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Demography of the endangered North Atlantic right whale.

Northern right whales (Eubalaena glacialis) were formerly abundant in the northwestern Atlantic, but by 1900 they had been hunted to near extinction. After the end of commercial whaling the population was thought to be recovering slowly; however, evidence indicates that it has been declining since about 1990 (ref. 1). There are now fewer than 300 individuals, and the species may already be functionally extinct owing to demographic stochasticity or the difficulty of females locating mates in the vast Atlantic Ocean (Allee effect). Using a data set containing over 10,000 sightings of photographically identified individuals we estimated trends in right whale demographic parameters since 1980. Here we construct, using these estimates, matrix population models allowing us to analyse the causes of right whale imperilment. Mortality has increased, especially among mother whales, causing declines in population growth rate, life expectancy and the mean lifetime number of reproductive events between the period 1980-1995. Increased mortality of mother whales can explain the declining population size, suggesting that the population is not doomed to extinction as a result of the Allee effect. An analysis of extinction time shows that demographic stochasticity has only a small effect, but preventing the deaths of only two female right whales per year would increase the population growth rate to replacement level.

Animals↗

Buoyant balaenids: the ups and downs of buoyancy in right whales.

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.

Animals↗

Sperm whale behaviour indicates the use of echolocation click buzzes "creaks" in prey capture.

During foraging dives, sperm whales (Physeter macrocephalus) produce long series of regular clicks at 0.5-2 s intervals interspersed with rapid-click buzzes called "creaks". Sound, depth and orientation recording Dtags were attached to 23 whales in the Ligurian Sea and Gulf of Mexico to test whether the behaviour of diving sperm whales supports the hypothesis that creaks are produced during prey capture. Sperm whales spent most of their bottom time within one or two depth bands, apparently feeding in vertically stratified prey layers. Creak rates were highest during the bottom phase: 99.8% of creaks were produced in the deepest 50% of dives, 57% in the deepest 15% of dives. Whales swam actively during the bottom phase, producing a mean of 12.5 depth inflections per dive. A mean of 32% of creaks produced during the bottom phase occurred within 10 s of an inflection (13x more than chance). Sperm whales actively altered their body orientation throughout the bottom phase with significantly increased rates of change during creaks, reflecting increased manoeuvring. Sperm whales increased their bottom foraging time when creak rates were higher. These results all strongly support the hypothesis that creaks are an echolocation signal adapted for foraging, analogous to terminal buzzes in taxonomically diverse echolocating species.

Acoustics↗

Deep-diving foraging behaviour of sperm whales (Physeter macrocephalus).

1. Digital tags were used to describe diving and vocal behaviour of sperm whales during 198 complete and partial foraging dives made by 37 individual sperm whales in the Atlantic Ocean, the Gulf of Mexico and the Ligurian Sea. 2. The maximum depth of dive averaged by individual differed across the three regions and was 985 m (SD = 124.3), 644 m (123.4) and 827 m (60.3), respectively. An average dive cycle consisted of a 45 min (6.3) dive with a 9 min (3.0) surface interval, with no significant differences among regions. On average, whales spent greater than 72% of their time in foraging dive cycles. 3. Whales produced regular clicks for 81% (4.1) of a dive and 64% (14.6) of the descent phase. The occurrence of buzz vocalizations (also called 'creaks') as an indicator of the foraging phase of a dive showed no difference in mean prey capture attempts per dive between regions [18 buzzes/dive (7.6)]. Sperm whales descended a mean of 392 m (144) from the start of regular clicking to the first buzz, which supports the hypothesis that regular clicks function as a long-range biosonar. 4. There were no significant differences in the duration of the foraging phase [28 min (6.0)] or percentage of the dive duration in the foraging phase [62% (7.3)] between the three regions, with an overall average proportion of time spent actively encountering prey during dive cycles of 0.53 (0.05). Whales maintained their time in the foraging phase by decreasing transit time for deeper foraging dives. 5. Similarity in foraging behaviour in the three regions and high diving efficiencies suggest that the success of sperm whales as mesopelagic predators is due in part to long-range echolocation of deep prey patches, efficient locomotion and a large aerobic capacity during diving.

Acoustics↗

Low-frequency whale and seismic airgun sounds recorded in the mid-Atlantic Ocean.

