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In vitro digestibility of different prey species of minke whales (Balaenoptera acutorostrata).

Information on diet composition, daily energy expenditure, energy storage and the utilization of energy in the prey are important factors when evaluating the food consumption of minke whales (Balaenoptera acutorostrata) during their summer stay in northern waters. The purpose of the present study was in this context to obtain information on the digestible energy (DE) of different prey selected by minke whales. An in vitro three-stage digestion technique, simulating the different compartments of the digestive system, has been developed. The initial step simulated the anaerobic microbial fermentation of substrate in the forestomach. The next stage included the addition of pepsin (EC 3.4.23.1)-HCl, simulating ventricle enzymic decomposition, and finally, in the third step, fresh extract from duodenal contents was used to simulate enzymic intestinal degradation of the remaining components of the food. The inoculum was normally obtained from animals which had recently eaten the prey to be tested. In such tests we obtained a dry matter disappearance (DMD) and a DE for herring (Clupea harengus) of 80.4 (SD 5.0)% (n 18) and 92.1 (SD 3.7)% (n 16) respectively, and a DMD of krill (Thysanoessa sp.) of 83.4 (SD 4.9)% (n 6). The DMD of krill was reduced to 73.8 (SD 7.3)% (n 8) while the DE was 70.6 (SD 10.4)% (n 7) when inoculum from whales which had recently eaten cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) was used. These results indicate a high digestibility of the most common species of prey in these animals, and also that the whales have little difficulty in changing from one prey species to another.

Animals↗

Digestibility of krill (Euphausia superba and Thysanoessa sp.) in minke whales (Balaenoptera acutorostrata) and crabeater seals (Lobodon carcinophagus).

Apparent digestible efficiency (% DE) was studied by use of dietary Mn as an inert marker, in minke whales (Balaenoptera acutorostrata) and crabeater seals (Lobodon carcinophagus) which had been eating krill. Median % DE in minke whales (n5) eating krill of the genus Thysanoessa sp. (energy density (ED) 23.8 kJ/g) was 93 (range 87-93). Median % DE in crabeater seals (n6) eating krill of the species Euphausia superba (ED 20.8 kJ/g) was 84 (range 79-85), which is significantly lower than the % DE of krill in minke whales (P = 0.008). Since the chemical composition in E. superba and in Thysanoessa sp. is similar, it is suggested that the complex multi-stomached system of minke whales, which contains both chitinase (EC 3.2.1.14)-producing as well as several other types of bacteria, is superior to the single-stomached system of crabeater seals with regard to krill digestion. It is worth noting, however, that the % DE of krill in the crabeater seal is still very high.

Animals↗

Cadmium, copper, lead, and zinc in five toothed whale species of the Mediterranean Sea.

The cadmium, lead, copper, and zinc levels were measured in six organs (lung, liver, kidney, skin, muscle, and bone) from 18 specimens of toothed whales, belonging to five species, found stranded along the Corsican coast between November 1993 and December 1998. The five species examined were the bottlenose dolphin Tursiops truncatus, the common dolphin Delphinus delphis, the striped dolphin Stenella coeruleoalba, the pilot whale Globicephala melas, and the Risso's dolphin Grampus griseus. The values obtained demonstrate that there is a great variability in the accumulation of toxic metals. A comparison of the values with those obtained in other geographical regions also demonstrates an extensive variability in metal levels. The presence of metals in the stomach contents of three of the individuals studied shows that the food source is responsible for a significant input of pollutants to the whales. Measured pollutant levels in whales are thus the result of an accumulation occurring throughout the animal's life.

Animals↗

The influence of maternal lineages on social affiliations among humpback whales (Megaptera novaeangliae) on their feeding grounds in the southern gulf of Maine.

