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Mysticete (baleen whale) relationships based upon the sequence of the common cetacean DNA satellite.

The genomes of all extant cetaceans are characterized by the presence of the so-called common cetacean DNA satellite. In the mysticetes (whalebone whales) the repeat length of the satellite is 1,760 bp. In the odontocetes (toothed whales), other than the family Delphinidae, the repeat length is usually approximately 1,740 bp. The Delphinidae are characterized by a repeat length of approximately 1,580 bp. It has been shown in odontocetes that the satellite evolves in concert and that differences between species, with respect to the sequence of the satellite, correspond reasonably well to their evolutionary distances. In the present study the sequence of the satellite was determined in three repeats in each of seven mysticete species, and a consensus for each species established. Parsimony and neighbor-joining analyses based upon sequences of all repeats showed that the primary evolutionary distinction among the mysticetes is between the Balaenidae sensu stricto (i.e., the bowhead whale and the right whale) and all remaining species, including the pygmy right whale, a species that usually has been included in the Balaenidae. The comparisons also showed that the humpback whale and the gray whale were approximately equidistant from the blue whale and the fin whale (genus Balaenoptera). Concerted evolution of the satellite was also demonstrated among the mysticetes, but it appeared to evolve more slowly in the mysticetes than in the odontocetes.

Animals↗

Measuring the off-axis angle and the rotational movements of phonating sperm whales using a single hydrophone.

The common use of the bent-horn model of the sperm whale sound generator describes sperm whale clicks as the pulse series {p0, p1, p2, p3,...}. Clicks, however, deviate from this standard when recorded using off-axis hydrophones. The existence of additional pulses within the {p0, p1, p2, p3, ...} series can be explained still using the bent-horn model. Multiple reflections on the whale's frontal and distal sacs of the p0 pulse lead to additional sets of pulses detectable using a farfield, off-axis hydrophone. The travel times of some of these additional pulses depend on the whale's orientation. The authors propose a method to estimate the off-axis angle of sperm whale clicks. They also propose a method to determine the nature of the movement (if it is pitch, yaw, or roll) of phonating sperm whales. The application of both methods requires the measurement of the travel time differences between pulses composing a sperm whale click. They lead, using a simple apparatus consisting of a single hydrophone at an unknown depth, to new measurements of the underwater movements of sperm whales. Using these methods shows that sperm whales would methodically scan seawater while searching for prey, by making periodic pitch and yaw movements in sync with their acoustic activity.

Acoustics↗

Three-dimensional localization of sperm whales using a single hydrophone.

A three-dimensional localization method for tracking sperm whales with as few as one sensor is demonstrated. Based on ray-trace acoustic propagation modeling, the technique exploits multipath arrival information from recorded sperm whale clicks and can account for waveguide propagation physics like interaction with range-dependent bathymetry and ray refraction. It also does not require ray identification (i.e., direct, surface reflected) while utilizing individual ray arrival information, simplifying automation efforts. The algorithm compares the arrival pattern from a sperm whale click to range-, depth-, and azimuth-dependent modeled arrival patterns in order to estimate whale location. With sufficient knowledge of azimuthally dependent bathymetry, a three-dimensional track of whale motion can be obtained using data from a single hydrophone. Tracking is demonstrated using data from acoustic recorders attached to fishing anchor lines off southeast Alaska as part of efforts to study sperm whale depredation of fishing operations. Several tracks of whale activity using real data from one or two hydrophones have been created, and three are provided to demonstrate the method, including one simultaneous visual and acoustic localization of a sperm whale actively clicking while surfaced. The tracks also suggest that whales' foraging is shallower in the presence of a longline haul than without.

Acoustics↗

Passive acoustic location of bowhead whales in a population census off Point Barrow, Alaska.