Beginning in February 1999, an array of six autonomous hydrophones was moored near the Mid-Atlantic Ridge (35 degrees N-15 degrees N, 50 degrees W-33 degrees W). Two years of data were reviewed for whale vocalizations by visually examining spectrograms. Four distinct sounds were detected that are believed to be of biological origin: (1) a two-part low-frequency moan at roughly 18 Hz lasting 25 s which has previously been attributed to blue whales (Balaenoptera musculus); (2) series of short pulses approximately 18 s apart centered at 22 Hz, which are likely produced by fin whales (B. physalus); (3) series of short, pulsive sounds at 30 Hz and above and approximately 1 s apart that resemble sounds attributed to minke whales (B. acutorostrata); and (4) downswept, pulsive sounds above 30 Hz that are likely from baleen whales. Vocalizations were detected most often in the winter, and blue- and fin whale sounds were detected most often on the northern hydrophones. Sounds from seismic airguns were recorded frequently, particularly during summer, from locations over 3000 km from this array. Whales were detected by these hydrophones despite its location in a very remote part of the Atlantic Ocean that has traditionally been difficult to survey.

Animal Communication↗

Click rates and silences of sperm whales at Kaikoura, New Zealand.

Analysis of the usual click rates of sperm whales (Physeter macrocephalus) at Kaikoura, New Zealand, confirms the potential for assessing abundance via "click counting." Usual click rates over three dive cycles each of three photographically identified whales showed that 5 min averages of usual click rate did not differ significantly within dives, among dives of the same whale or among whales. Over the nine dives (n= 13 728 clicks) mean usual click rate was 1.272 clicks s(-1) (95% CI= 0.151). On average, individual sperm whales at Kaikoura spent 60% of their time usual clicking in winter and in summer. There was no evidence that whale identity or stage of the dive recorded affects significantly the percentage of time spent usual clicking. Differences in vocal behavior among sperm whale populations worldwide indicate that estimates of abundance that are based on click rates need to based on data from the population of interest, rather than from another population or some global average.

Acoustics↗

Sounds and source levels from bowhead whales off Pt. Barrow, Alaska.

Sounds were recorded from bowhead whales migrating past Pt. Barrow, AK, to the Canadian Beaufort Sea. They mainly consisted of various low-frequency (25- to 900-Hz) moans and well-defined sound sequences organized into "song" (20-5000 Hz) recorded with our 2.46-km hydrophone array suspended from the ice. Songs were composed of up to 20 repeated phrases (mean, 10) which lasted up to 146 s (mean, 66.3). Several bowhead whales often were within acoustic range of the array at once, but usually only one sang at a time. Vocalizations exhibited diurnal peaks of occurrence (0600-0800, 1600-1800 h). Sounds which were located in the horizontal plane had peak source spectrum levels as follows--44 moans: 129-178 dB re: 1 microPa, 1 m (median, 159); 3 garglelike utterances: 152, 155, and 169 dB; 33 songs: 158-189 dB (median, 177), all presumably from different whales. Based on ambient noise levels, measured total propagation loss, and whale sound source levels, our detection of whale sounds was theoretically noise-limited beyond 2.5 km (moans) and beyond 10.7 km (songs), a model supported by actual localizations. This study showed that over much of the shallow Arctic and sub-Arctic waters, underwater communications of the bowhead whale would be limited to much shorter ranges than for other large whales in lower latitude, deep-water regions.

Alaska↗

Time and frequency domain characteristics of sperm whale clicks.

Regular clicks from diving sperm whales, both large bull males and smaller females, were recorded in deep oceanic water off the Azores and subsequently sampled to computer disks for digital analysis. A total of 8540 clicks were marked and analyzed. Simple temporal analysis of the interclick intervals during feeding dives revealed mean click rates for male sperm whales of 1.1713 s-1 and 1.9455 s-1 for females. Fourier analysis showed distinctive peaks in the spectra of bull male sperm whales at 400 Hz and 2 kHz which were stable over extended periods of up to 20 mins. The clicks contained higher frequency components with energy ranging up to at least 12 kHz but not concentrated at any sharply defined frequency. The clicks of smaller female sperm whales showed similar spectral peaks, shifted to 1.2 and 3 kHz, respectively, but these peaks were less pronounced than those in the male click spectra and less stable with time. Higher frequencies were also present up to at least 15 kHz. The previously reported multiple pulse structure of sperm whale clicks is confirmed, but digital filtering reveals this structure to be frequency dependent. Analysis using the short-time Fourier transform confirms the complex time-frequency structure of individual clicks. The frequencies at which the multiples emerge in male and female clicks supports the idea of air cavities in the sperm whale head acting as sound reflectors, although the magnitude of the second pulse at high frequencies suggests some form of off axis distortion. It is also possible that air cavity resonance in the head of the sperm whale may act to reinforce the high-frequency components of the click, and that such components may have superior range and resolution performance in terms of echolocation.