Humpback whales on their feeding grounds in the Gulf of Maine typically form fluid fission/fusion groups of two to three individuals characterized by noncompetitive and, at times, cooperative behavior. Here we test the hypothesis that, despite the apparent absence of close kinship bonds, the fluid associations between feeding whales are influenced by "maternal lineages" as represented by mtDNA haplotypes. Using skin samples collected with a biopsy dart, variation in the hypervariable segment of the mtDNA control region identified 17 unique haplotypes among 159 individually identified whales from the southern Gulf of Maine. The haplotypes of a further 143 individuals were inferred from known direct maternal (cow-calf) relationships. The frequencies of associations among these 302 individuals were calculated from 21,617 sighting records collected from 1980 to 1995, excluding associations between a cow and her dependent calf. For groups of two where the haplotypes of both individuals were known (n = 3,151), individuals with the same haplotype were together significantly more often (26%) than expected by random association (20%). To account for different group sizes and associations with individuals of unknown haplotype and sex, we used Monte Carlo simulations to test for nonrandom associations in the full data set, as well as known female-only (n = 1,512), male-only (n = 730), and mixed-sex (n = 2,745) groups. Within-haplotype associations were significantly more frequent than expected at random for all groups (P = .002) and female-only groups (P = .011) but not male-only groups, while mixed-sex groups approached significance (P = .062). A Mantel test of individual pairwise association indices and haplotype identity confirmed that within-haplotype associations were more frequent than expected for all sex combinations except male-male associations, with females forming within-haplotype associations 1.7 times more often than expected by random assortment. Partial matrix correlations and permutation analyses indicated that the skew toward within-haplotype associations could not be accounted for by short-term temporal co-occurrence or fine-scale spatial distributions of individuals with shared haplotypes. While the mechanism by which individuals with a common mtDNA haplotype assort remains unknown, our results strongly suggest an influence of maternal lineages on the social organization of humpback whales within a regional feeding ground.

Animals↗

1H-NMR comparative study of the active site in shark (Galeorhinus japonicus), horse, and sperm whale deoxy myoglobins.

1H-NMR spectra of deoxy myoglobins (Mbs) from shark (Galeorhinus japonicus), horse, and sperm whale have been studied to gain insights into their active site structure. It has been demonstrated for the first time that nuclear Overhauser effect (NOE) can be observed between heme peripheral side-chain proton resonances of these paramagnetic complexes. Val-E11 methyl and His-F8 C delta H proton resonances of these Mbs were also assigned from the characteristic shift and line width. The hyperfine shift of the former resonance was used to calculate the magnetic anisotropy of the protein. The shift analysis of the latter resonance, together with the previously assigned His-F8 N delta H proton resonance, revealed that the strain on the Fe-N epsilon bond is in the order horse Mb approximately whale Mb < shark Mb and that the hydrogen bond strength of the His-F8 N delta H proton to the main-chain carbonyl oxygen in the preceding turn of the F helix is in the order shark Mb < horse Mb < whale Mb. Weaker Feporphyrin interaction in shark Mb was manifested in a smaller shift of the heme methyl proton resonance and appears to result from distortion of the coordination geometry in this Mb. Larger strain on the Fe-N epsilon bond in shark Mb should be to some extent attributed to its lowered O2 affinity (P50 = 1.1 mmHg at 20 degrees C), compared to whale and horse Mbs.

Animals↗

Population characteristics of DNA fingerprints in humpback whales (Megaptera novaeangliae).

Humpback whales exhibit a remarkable social organization that is characterized by seasonal long-distance migration (> 10,000 km/year) between summer feeding grounds in high latitudes and winter calving and breeding grounds in tropical or near-tropical waters. All populations are currently considered endangered as a result of intensive commercial exploitation during the last 200 years. Using three hypervariable minisatellite DNA probes (33.15, 3'HVR, and M13) originally developed for studies of human genetic variation, we examined genetic variation within and among three regional subpopulations of humpback whales from the North Pacific and one from the North Atlantic oceans. Analysis of DNA extracted from skin tissues collected by biopsy darting from free-ranging whales revealed considerable variation in each subpopulation. The extent of this variation argues against a recent history of inbreeding among humpback whales as a result of nineteenth- and twentieth-century hunting. A canonical variate analysis suggested a relationship between scaled genetic distance, based on similarities of DNA fingerprints, and geographic distance (i.e., longitude of regional subpopulation). Significant categorical differences were found between the two oceanic populations using a multivariate analysis of variance (MANOVA) with a modification of the Mantel nonparametric permutation test. The relationship between DNA fingerprint similarities and geographic distance suggests that nuclear gene flow between regional subpopulations within the North Pacific is restricted by relatively low rates of migratory interchange between breeding grounds or assortative mating on common wintering grounds.

Animals↗

Discovery of a blue whale feeding and nursing ground in southern Chile.

