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Ultrastructure of the spermatozoa from three odontocetes: a killer whale (Orcinus orca), a Pacific white-sided dolphin (Lagenorhynchus obliquidens) and a beluga (Delphinapterus leucas).

Semen was collected from three captive odontocetes: a killer whale (Orcinus orca); a Pacific white-sided dolphin (Lagenorhynchus obliquidens) and a beluga (Delphinapterus leucas). Semen was collected from the killer whale and Pacific white-sided dolphin using behavioural commands, whereas semen from the beluga was collected postmortem. Ultrastructure of the spermatozoa was examined using scanning and transmission electron microscopy. Species differences were more pronounced in the head sections followed by the midpiece and finally, the tail sections. Spermatozoa from the killer whale and beluga were similar but differed from the Pacific white-sided dolphin spermatozoa.

Animals↗

Toxoplasmosis in beluga whales (Delphinapterus leucas) from the St Lawrence estuary: two case reports and a serological survey.

Toxoplasmosis was diagnosed in two free-ranging beluga whales from the St Lawrence estuary, Quebec, Canada, in 1988 and 1998. Histologically, tachyzoites and bradyzoites were present in the brain, spleen, lymph nodes, adrenals and lungs of both animals, and in the thymus of one. These organisms were readily labelled by an indirect immunohistochemical method for Toxoplasma gondii antigens. In the lymph nodes, spleen and lungs the organisms were associated with histiocytic infiltration. In the brain of one animal they were associated with mild multifocal gliosis and haemorrhages. There was no evidence of concomitant morbillivirus infection. Serum samples were collected from 22 beluga whales stranded between 1995 and 1998 on the shores of the St Lawrence Estuary and examined for antibodies to T. gondii by the modified agglutination test. Antibody titres of s25 were found in six (27%) of the animals. This is the first confirmed report of toxoplasmosis in beluga whales.

Animals↗

Pathological findings in two fin whales (Balaenoptera physalus) with evidence of morbillivirus infection.

Two immature female fin whales stranded on the Belgian and French coastlines, were examined post mortem. The main gross findings were massive parasitic infestation, associated with a large thrombus in one whale, and severe emaciation. Microscopical investigations revealed multinucleated syncytia with large intranuclear inclusion bodies in various tissues, and positive immunolabelling for morbillivirus antigens. Other evidence of morbillivirus infection was provided by the demonstration of specific viral structures in syncytia and in cell cultures, and the detection of neutralizing antibodies to canine distemper virus. To the authors>> knowledge, this is the first firm report of morbillivirus infection in baleen whales.

Animals↗

Comparative studies on mammalian Hoxc8 early enhancer sequence reveal a baleen whale-specific deletion of a cis-acting element.

Variations in regulatory regions of developmental control genes have been implicated in the divergence of axial morphologies. To find potentially significant changes in cis-regulatory regions, we compared nucleotide sequences and activities of mammalian Hoxc8 early enhancers. The nucleotide sequence of the early enhancer region is extremely conserved among mammalian clades, with five previously described cis-acting elements, A-E, being invariant. However, a 4-bp deletion within element C of the Hoxc8 early enhancer sequence is observed in baleen whales. When assayed in transgenic mouse embryos, a baleen whale enhancer (unlike other mammalian enhancers) directs expression of the reporter gene to more posterior regions of the neural tube but fails to direct expression to posterior mesoderm. We suggest that regulation of Hoxc8 in baleen whales differs from other mammalian species and may be associated with variation in axial morphology.

Animals↗

Do minke whales (Balaenoptera acutorostrata) digest wax esters?