Animal Communication↗

Blue and fin whales observed on a seafloor array in the northeast pacific.

Calling blue and fin whales have been tracked using relative travel times and amplitudes from both direct and multipath arrivals to a seafloor array of seismometers. Calls of three fin whales swimming in the same general direction, but several kilometers apart, are believed to represent communication between the whales because of signature differences in call character, an alternating call pattern, and coordination of call and respiration times. Whale call tracks, call patterns, call character, and swimming speeds were examined during periods with and without the presence of noise. Noise sources included airguns, when the whales were subject to sound levels of up to 143 dB P-P (peak-to-peak) re: 1 microPa over the 10 to 60-Hz band, and transits of merchant ships, when the whales received continuous levels up to 106 dB rms re: 1 microPa over the 10 to 60-Hz band (115 dB P-P). Whale responses associated with these noises remain arguable.

Animals↗

Swimming gaits, passive drag and buoyancy of diving sperm whales Physeter macrocephalus.

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.

Acceleration↗

The mechanical properties of fin whale arteries are explained by novel connective tissue designs.

The aortic arch and the descending aorta in the fin whale (Balaenoptera physalus) are structurally and mechanically very different from comparable vessels in other mammals. Although the external diameter of the whale's descending thoracic aorta (approximately 12 cm) is similar to that predicted by scaling relationships for terrestrial mammals, the wall thickness:diameter ratio in the whale (0.015) is much smaller than the characteristic value for other mammals (0.05). In addition, the elastic modulus of the thoracic aorta (12 MPa at 13 kPa blood pressure) is about 30 times higher than in other mammals. In contrast, the whale's aortic arch has a wall thickness/diameter ratio (0.055) and an elastic modulus (0.4 MPa) that are essentially identical to those for other mammals. However, the aortic arch is unusual in that it can be deformed biaxially to very large strains without entering a region of high stiffness caused by the recruitment of fully extended collagen fibres. Chemical composition studies indicate that the elastin:collagen ratio is high in the aortic arch (approximately 2:1) and that this ratio falls in the thoracic (approximately 1:2) and abdominal (approximately 1:3) aortas, but the magnitude of the change in composition does not account for the dramatic difference in mechanical properties. This suggests that there are differences in the elastin and collagen fibre architecture of these vessels. The descending aorta contains dense bands of tendon-like, wavy collagen fibres that run in the plane of the arterial wall, forming a fibre-lattice that runs in parallel to the elastin lamellae and reinforces the wall, making it very stiff. The aortic arch contains a very different collagen fibre-lattice in which fibres appear to have a component of orientation that runs through the thickness of the artery wall. This suggests that the collagen fibres may be arranged in series with elastin-containing elements, a difference in tissue architecture that could account for both the lower stiffness and the extreme extensibility of the whale's aortic arch. Thus, both the structure and the mechanical behaviour of the lamellar units in the aortic arch and aorta of the whale have presumably been modified to produce the unusual mechanical and haemodynamic properties of the whale circulation.

Animals↗

Hearing and whistling in the deep sea: depth influences whistle spectra but does not attenuate hearing by white whales (Delphinapterus leucas) (Odontoceti, Cetacea).

Hearing is attenuated in the aerial ear of humans and other land mammals tested in pressure chambers as a result of middle ear impedance changes that result from increased air density. We tested the hypothesis, based on recent middle ear models, that increasing the density of middle ear air at depth might attenuate whale hearing. Two white whales Delphinapterus leucas made dives to a platform at a depth of 5, 100, 200 or 300 m in the Pacific Ocean. During dives to station on the platform for up to 12 min, the whales whistled in response to 500 ms tones projected at random intervals to assess their hearing threshold at each depth. Analysis of response whistle spectra, whistle latency in response to tones and hearing thresholds showed that the increased hydrostatic pressure at depth changed each whale's whistle response at depth, but did not attenuate hearing overall. The finding that whale hearing is not attenuated at depth suggests that sound is conducted through the head tissues of the whale to the ear without requiring the usual ear drum/ossicular chain amplification of the aerial middle ear. These first ever hearing tests in the open ocean demonstrate that zones of audibility for human-made sounds are just as great throughout the depths to which these whales dive, or at least down to 300 m.