After the extensive exploitation that reduced the Southern Hemisphere blue whale (Balaenoptera musculus) populations to less than 3% of its original numbers, studies on its recovery have been compounded by the inaccessibility of most populations and the extensive migrations between low and high latitudes, thus ensuring that knowledge about blue whale ecology and status remains limited. We report the recent discovery of, arguably, the most important blue whale feeding and nursing ground known to date in the Southern Hemisphere, which is located near the fjords off southern Chile. Through aerial and marine surveys (n = 7) 47 groups, comprising 153 blue whales including at least 11 mother-calf pairs, were sighted during the austral summer and early autumn of 2003. The implications of this discovery on the biological understanding and conservation of this endangered species are discussed.

Animal Migration↗

Populations genetic analysis of nuclear and mitochondrial loci in skin biopsies collected from central and northeastern North Atlantic humpback whales (Megaptera novaeangliae): population identity and migratory destinations.

It has been speculated that humpback whales, Megaptera novaeangliae, from the northeastern North Atlantic breed in tropical waters off the coast of West Africa and therefore that they represent a separate breeding population from that which winters in the West Indies. We determined the genotype at six microsatellite loci as well as the sequence of the first 288 nucleotides in the mitochondrial control region of 133 skin biopsies collected from humpback whales in the central North Atlantic (Iceland and Jan Mayen) and the northeastern North Atlantic (Bear Island and the northern coast of Norway). We detected no significant deviations from Hardy-Weinberg proportions nor any differences in genotype frequencies between localities at the nuclear loci. However, the mitochondrial analyses revealed two distinct matrilineal aggregations: the central and the northeastern North Atlantic. Our findings were not compatible with the idea of a separate eastern North Atlantic breeding ground unless one has been established recently. The proposed alternative hypothesis of a common North Atlantic panmictic population (wintering primarily in the West Indies) in which individual whales display maternally directed site-fidelity to specific feeding grounds was supported by re-sightings of two northeastern North Atlantic humpback whales in the West Indies.

Animals↗

Phenotypic and molecular characterization of a Brucella strain isolated from a minke whale (Balaenoptera acutorostrata).

Isolation of Brucella spp. in marine mammals has been reported during the past several years. A Brucella strain from the spleen and liver of a minke whale (Balaenoptera acutorostrata) was isolated. Conventional typing methods indicated that this isolate was related to the genus Brucella but did not match the profiles of any known Brucella species or biovar. Successful PCR amplification of the Brucella rrs-rrl spacer sequence and of the insertion sequence IS6501 also indicated that the minke whale strain was related to the genus Brucella. In addition, the rrs gene of this strain shared a very high degree of nucleotide identity (>98%) with published Brucella spp. rrs sequences. However, RFLP studies using an IS6501-specific probe showed a unique profile for this strain in comparison with the profiles of the six known Brucella species. Moreover, analysis of the omp2 locus by PCR-RFLP, by Southern hybridization using omp2a- and omp2b-specific probes, and by DNA sequencing showed that the minke whale isolate possesses two copies of the omp2b gene instead of one omp2a and one omp2b gene copy or two copies of the omp2a gene described in the six known Brucella species. Thus, molecular typing methods showed that this isolate is clearly distinct from all other known Brucella species and strains. The specific molecular features of this minke whale Brucella isolate raise questions about the lineage between the Brucella strains isolated from marine mammals and the Brucella species isolated from terrestrial mammals.

Animals↗

Nuclear and mitochondrial markers reveal distinctiveness of a small population of bottlenose whales (Hyperoodon ampullatus) in the western North Atlantic.

Small populations at the edge of a species' distribution can represent evolutionary relics left behind after range contractions due to climate change or human exploitation. The distinctiveness and genetic diversity of a small population of bottlenose whales in the Gully, a submarine canyon off Nova Scotia, was quantified by comparison to other North Atlantic populations using 10 microsatellites and mitrochondrial DNA (mtDNA) control region sequences (434 bp). Both markers confirmed the distinctiveness of the Gully (n = 34) from the next nearest population, off Labrador (n = 127; microsatellites -F(ST)= 0.0243, P < 0.0001; mtDNA -Phi(ST) = 0.0456, P < 0.05). Maximum likelihood microsatellite estimates suggest that less than two individuals per generation move between these areas, refuting the hypothesis of population links through seasonal migration. Both males and females appear to be philopatric, based on significant differentiation at both genomes and similar levels of structuring among the sexes for microsatellites. mtDNA diversity was very low in all populations (h = 0.51, pi = 0.14%), a pattern which may be due to selective sweeps associated with this species' extreme deep-diving ecology. Whaling had a substantial impact on bottlenose whale abundance, with over 65 000 animals killed before the hunt ceased in the early 1970s. Genetic diversity was similar among all populations, however, and no signal for bottlenecks was detected, suggesting that the Gully is not a relic of a historically wider distribution. Instead, this unique ecosystem appears to have long provided a stable year-round habitat for a distinct population of bottlenose whales.