A sonobuoy array placed in the nearshore lead was used for locating bowhead whale sounds to determine if whales migrated past census stations beyond visual range and were uncounted. Based on a sample of 182 whale sounds (over 48 h) from closest point of approach (CPA) distances out to more than 10 km, 68% originated beyond 2 km (CPA), where only 1% of the 242 whales were sighted. No whales were sighted beyond 3 km during this time, but 53% of the located sounds originated that far and beyond. Thirty-seven other bowhead sounds over 15 h were distributed out to 6 km. Two tracked whales moved at average speeds of 1.5 and 1.8 kn. Maximum location error was 1%-25% in a sector of 120 degrees X 5-10 km, depending upon bearing and range. Most whale sounds were low-frequency moans, trumpeting roars, and repetitive sequences (songs) with peak spectrum source level up to 189 dB re: 1 microPa, 1 m. Lack of correlations between numbers of sounds and sighted whales precluded using bowhead sounds to count individuals or even to extrapolate ratios of unseen to observed whales.

Alaska↗

Review of welfare concerns relating to commercial and special permit (scientific) whaling.

This paper reviews recent data relating to the efficiency and humaneness of the methods used to kill minke whales in commercial and special permit whaling operations. In the 1980s a grenade-headed harpoon was developed for minke whaling. Most recent results indicate that in the Norwegian industry approximately 60 per cent of whales were considered to be killed immediately by this harpoon, with 40 per cent being wounded. However, in Japanese whaling operations, generally only 30 per cent of whales were killed immediately, with 70 per cent being wounded. In both operations, more than six minutes elapsed before half the wounded whales were pronounced dead, with some whales surviving for more than an hour after being harpooned. Many of the long killing times were associated with a failure of a part of the whaling equipment

Animal Welfare↗

Identification of whale species by thin-layer isoelectric focusing of sarcoplasmic proteins.

Thin-layer isoelectric focusing was applied to the identification of whale (Cetacea) species by using water-soluble sarcoplasmic proteins of skeletal muscles. Twenty-eight samples consisting of 4 species (10 samples) of baleen whales (Mysticeti) and 8 species (18 samples) of toothed whales (Odontoceti) were analyzed. Each sample (approximately 1 g) was electrophoresed with Ampholine PAGplate, pH 3.5-9.5. The electrophoretic profiles were species-specific on the 4 toothed whale species that did not have a marked intra-species difference, and all 4 baleen whale species. However, the profiles were not specific on the 4 other dolphin species, even though they were discriminable from the other 4 toothed whale species. Numerical values of pIs and relative peak heights were obtained by densitometric analysis of the isoelectro-focused protein bands. The bands were also species-specific for the 8 toothed whale species mentioned. The values may be applicable to species identification without the need for a standard sample, which may not be readily obtainable. Experiments on test samples of minke and sel whales showed that bloodletting with ice water made the densities of isoelectro-focused bands thinner, although species identification was still possible by using the inside part of muscles. Heat treatment at below 60 degrees C for 10 min caused little denaturation; at higher temperatures the protein bands were diminished in a temperature-dependent fashion. Therefore, the present isoelectric focusing analysis should be applicable to small samples of whale meat, excluding several species of dolphins.

Animals↗

Anatomy of the sinus node, AV node, and His bundle of the heart of the sperm whale (Physeter macrocephalus), with a note on the absence of an os cordis.

BACKGROUND: Atrioventricular (AV) conduction time in large whales is only slightly greater than in smaller mammals even though their hearts are enormously larger. Little is known of the detailed histology or cytology of the conduction system of large whales. Such knowledge could be useful in defining the nature of cardiac rhythm and conduction of the whale as well as smaller mammals including humans. METHODS: We studied hearts from seven sperm whales. After fixation in formaldehyde and later dissection, specimens were prepared for histological examination. RESULTS: Cell size, histological organization, and innervation of the sperm whale's sinus node, AV node, and His bundle are similar to most mammalian hearts, except the sinus node is substantially larger. There is no central fibrous body between the atrial and ventricular septa, and the whale has no os cordis. Only the upper quarter of the interventricular septum is fully formed; below that there is only a thin layer of fatty connective tissue between the two ventricles. CONCLUSIONS: Given our morphological findings, we believe that the whale's comparatively short AV conduction time may be best explained by the sinus node and AV node functioning as coupled relaxation oscillators. Absence of an os cordis or central fibrous body or strong attachment between the two ventricles may pose both electrophysiological and hemodynamic hazards when the whale is no longer in its normally buoyant aquatic environment.