Animals↗

The pattern of ovarian development in the prepubertal antarctic minke whale (Balaenoptera bonaerensis).

This study describes the morphological and morphometrical changes associated with prepubertal ovarian development in the Antarctic minke whale (Balaenoptera bonaerensis). Ovaries were harvested from 94 immature minke whales caught in the Antarctic Ocean during the summer feeding season (December-March). Notable differences in ovarian size and morphology were found among animals. Up to 10 folds difference in ovarian weight was found among prepubertal whales of similar body size. During the prepubertal period, ovaries grew slowly and approximately doubled their weight. The morphologies of right and left ovaries were almost identical while the growth of the ovary appears to occur preferentially on the right side. The most striking morphological feature was numerous small antral follicles less than 5 mm in diameter found in ovaries of younger immature whales. The occurrence of these ovaries was highest in whales less than 6 m long and gradually decreased as body length increased. In larger whales, the occurrence of ovaries with a smaller number of follicles up to 10 mm and thick tunica albuginea increased. Thus, the ovary of the Antarctic minke whale experiences bursts of small follicular development during the early prepubertal period before becoming a more developed ovary with fewer but larger follicles, and thick tunica albuginea.

Animals↗

Organochlorine contaminant levels in Eskimo harvested bowhead whales of arctic Alaska.

Organochlorine (OC) levels in liver and blubber of 20 bowhead whales (Balaena mysticetus) collected during the Eskimo subsistence harvest at Barrow (Alaska, USA) in 1992 and 1993 are presented. Liver sum DDT (lipid weight) was significantly greater in male whales than in females. Most of the organochlorines measured were at higher levels in longer (older) than in shorter (younger) males. For female bowhead whales, hexachlorobenzene and lipid levels decreased and other OC levels did not change significantly with increasing length. Most organochlorine contaminants have low concentrations in tissues of the bowhead whale compared to concentrations in tissues of other cetaceans, especially Odontocetes. Based on allowable daily intakes (ADI) levels established by the Canadian Northern Contaminants Program (Ottawa, Ontario, Canada) "safe" levels of blubber to consume were calculated. Chlordane levels in bowhead whale blubber results in the most restrictive consumption amount (50 g blubber/day). We expect no adverse effects related to these organochlorine contaminants to occur in bowhead whales or in consumers of their tissues. However, investigation of low level chronic exposure effects and a more rigorous assessment of histopathology, biomarkers, and immune status in the bowhead whale would be required to conclude "no effect" with more certainty.

Adipose Tissue↗

Cardiomyopathy in stranded pygmy and dwarf sperm whales.

Necropsy and histologic examinations were performed in 23 pygmy sperm whales (Kogia breviceps) and 6 dwarf sperm whales (Kogia simus) that had been stranded singly or in cow-calf pairs along the southeastern coastline of the United States. At necropsy, the gross findings in the adult whales included pale, flabby right ventricles. Microscopically, lesions in the hearts of the whales were characterized by moderate to extensive myocellular degeneration, atrophy, and fibrosis. Similar changes were not seen in 5 of 6 sexually immature whales or in the whale calves. Hepatic changes were consistent with heart failure. The cause of the myocardial lesions was not determined. The systemic effects of failing myocardium probably were a major reason for the stranding of the adult whales.

Age Factors↗

Titration behavior of individual tyrosine residues of myoglobins from sperm whale, horse, and red kangaroo.