Animals↗

The acoustic calls of blue whales off California with gender data.

The acoustic calls of blue whales off California are described with visual observations of behavior and with acoustic tracking. Acoustic call data with corresponding position tracks are analyzed for five calling blue whales during one 100-min time period. Three of the five animals produced type A-B calls while two produced another call type which we refer to as type D. One of the animals producing the A-B call type was identified as male. Pauses in call production corresponded to visually observed breathing intervals. There was no apparent coordination between the calling whales. The average call source level was calculated to be 186 dB re: 1 muPa at 1 m over the 10-110-Hz band for the type B calls. On two separate days, female blue whales were observed to be silent during respective monitoring periods of 20 min and 1 h.

Animals↗

The monopulsed nature of sperm whale clicks.

Traditionally, sperm whale clicks have been described as multipulsed, long duration, nondirectional signals of moderate intensity and with a spectrum peaking below 10 kHz. Such properties are counterindicative of a sonar function, and quite different from the properties of dolphin sonar clicks. Here, data are presented suggesting that the traditional view of sperm whale clicks is incomplete and derived from off-axis recordings of a highly directional source. A limited number of assumed on-axis clicks were recorded and found to be essentially monopulsed clicks, with durations of 100 micros, with a composite directionality index of 27 dB, with source levels up to 236 dB re: 1 microPa (rms), and with centroid frequencies of 15 kHz. Such clicks meet the requirements for long-range biosonar purposes. Data were obtained with a large-aperture, GPS-synchronized array in July 2000 in the Bleik Canyon off Vesterålen, Norway (69 degrees 28' N, 15 degrees 40' E). A total of 14 h of sound recordings was collected from five to ten independent, simultaneously operating recording units. The sound levels measured make sperm whale clicks by far the loudest of sounds recorded from any biological source. On-axis click properties support previous work proposing the nose of sperm whales to operate as a generator of sound.

Acoustics↗

Acoustic properties of humpback whale songs.

A vertical array of five hydrophones was used to measure the acoustic field in the vertical plane of singing humpback whales. Once a singer was located, two swimmers with snorkel gear were deployed to determine the orientation of the whale and position the boat so that the array could be deployed in front of the whale at a minimum standoff distance of at least 10 m. The spacing of the hydrophones was 7 m with the deepest hydrophone deployed at a depth of 35 m. An eight-channel TASCAM recorder with a bandwidth of 24 kHz was used to record the hydrophone signals. The location (distance and depth) of the singer was determined by computing the time of arrival differences between the hydrophone signals. The maximum source level varied between individual units in a song, with values between 151 and 173 dB re 1 microPa. One of the purposes of this study was to estimate potential sound exposure of nearby conspecifics. The acoustic field determined by considering the relative intensity of higher frequency harmonics in the signals indicated that the sounds are projected in the horizontal direction despite the singer being canted head downward anywhere from about 25 degrees to 90 degrees. High-frequency harmonics extended beyond 24 kHz, suggesting that humpback whales may have an upper frequency limit of hearing as high as 24 kHz.

Acoustics↗

Sperm whales (Physeter catodon L. 1758) do not react to sounds from detonators.

A number of observations show that sperm whales (Physeter catodon L. 1758) react to various man-made pulses with moderate source levels. The behavioral responses are described to vary from silence to fear. Click rates of five submerged male sperm whales were measured during the discharge of eight detonators off Andenes, northern Norway. In addition, the behavioral response of a surfaced specimen was observed. Click rates of the submerged whales and the behavior of the surfaced specimen did not change during the discharges with received sound levels of some 180 dB re 1 microPa peRMS. The apparent lack of response to the discharges could be due to similarity between sperm whale clicks and detonations. Accordingly, it can be speculated that the discharges may have been perceived as isolated clicks from conspecifics.

Animals↗

Baleen whales: preliminary evidence for forestomach microbial fermentation.