Mammals are known to utilize wax esters with an efficiency of less than 50%. The purpose of the present study was to examine whether or not minke whales (Balaenoptera acutorostrata), which at times may eat considerable amounts of wax-ester-rich krill, represent an exception to this general pattern. Samples of fresh undigested forestomach, as well as colon, contents were obtained from minke whales (n 5) that had been feeding on krill (Thysanoessa inermis) for some time. The samples were analysed for dry mass, energy density, lipid content and the major lipid classes, including wax esters. The concentrations of wax esters were compared with previous estimates of dry-matter disappearance of the same type of prey using an in vitro technique, to calculate the dry-matter digestibility of wax esters (DMDwax). Wax esters contributed 21% of the energy and 47% of total lipids in the krill diet. The energy density of gut contents decreased by 50% after their passage from forestomach to the end of the colon. The DMDwax was 94.1 (SD 2.8)% (n 5). This high DMDwax and the occurrence of fatty alcohols, one of the products of wax-ester hydrolysis, in faeces show that minke whales are very efficient digesters of wax esters and absorb most of the energy-rich products of this process.

Animals↗

Distribution of inorganic mercury in liver and kidney of beluga and bowhead whales through autometallographic development of light microscopic tissue sections.

Inorganic mercury was localized through autometallography (AMG) in kidney and liver of free-ranging, subsistence-harvested beluga (Delphinapterus leucus: n = 20) and bowhead (Balaena mysticetrus: n = 5) whales. AMG granules were not evident in bowhead tissues, confirming nominal mercury (Hg) concentrations (range = 0.011 to 0.038 microg/g ww for total Hg). In belugas, total Hg ranged from 0.30 to 17.11 and from 0.33 to 82.47 microg/g ww in liver and kidney, respectively. AMG granules were restricted to cortical tubular epithelial cytoplasm in belugas with lower tissue burdens; whales with higher tissue burdens had granules throughout the uriniferous tubular epithelium. In liver, AMG granular densities differed between lobular zones, concentrating in stellate macrophages and bile cannalicular domains of hepatocytes. AMG granules aggregated in periportal regions in belugas with lower hepatic Hg concentrations, yet among whales with higher Hg, AMG granule deposition extended to pericentral and midzonal regions of liver lobules. Mean areas occupied by AMG granules correlated well with hepatic Hg concentrations and age. In beluga livers, AMG staining density was not associated with lipofuscin quantity (an index of oxidative damage). Occasionally, AMG granules and lipofuscin were colocalized, but more often were not, implying that Hg was not a prominent factor in hepatic lipofuscin deposition in belugas.

Animals↗

Dorsal skin color patterns among southern right whales (Eubalaena australis): genetic basis and evolutionary significance.

Distribution and inheritance of dorsal skin color markings among two populations of southern right whales (Eubalaena australis) suggest that two genes influence dorsal skin color. The grey-morph and partial-grey-morph phenotypes (previously known as partial albino and grey-blaze, respectively) appear to be controlled by an X-linked gene, whereas the white blaze appears controlled by an autosomal gene (recessive phenotype). Calving intervals, calf size, and length of sighting history data suggest that partial-grey-morph, white-blaze, and black cows experience similar levels of reproductive success. Grey-morph cows (XgXg) are rare or absent in the two populations, but this was not unexpected given observed population frequencies of grey-morph males (XgY) and partial-grey-morph females (XGXg). The proportion of partial-grey-morph calves produced by black cows (XGXG) suggests that the reproductive success of grey-morph males was equal to that of black males, however, larger sample sizes are required to determine whether grey-morph males tend to have shorter sighting histories. The reproductive success of white-blaze males appeared similar to that of black males among whales off Argentina. There were significantly fewer white-blaze calves than expected off South Africa, which could be due to white-blaze males experiencing reduced reproductive success or to sighting blases that result in white-marked calves being misidentified as black calves. The relative frequencies of both types of dorsal color markings varied between the South African and Argentinian right whale populations, suggesting limited nuclear gene flow between these populations; analyses using other nuclear markers are under way to confirm the extent of gene flow.

Animals↗

Major histocompatibility complex class I loci from the gray whale (Eschrichtius robustus).