The titration behavior of individual tyrosine residues of myoglobins has been studied by observing the pH dependence of the chemical shifts of Czeta and Cgamma of these residues in natural abundance of 13C Fourier transform NMR spectra (at 15.18 MHz, in 20-mm sample tubes, at 37 degrees) of cyanoferrimyoglobins from sperm whale, horse, and red kangaroo. A comparison of the pH dependence of the spectra of the three proteins yielded specific assignments for the resonance of Tyr-151 (sperm whale) and Tyr-103 (sperm whale and horse). Selective proton decoupling yielded specific assignments for Czeta of Tyr-146 of the cyanoferrimyoglobins from horse and kangaroo, but not the corresponding assignment for sperm whale. The pH dependence of the chemical shifts indicated that only Tyr-151 and Tyr-103 are titratable tyrosine residues. Even at pH 12, Tyr-146 did not begin to titrate. The titration behavior of C zeta and Cgamma of Tyr-151 of sperm whale cyanoferrimyoglobin yielded a single pK value of 10.6. The pH dependence of the chemical shift of each of the resonances of Tyr-103 of the cyanoferrimyoglobins from horse and sperm whale could not be fitted with the use of a single pK value, but was consistent with two pK values (about 9.8 and 11.6). Furthermore, the resonances of Czeta and Cgamma of Tyr-103 broadened at high pH. The titration behavior of the tyrosines of sperm whale carbon monoxide myoglobin and horse ferrimyoglobin was also examined. A comparison of all the experimental results indicated that Tyr-151 is exposed to solvent, Tyr-146 is not exposed, and Tyr-103 exhibits intermediate behavior. These results for myoglobins in solution are consistent with expectations based on the crystal structure.

Animals↗

Accumulation, metabolism, and food-chain transfer of chlorinated and brominated contaminants in subadult white whales (Delphinapterus leucas) and narwhals (Monodon monoceros) from Svalbard, Norway.

The concentrations and patterns of polychlorinated biphenyls (PCBs), chlorinated pesticides, and polybrominated diphenyl ethers (PBDEs) were studied in white whales (Delphinapterus leucas) and narwhals (Monodon monoceros) from Svalbard, Norway. In addition, their main food items were included in the study. In the whales, a broad range of pollutants was found in relatively high concentrations. PCBs and pesticides were approximately 3000 and 8000 ng/g lipid, respectively, for white whales and three times higher for narwhals. PBDEs 47 were approximately 70 ng/g lipid for white whales and 170 ng/g lipid for narwhals. Compared with other marine mammals from the same area, contaminant levels are among the highest levels ever measured. These high levels are likely in part because of a decreased capacity to metabolize contaminants. Metabolic indices indicated that most compounds accumulate to the same degree in white whales and narwhals, but for some toxaphenes and chlordanes, narwhals might have a decreased metabolism and consequently a higher accumulation. The three-times-higher contaminant levels in blubber of narwhals was further explained by substantially higher contaminant levels in their more benthic diet. The high levels and broad pattern of accumulating pollutants make white whales and narwhals excellent indicators for a wide range of contaminants in the Arctic.

Animals↗

Ontogenesis of the sperm whale brain.

The development of the sperm whale brain (Physeter macrocephalus) was investigated in 12 embryos and early fetuses to obtain a better understanding of the morphological and physiological adaptations in this most exotic cetacean concerning locomotion, deep diving, and orientation. In male adult sperm whales, the average absolute brain mass and the relative size of the telencephalic hemisphere are the largest within the mammalia, whereas the ratio of the brain mass to the total body mass is one of the smallest. In the early sperm whale fetus, the rostral part of the olfactory system (olfactory nerves and bulbs) is lost, whereas the nervus terminalis seems to persist. Several components of the limbic system show signs of regression (hippocampus, fornix, mamillary body). In contrast, some components of the auditory system (trapezoid body, inferior colliculus) show marked enlargement in the early fetal period, thereby reflecting their dominant position in the adult. The cerebellum and pons grow slower than in most smaller toothed whales. The pyramidal tract develops poorly (reduction of the limbs), whereas marked growth of the striatum and inferior olive may be related to the animal's locomotion via trunk and tail. In the early fetal period, the trigeminal, vestibulocochlear, and facial nerves are the dominant cranial nerves (besides the vagus nerve). Whereas the number of axons in the vestibulocochlear nerve is high in adult, toothed whales and their diameters are considerable, the trigeminal nerve of the sperm whale may be the thickest of all cranial nerves and has the largest number of axons (innervation of the huge forehead region). A similar situation seems to exist for the facial nerve: It innervates the blowhole musculature that surrounds the very large spermaceti organ and melon (generation and emission of sonar clicks).

Animals↗