed investigation into the locations and tracks of the signal-generating whales using a seismological hypocentral location code. Individual pulses of different sequences have been chosen to study such tracks. Problems related to the correct identification of pulses, discrimination between direct and multiple arrivals, and the presence of more than one individual have been considered prior to location. Fin calls were concentrated in the last two weeks of the deployment and the locations were spread around the area covered by the array. These results illustrate that, besides its classical seismological aim, deployment of semipermanent seafloor seismic arrays can also provide valuable data for marine mammal behavior studies.

Algorithms↗

Demonstration of adaptation in beluga whale echolocation signals.

The echolocation signals of the same beluga whale (Delphinapterus leucas) were measured first in San Diego Bay, and later in Kaneohe Bay, Oahu, Hawaii. The ambient noise level in Kaneohe Bay is typically 12-17 dB greater than in San Diego Bay. The whale demonstrated the adaptiveness of its biosonar by shifting to higher frequencies and intensities after it was moved to Kaneohe. In San Diego, the animal emitted echolocation signals with peak frequencies between 40 and 60 kHz, and bandwidths between 15 and 25 kHz. In Kaneohe, the whale shifted its signals approximately an octave higher in frequencies with peak frequencies between 100 and 120 kHz, and bandwidths between 20 and 40 kHz. Signal intensities measured in Kaneohe were up to 18 dB higher than in San Diego. The data collected represent the first quantitative evidence of the adaptive capability of a cetacean biosonar system.

Adaptation, Physiological↗

Reactions of bowhead whales, Balaena mysticetus, to seismic exploration in the Canadian Beaufort Sea.

The behavioral reactions of bowhead whales to distant seismic vessels not under our control, a controlled approach by a seismic vessel, and controlled tests with a single airgun were observed. On 21 occasions in the summers of 1980-84, general activities of bowheads exposed to pulses of underwater noise (107-158 dB re: 1 mu Pa) from seismic vessels 6-99 km away were observed. Activities were indistinguishable from those without seismic noise; there was no detectable avoidance. Hints of subtle changes in surfacing, respiration, and diving behavior were unconfirmed, but were consistent with reactions to stronger noise pulses from closer seismic boats. In a test with a full-scale seismic boat (30 airguns totaling 471, source level 248 dB re: 1 mu Pa, closest point of approach = 1 1/2 km), bowheads began to orient away when the airgun array began to fire 7 1/2 km away. However, some whales continued apparent near-bottom feeding until the vessel was 3 km away. Whales were displaced by about 2 km. Reactions were not much stronger than those to any conventional vessel. Tests with one 0.66-1 airgun showed that some bowheads move away from sources of strong seismic impulses even in the absence of boat noise, and that bowheads can detect the direction from which seismic impulses arrive. In general, bowheads exhibit avoidance reactions when they receive seismic pulses stronger than about 160 dB re: 1 mu Pa. Evidence of reactions to lower received levels remains inconclusive.

Animals↗

Sounds, source levels, and associated behavior of humpback whales, southeast Alaska.

Humpback whales in Southeast Alaskan waters produced five categories of sounds: moans, grunts, pulse trains, blowhole-associated sounds, and surface impacts. Frequencies (Hz) of moans and grunts were 20-1900. Major energy in low-frequency pulse trains was in a band of 25-80 Hz with pulse duration of 300-400 ms. Blowhole-associated sounds, recorded as transiting whales encountered one another, were of two types: shrieks, 555-2000 Hz, and trumpetlike horn blasts with fundamental at 414 Hz (median). Pulses and spread spectrum noise were associated with gas bubble formation and explosive bursts, respectively, in connection with spiral feeding maneuvers. Surface impacts resulted from fluke or flipper slaps in sequences of 3-21 sounds. Source levels ranged from 162 (low-frequency pulse trains) to 192 dB (surface impacts), re: 1 microPa, 1 m. Songs, commonly heard on winter breeding grounds, were absent from our recordings. Feeding and perhaps certain other whale activities can be monitored based on sound production.

Animals↗