Animals↗

Diversity and duplication of DQB and DRB-like genes of the MHC in baleen whales (suborder: Mysticeti).

The molecular diversity and phylogenetic relationships of two class II genes of the baleen whale major histocompatibility complex were investigated and compared to toothed whales and out-groups. Amplification of the DQB exon 2 provided sequences showing high within-species and between-species nucleotide diversity and uninterrupted reading frames consistent with functional class II loci found in related mammals (e.g., ruminants). Cloning of amplified products indicated gene duplication in the humpback whale and triplication in the southern right whale, with average nucleotide diversity of 5.9 and 6.3%, respectively, for alleles of each species. Significantly higher nonsynonymous divergence at sites coding for peptide binding (32% for humpback and 40% for southern right) suggested that these loci were subject to positive (overdominant) selection. A population survey of humpback whales detected 23 alleles, differing by up to 21% of their inferred amino acid sequences. Amplification of the DRB exon 2 resulted in two groups of sequences. One was most similar to the DRB3 of the cow and present in all whales screened to date, including toothed whales. The second was most similar to the DRB2 of the cow and was found only in the bowhead and right whales. Both loci showed low diversity among species and apparent loss of function or altered function including interruption of reading frames. Finally, comparison of inferred protein sequence of the DRB3-like locus suggested convergence with the DQB, perhaps resulting from intergenic conversion or recombination.

Amino Acid Sequence↗

Organochlorine and metal contaminants in baleen whales: a review and evaluation of conservation implications.

The literature on organochlorine and metal contaminants in tissues of baleen whales includes data for approximately 1000 individuals in 10 species from various oceans of the world. Concentrations of these contaminants in tissues of baleen whales are low. Sources of variation in organochlorine concentrations in whales include age, sex, region of exposure, and feeding habits. Concentrations of sigma DDT and PCBs in baleen whales appear higher in the northern hemisphere than in the southern oceans, perhaps due to greater contamination of northern ecosystems and a higher incidence of fish in the diet. However, maximum concentrations are generally lower in tissues of baleen whales than in other marine mammal species, both on global and local scales. This is predictable based on the general distribution, habitats and trophic levels of baleen whales. We reviewed laboratory studies on the effects of selected organochlorine contaminants on direct mortality and impaired reproduction in other mammals, and critically examined observations attempting to link organochlorines to reproductive and population effects in marine mammals. There is no firm basis to conclude that the contaminants reviewed herein have affected baleen whale populations. Although more information on contaminants in baleen whales would be of value, as a matter of priority, research and management actions should focus on reducing human-caused mortality and increasing habitat carrying capacity for these species.

Animals↗

cDNA-derived amino acid sequences of myoglobins from nine species of whales and dolphins.

We determined the myoglobin (Mb) cDNA sequences of nine cetaceans, of which six are the first reports of Mb sequences: sei whale (Balaenoptera borealis), Bryde's whale (Balaenoptera edeni), pygmy sperm whale (Kogia breviceps), Stejneger's beaked whale (Mesoplodon stejnegeri), Longman's beaked whale (Indopacetus pacificus), and melon-headed whale (Peponocephala electra), and three confirm the previously determined chemical amino acid sequences: sperm whale (Physeter macrocephalus), common minke whale (Balaenoptera acutorostrata) and pantropical spotted dolphin (Stenella attenuata). We found two types of Mb in the skeletal muscle of pantropical spotted dolphin: Mb I with the same amino acid sequence as that deposited in the protein database, and Mb II, which differs at two amino acid residues compared with Mb I. Using an alignment of the amino acid or cDNA sequences of cetacean Mb, we constructed a phylogenetic tree by the NJ method. Clustering of cetacean Mb amino acid and cDNA sequences essentially follows the classical taxonomy of cetaceans, suggesting that Mb sequence data is valid for classification of cetaceans at least to the family level.