Sequences from exons encoding the peptide binding region of MHC class I (MHC-I) molecules were isolated from California gray whale (Eschrichtius robustus) genomic DNA to initiate an investigation of variation in these genes in a cetacean. These represent the first mysticete MHC-I sequences to be reported. The analysis of gray whale MHC-I sequences suggests the presence of at least three loci, which share greatest similarity to MHC-I in the ungulates, consistent with current views on cetacean phylogenetics. The peptide binding region of MHC is the most polymorphic part of the molecule and analysis of the variation and synonymous to nonsynonymous substitution ratios in gray whale sequences found these genes to display polymorphism characteristics similar to that attributed to selection in other species.

Amino Acid Sequence↗

The effect of differential reproductive success on population genetic structure: correlations of life history with matrilines in humpback whales of the gulf of maine.

To examine whether demographic and life-history traits are correlated with genetic structure, we contrasted mtDNA lineages of individual humpback whales (Megaptera novaeangliae) with sighting and reproductive histories of female humpback whales between 1979 and 1995. Maternal lineage haplotypes were obtained for 323 whales, either from direct sequencing of the mtDNA control region (n = 159) or inferred from known relationships along matrilines from the sequenced sample of individuals (n = 164). Sequence variation in the 550 bp of the control region defined a total of 19 maternal lineage haplotypes that formed two main clades. Fecundity increased significantly over the study period among females of several lineages among the two clades. Individual maternal lineages and other clades were characterized by significant variation in fecundity. The detected heterogeneity of reproductive success has the potential to substantially affect the frequency and distribution of maternal lineages found in this population over time. There were significant yearly effects on adult resighting rates and calf survivorship based on examination of sighting histories with varying capture-recapture probability models. These results indicate that population structure can be influenced by interactions or associations between reproductive success, genetic structure, and environmental factors in a natural population of long-lived mammals.

Animals↗

Purification and properties of four biologically active components of whale luteinizing hormone.

Four biologically active components of luteinizing hormone (LH), designated as LH I, II, III, and IV, were isolated from whale pituitary glands. These components migrated as single bands with different mobilities in disc electrophoresis, and their isoeletric points were at pH 8.1, 8.3, 8.5, and 8.7, respectively. Their amino acid and carbohydrate compositions showed close similarity. The molecular weight of whale LH was determined to be 31,000 by sedimentation equilibrium, and no difference was detected among the four components. The molecular properties of whale LH were compared with the previously reported data for LH from other animal species.

Amino Acids↗

Thrombin-inhibitory activity of whale heparin oligosaccharides.

Whale heparin was partially digested with a purified heparinase and the oligosaccharide fractions with 8-20 monosaccharide units were isolated from the digest by gel filtration on Sephadex G-50, followed by affinity chromatography on a column of antithrombin III immobilized on Sepharose 4B. A marked difference in the inhibitory activity for thrombin in the presence of antithrombin III was observed between the high-affinity fractions for antithrombin III of octasaccharide approximately hexadecasaccharide and those of octadecasaccharide approximately eicosasaccharide. The disaccharide compositions of these hexadeca-, octadeca-, and eicosasaccharides were analyzed by high-performance liquid chromatography after digestion with a mixture of purified heparitinases 1 and 2 and heparinase. The analytical data indicated that the proportions of trisulfated disaccharide (IdUA(2S)alpha 1----4GlcNS(6S)) and disulfated disaccharide (UA1----4GlcNS(6S)) increased with the manifestation of high thrombin-inhibitory activity, while that of monosulfated disaccharide (UA1----4GlcNS) decreased. The present observations, together with those so far reported, suggest that the presence of the former structural elements, specifically IdUA(2S)alpha 1----4GlcNS(6S), as well as the antithrombin III-binding pentasaccharide at the proper positions in the molecules of whale heparin oligosaccharides is essential for the manifestation of high inhibitory activity for thrombin in the presence of antithrombin III. The structural bases for the manifestation of the anticoagulant activity of whale and porcine heparins and their oligosaccharides are also discussed.

Animals↗

Low diversity and biased substitution patterns in the mitochondrial DNA control region of sperm whales: implications for estimates of time since common ancestry.