Baleen whales have a multichambered stomach divided into three distinct compartments. The forestomach (first compartment) consists of noncornified and nonglandular tissue and appears to be analogous to the tissue of the rumen. The exact function of the forestomach is unknown; however, we have detected volatile fatty acids in forestomach samples from seven bowhead (Balaena mysticetus) and four gray (Escherichtius robustus) whales. Acetic, propionic, and butyric acids were found in all samples. Seven whale samples also contained valeric, isobutyric, isovaleric, and isocaproic acids. Bacterial counts in 10 of the samples ranged from 5 X 10(8) to 630 X 10(8)/ml of fluid. The types, concentrations, and relative proportions of volatile fatty acids, the presence of significant levels of bacteria, and the morphology of the digestive tract support the hypothesis that a microbial forestomach fermentation occurs in these two species of baleen whales.

Animals↗

Chitinolytic bacteria in the minke whale forestomach.

Minke whales consume large amounts of pelagic crustaceans. Digestion of the prey is initiated by indigenous bacteria in a rumen-like forestomach system. A major structural component of the crustacean exoskeleton is chitin, the beta-1,4-linked polymer of N-acetyl-D-glucosamine. The exoskeletons appear to dissolve completely in the non-glandular forestomach. Bacteria in the forestomach fluid of six krill-eating minke whales were enumerated and isolated using an anaerobic habitat-simulating culture medium. Median viable population densities ranged between 6.0 x 10(6) and 9.9 x 10(9) bacterial cells per mL forestomach fluid. Bacterial isolates (n = 44) cultured from the forestomach fluid of one minke whale mainly resembled strains of Eubacterium (25%), Streptococcus (18%), Clostridium (14%), and Bacteroides (11%). As much as 12% of the bacterial isolates were chitinolytic, while beta-N-acetylglucosaminidase activity was demonstrated in 54% of the isolates, and utilisation of N-acetyl-D-glucosamine was observed in 73%. The chitinolytic isolates resembled strains of Bacteroides, Bacteroidaceae, Clostridium, and Streptococcus. Scanning and transmission electron microscopy of partly digested krill from the minke whale forestomach revealed bacteria close to and inside the chitinous exoskeleton. The bacterial chitinase may act on the chitinous crustacean exoskeletons, thereby allowing other bacteria access to the nutritious soft inner tissues of the prey, and thus initiating its degradation and fermentation.

Animals↗

Extreme diving of beaked whales.

Sound-and-orientation recording tags (DTAGs) were used to study 10 beaked whales of two poorly known species, Ziphius cavirostris (Zc) and Mesoplodon densirostris (Md). Acoustic behaviour in the deep foraging dives performed by both species (Zc: 28 dives by seven individuals; Md: 16 dives by three individuals) shows that they hunt by echolocation in deep water between 222 and 1885 m, attempting to capture about 30 prey/dive. This food source is so deep that the average foraging dives were deeper (Zc: 1070 m; Md: 835 m) and longer (Zc: 58 min; Md: 47 min) than reported for any other air-breathing species. A series of shallower dives, containing no indications of foraging, followed most deep foraging dives. The average interval between deep foraging dives was 63 min for Zc and 92 min for Md. This long an interval may be required for beaked whales to recover from an oxygen debt accrued in the deep foraging dives, which last about twice the estimated aerobic dive limit. Recent reports of gas emboli in beaked whales stranded during naval sonar exercises have led to the hypothesis that their deep-diving may make them especially vulnerable to decompression. Using current models of breath-hold diving, we infer that their natural diving behaviour is inconsistent with known problems of acute nitrogen supersaturation and embolism. If the assumptions of these models are correct for beaked whales, then possible decompression problems are more likely to result from an abnormal behavioural response to sonar.

Activity Cycles↗

Evidence that fin whales respond to the geomagnetic field during migration.

We challenge the hypothesis that fin whales use a magnetic sense to guide migration by testing for associations between geophysical parameters and the positions where fin whales were observed over the continental shelf off the northeastern United States. Monte Carlo simulations estimated the probability that the distribution of fin whale sighting was random with respect to bottom depth, bottom slope and the intensity and gradient of the geomagnetic field. The simulations demonstrated no overall association of sighting positions with any of these four geophysical parameters. Analysis of the data by season, however, demonstrated statistically reliable associations of sighting positions with areas of low geomagnetic intensity and gradient in winter and fall, respectively, but no association of sighting positions with bathymetric parameters in any season. An attempt to focus on migrating animals by excluding those observed feeding confirmed the associations of sighting positions with low geomagnetic intensity and gradient in winter and fall, respectively, and revealed additional associations with low geomagnetic gradients in winter and spring. These results are consistent with the hypothesis that fin whales, and perhaps other mysticete species, possess a magnetic sense that they use to guide migration.

Animals↗