Amino Acid Sequence↗

Chemical contaminants in juvenile gray whales (Eschrichtius robustus) from a subsistence harvest in Arctic feeding grounds.

Gray whales are coastal migratory baleen whales that are benthic feeders. Most of their feeding takes place in the northern Pacific Ocean with opportunistic feeding taking place during their migrations and residence on the breeding grounds. The concentrations of organochlorines and trace elements were determined in tissues and stomach contents of juvenile gray whales that were taken on their Arctic feeding grounds in the western Bering Sea during a Russian subsistence harvest. These concentrations were compared to previously published data for contaminants in gray whales that stranded along the west coast of the US during their northbound migration. Feeding in coastal waters during their migrations may present a risk of exposure to toxic chemicals in some regions. The mean concentration (standard error of the mean, SEM) of sigmaPCBs [1400 (130) ng/g, lipid weight] in the blubber of juvenile subsistence whales was significantly lower than the mean level [27,000 (11,000) ng/g, lipid weight] reported previously in juvenile gray whales that stranded in waters off the west coast of the US. Aluminum in stomach contents of the subsistence whales was high compared to other marine mammal species, which is consistent with the ingestion of sediment during feeding. Furthermore, the concentrations of potentially toxic chemicals in tissues were relatively low when compared to the concentrations in tissues of other marine mammals feeding at higher trophic levels. These chemical contaminant data for the subsistence gray whales substantially increase the information available for presumably healthy animals.

Age Factors↗

Pathological and serological evidence of Brucella-infection in baleen whales (Mysticeti) in the western North Pacific.

Abnormal testes and uterus were observed in 13 males (33%) and one female (3%) out of 40 common minke whales (Balaenoptera acutorostrata) in the western North Pacific. Similar lesions were found in testis and ovary, respectively, in one male (2%) and female (2%) out of 43 Bryde's whales (Balaenoptera edeni) in the western North Pacific. Grossly, granular lesions with caseation and calcification were main pathological signs, and they were restricted to reproductive organs of mature whales. Chronic purulent or granulomatous orchitis was observed by microscopic analysis. Antibodies to Brucella species were detected in the serum samples of 15/40 (38%) of common minke whales and 4/43 (9%) of Bryde's whales. Neither pathological nor serological change was found in the examined sperm whales (Physeter macrocephalus) in the western North Pacific and Antarctic minke whales (Balaenoptera bonaerensis). These results strongly suggest that Brucella infection was involved in two species of baleen whales (Mysticeti) in the North Pacific.

Agglutination Tests↗

PCBs and organochlorine pesticides in blubber biopsies from free-ranging St. Lawrence River Estuary beluga whales (Delphinapterus leucas), 1994-1998.

For the first time, organochlorine (OC) contaminants were measured in blubber biopsies from free-ranging St. Lawrence River Estuary beluga whales (Delphinapterus leucas), to compare contaminant levels to those previously measured in dead stranded belugas. PCBs, DDTs, toxaphene and chlordane-related compounds were the major OC contaminants detected in 44 belugas biopsied in 1994-1998. SigmaPCB (the sum of 104 congeners) ranged from 2080 to 128,000 ng/g lipid in males (n=34; minimum estimated ages 8-22 years), and from 148 to 44,100 ng/g lipid in females (n=10; minimum estimated ages 7-22 years). The concentrations of PCBs and OC pesticides in the blubber of these whales overlapped those observed in stranded belugas from an earlier study, and demonstrated comparable age and sex-related trends. However, lower proportions of mirex, HCB, DDTs, and many of the highly chlorinated PCBs occurred in the biopsy samples compared to results for blubber from stranded carcasses. Most major OC compounds were present at lower concentrations in the biopsies, but this does not appear to be solely related to age differences between the two groups, or to emaciation in the stranded whales. Nor does it appear to be associated with the use of superficial biopsies, and the possible stratification of lipids and OCs in the blubber layer. Nevertheless, given these possible confounding factors, and the uncertainty in age estimates for the biopsied whales, the results point to the need for careful interpretation of biopsy results when comparing with data taken from the full depth of the blubber mantle in stranded whales. Taken together, results from both biopsied whales and previously studied stranded belugas indicate that PCB and OC pesticide contamination of St. Lawrence beluga whales may occur across a broader range of levels than previously thought, at least for males which formed the largest group in this study, possibly due to different degrees of dietary exposure. It also appears that measuring contaminant concentrations only in stranded whales, may overestimate OC levels in the population as a whole, especially for highly chlorinated OCs.