The mitochondrial DNA (mtDNA) control region was sequenced in 37 sperm whales from a large part of the global range of the species. Nucleotide diversity was several-fold lower than that reported for control regions of abundant and outbred mammals, but similar to that for populations known to have experienced bottlenecks. Relative neck tests did not suggest that the low diversity is due to a lower substitution rate in sperm whale mtDNA. Rather, it is more likely that demographic factors have reduced diversity. The pattern of nucleotide substitutions was examined by cladistic methods, facilitated by the apparent monophyly of lineages from the Southern Hemisphere, as defined by a single base pair deletion. Substitutions were nonrandom in nature, confined to a few "hot spots," and parallel substitutions constituted a majority of the inferred changes. The substitution pattern fitted a negative binomial distribution better than a Poisson distribution, and the bias in number of substitutions among sites was considerably higher than previously reported for the mtDNA control region of any species. A novel method of estimating time since common ancestry was developed, which utilizes the transition/transversion ratio R and the number of substitutions inferred from a parsimony analysis. Using this method, we estimated the age of sperm whale mtDNA diversity to be about 6,000-25,000 years, and when the uncertainty of R was accounted for, a range of about 1,000-100,000 years was obtained.

Animals↗

Urinary concentrations of ovarian steroid hormone metabolites and bioactive follicle-stimulating hormone in killer whales (Orcinus orchus) during ovarian cycles and pregnancy.

Reproductive hormone profiles of six captive killer whales (Orcinu orcus) from three Sea World aquaria were studied for intervals up to 2 yr. Daily urine samples and bimonthly blood samples were collected and analyzed for hormone concentration. Immunoreactive estrone conjugates, pregnanediol-3-glucoruonide, 20-alpha-hydroxyprogesterone as well as bioactive follicle-stimulating hormone (FSH) were measured in urine samples and indexed by creatinine concentrations of the same sample. In selected cases, serum progesterone concentrations were also measured. Three of the animals in the study became pregnant during the study period and two of these animals were evaluated during the time of conception and throughout most of gestation. From the data of the three animals that conceived, hormone profiles of the complete ovarian cycle, early pregnancy, and mid- to late gestation are described. The remaining three animals did not conceive and only one of these demonstrated hormone changes that indicated regular ovarian activity. The female reproductive pattern of the killer whale is characterized by a gestation of 17 mo and an ovarian cycle of 6-7 wk in duration. The hormone changes associated with the ovarian cycle of the killer whale are similar to those of most other mammalian species. A bimodal pattern of bioactive FSH with a pronounced rise of estrogen predominates the preovulatory hormone profile. After ovulation, increased progesterone production is observed for approximately 4 wk in the nonconceptive ovarian cycle. During the luteal phase and early pregnancy, when progesterone metabolites are elevated, estrogen metabolite excretion remains low. These data extend the application of urine collections for longitudinal studies involving hormone changes, particularly those involving nondomesticated species.(ABSTRACT TRUNCATED AT 250 WORDS)

20-alpha-Dihydroprogesterone↗

Against the current: an inter-oceanic whale migration event.

Humpback whales seasonally migrate long distances between tropical and polar regions. However, inter-oceanic exchange is rare and difficult to document. Using skin biopsy samples collected in the Indian Ocean and in the South Atlantic Ocean, and a genetic capture-recapture approach based on microsatellite genotyping, we were able to reveal the first direct genetic evidence of the inter-oceanic migration of a male humpback whale. This exceptional migration to wintering grounds of two different ocean basins questions traditional notions of fidelity to an ocean basin, and demonstrates how the behaviour of highly mobile species may be elucidated from combining genetics with long-term field studies. Our finding has implications for management of humpback whale populations, as well as for hypotheses concerning cultural transmission of behaviour.

Animal Migration↗

Global climate drives southern right whale (Eubalaena australis) population dynamics.

Sea surface temperature (SST) time-series from the southwest Atlantic and the El Niño 4 region in the western Pacific were compared to an index of annual calving success of the southern right whale (Eubalaena australis) breeding in Argentina. There was a strong relationship between right whale calving output and SST anomalies at South Georgia in the autumn of the previous year and also with mean El Niño 4 SST anomalies delayed by 6 years. These results extend similar observations from other krill predators and show clear linkages between global climate signals and the biological processes affecting whale population dynamics.

Animals↗

A killer whale social network is vulnerable to targeted removals.

Individuals play various roles in maintaining social integrity of mammalian populations. However, many models developed for managing wildlife resources assume that all individuals are equal. Killer whales are social animals that rely on relationships within and among family groups for survival. In the northeastern Pacific, fish-eating, 'resident' killer whale populations are composed of matrilines from which offspring do not disperse. We analysed the influence of various individuals' age, sex and matrilineal affiliation on their position in a social network. Here, we show that some matrilines appeared to play more central roles than others in the network. Furthermore, juvenile whales, especially females, appeared to play a central role in maintaining network cohesion. These two key findings were supported subsequently by simulating removal of different individuals. The network was robust to random removals; however, simulations that mimicked historic live-captures from the northeastern Pacific were likely to break the network graph into isolated groups. This finding raises concern regarding targeted takes, such as live-capture or drive fisheries, of matrilineal cetaceans.

Animals↗

Nuclear markers confirm taxonomic status and relationships among highly endangered and closely related right whale species.

Right whales (genus: Eubalaena) are among the most endangered mammals, yet their taxonomy and phylogeny have been questioned. A phylogenetic hypothesis based on mitochondrial DNA (mtDNA) variation recently prompted a taxonomic revision, increasing the number of right whale species to three. We critically evaluated this hypothesis using sequence data from 13 nuclear DNA (nuDNA) loci as well as the mtDNA control region. Fixed diagnostic characters among the nuclear markers strongly support the hypothesis of three genetically distinct species, despite lack of any diagnostic morphological characters. A phylogenetics analysis of all data produced a strict consensus cladogram with strong support at nodes that define each right whale species as well as relationships among species. Results showed very little conflict among the individual partitions as well as congruence between the mtDNA and nuDNA datasets. These data clearly demonstrate the strength of using numerous independent genetic markers during a phylogenetics analysis of closely related species. In evaluating phylogenetic support contributed by individual loci, 11 of the 14 loci provided support for at least one of the nodes of interest to this study. Only a single marker (mtDNA control region) provided support at all four nodes. A study using any single nuclear marker would have failed to support the proposed phylogeny, and a strong phylogenetic hypothesis was only revealed by the simultaneous analysis of many nuclear loci. In addition, nu DNA and mtDNA data provided complementary levels of support at nodes of different evolutionary depth indicating that the combined use of mtDNA and nuDNA data is both practical and desirable.

Animals↗

A bizarre new toothed mysticete (Cetacea) from Australia and the early evolution of baleen whales.

Extant baleen whales (Cetacea, Mysticeti) are all large filter-feeding marine mammals that lack teeth as adults, instead possessing baleen, and feed on small marine animals in bulk. The early evolution of these superlative mammals, and their unique feeding method, has hitherto remained enigmatic. Here, I report a new toothed mysticete from the Late Oligocene of Australia that is more archaic than any previously described. Unlike all other mysticetes, this new whale was small, had enormous eyes and lacked derived adaptations for bulk filter-feeding. Several morphological features suggest that this mysticete was a macrophagous predator, being convergent on some Mesozoic marine reptiles and the extant leopard seal (Hydrurga leptonyx). It thus refutes the notions that all stem mysticetes were filter-feeders, and that the origins and initial radiation of mysticetes was linked to the evolution of filter-feeding. Mysticetes evidently radiated into a variety of disparate forms and feeding ecologies before the evolution of baleen or filter-feeding. The phylogenetic context of the new whale indicates that basal mysticetes were macrophagous predators that did not employ filter-feeding or echolocation, and that the evolution of characters associated with bulk filter-feeding was gradual.

Animals↗