Animals↗

Relationship of baleen whales established by cytochrome b gene sequence comparison.

A recent revision of whale phylogeny suggested that the sperm whale was more closely related to rorquals than to other toothed whales. This made the suborder Odontoceti (toothed whales) paraphyletic, and implied that the latest common ancestor of rorquals and sperm whales may have lived only 10-13 million years ago. This is at variance with palaeontological evidence for the greater antiquity for both mysticetes (baleen whales) and sperm whales, so the Mysticeti, as well as the Odontoceti, must also be paraphyletic if the dates implied in ref. 1 were correct. Here we present a more comprehensive phylogenetic analysis that demonstrates the monophyly of mysticetes and identifies no particular affinity between the sperm whales and rorquals.

Animals↗

Molecular genetic identification of southern hemisphere beaked whales (Cetacea: Ziphiidae).

To assist in the species-level identification of stranded and hunted beaked whales, we compiled a database of 'reference' sequences from the mitochondrial DNA control region for 15 of the 20 described ziphiid species. Reference samples for eight species were obtained from stranded animals in New Zealand and South Australia. Sequences for a further seven species were obtained from a previously published report. This database was used to identify 20 'test' samples obtained from incompletely documented strandings around New Zealand. Analyses showed that four of these 'test' specimens (20%) had initially been misidentified. These included two animals of particular interest: (i) a Blainville's beaked whale (Mesoplodon densirostris), the first record of this species in New Zealand waters; and, (ii) a juvenile Andrews' beaked whale (Mesoplodon bowdoini), a species known from just over 20 strandings worldwide. A published sequence from a beaked whale product purchased in the Republic of Korea was identified as a Cuvier's beaked whale (Ziphius cavirostris). Levels of intra- and interspecific variation were compared to determine the potential for misidentification when the database or taxonomy is incomplete. Intraspecific variation was generally < 2%, and interspecific divergence was generally > 4.7%. Exceptions were within-species variation in Hyperoodon planifrons, southern bottlenosed whale (4.12%), which exceeded the variation between the two species of Berardius (3.78%), and variation between the two specimens assigned to M. hectori, Hector's beaked whale (7.14%). The latter case appears to be an error in species identification, and could represent the discovery of a new species of beaked whale.

Animals↗

World-wide genetic differentiation of Eubalaena: questioning the number of right whale species.

Few studies have examined systematic relationships of right whales (Eubalaena spp.) since the original species descriptions, even though they are one of the most endangered large whales. Little morphological evidence exists to support the current species designations for Eubalaena glacialis in the northern hemisphere and E. australis in the southern hemisphere. Differences in migratory behaviour or antitropical distribution between right whales in each hemisphere are considered a barrier to gene flow and maintain the current species distinctions and geographical populations. However, these distinctions between populations have remained controversial and no study has included an analysis of all right whales from the three major ocean basins. To address issues of genetic differentiation and relationships among right whales, we have compiled a database of mitochondrial DNA control region sequences from right whales representing populations in all three ocean basins that consist of: western North Atlantic E. glacialis, multiple geographically distributed populations of E. australis and the first molecular analysis of historical and recent samples of E. glacialis from the western and eastern North Pacific Ocean. Diagnostic characters, as well as phylogenetic and phylogeographic analyses, support the possibility that three distinct maternal lineages exist in right whales, with North Pacific E. glacialis being more closely related to E. australis than to North Atlantic E. glacialis. Our genetic results provide unequivocal character support for the two usually recognized species and a third distinct genetic lineage in the North Pacific under the Phylogenetic Species Concept, as well as levels of genetic diversity among right whales world-wide.

Animals↗

Mitochondrial phylogenetics and evolution of mysticete whales.

The phylogenetic relationships among baleen whales (Order: Cetacea) remain uncertain despite extensive research in cetacean molecular phylogenetics and a potential morphological sample size of over 2 million animals harvested. Questions remain regarding the number of species and the monophyly of genera, as well as higher order relationships. Here, we approach mysticete phylogeny with complete mitochondrial genome sequence analysis. We determined complete mtDNA sequences of 10 extant Mysticeti species, inferred their phylogenetic relationships, and estimated node divergence times. The mtDNA sequence analysis concurs with previous molecular studies in the ordering of the principal branches, with Balaenidae (right whales) as sister to all other mysticetes base, followed by Neobalaenidae (pygmy right whale), Eschrichtiidae (gray whale), and finally Balaenopteridae (rorquals + humpback whale). The mtDNA analysis further suggests that four lineages exist within the clade of Eschrichtiidae + Balaenopteridae, including a sister relationship between the humpback and fin whales, and a monophyletic group formed by the blue, sei, and Bryde's whales, each of which represents a newly recognized phylogenetic relationship in Mysticeti. We also estimated the divergence times of all extant mysticete species, accounting for evolutionary rate heterogeneity among lineages. When the mtDNA divergence estimates are compared with the mysticete fossil record, several lineages have molecular divergence estimates strikingly older than indicated by paleontological data. We suggest this discrepancy reflects both a large amount of ancestral polymorphism and long generation times of ancestral baleen whale populations.

Animals↗

Calcium-binding protein-containing neuronal populations in mammalian visual cortex: a comparative study in whales, insectivores, bats, rodents, and primates.

This study is focused on comparative analysis of gamma-aminobutyric acid-positive (GABAergic) neuronal populations in primary visual cortex of totally aquatic toothed whales and select terrestrial mammals with different evolutionary histories and various ecological adaptations. The distribution of neuronal populations containing the calcium-binding proteins calbindin and parvalbumin, which are recognized markers for the GABAergic neurons in cerebral cortex, is compared in five species of toothed whales and in representatives (one species each) of insectivores, bats, rodents, and primates. Computerized image analysis has shown that overall quantitative characteristics of GABAergic cortical neurons in toothed whales are similar to those in other mammalian orders. Thus, GABA-positive neurons represent 26% of the total population of cortical neurons in the visual cortex of whales. Some 97% of GABA-positive cells contain calcium-binding proteins, which is numerically similar to these parameters found in primates and other mammals. On the other hand, the typology and laminar distribution of calcium-binding protein-containing neurons in the primary visual cortex of five whale species (Delphinapterus leucas, Globicephala melaena, Phocoena phocoena, Stenella coeruleoalba, and Tursiops truncatus) differ significantly from those of primates (Macaca mulatta) and rodents (Rattus rattus) and are similar to those found in insectivorous bats (Eptesicus fuscus) and hedgehogs (Erinaceus europaeus). In whales, bats, and hedgehogs a significant concentration of calbindin-positive, vertically oriented bipolar and bitufted neurons was found in layers I, II, and IIIc/V with their axons arranged in a three-dimensional network. In primates and rodents they are distributed evenly across all cortical layers and are predominantly multipolar or bitufted neurons found in all cortical layers with their axons oriented along the vertical axis of the cortical plate. The parvalbumin-positive neurons in all mammalian species, including toothed whales, are represented by variously sized multipolar non-pyramidal cells. As opposed to all other mammalian species, the major concentrations of parvalbumin-positive neurons in whales are found in layers IIIc/V and VI, whereas in other cortical layers there are only scattered parvalbumin-positive neurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