The systematics of right whales (Mysticeti: Balaenidae)

Balaenidae (right whales) are large, critically endangered baleen whales represented by four living species. The evolutionary relationships of balaenids are poorly known, with the number of genera, relationships to fossil taxa, and position within Mysticeti in contention. This study employs a comprehensive set of morphological characters to address aspects of balaenid phylogeny. A sister‐group relationship between neobalaenids and balaenids is strongly supported, although this conflicts with molecular evidence, which may be an artifact of long‐branch attraction (LBA). Monophyly of Balaenidae is supported, and three major clades are recognized: (1) extinct genus Balaenula, (2) extant and extinct species of the genus Eubalaena, and (3) extant and extinct species of the genus Balaena plus the extinct taxon, Balaenella. The relationships of these clades to one another, as well as to the early Miocene stem balaenid, Morenocetus parvus, remain unresolved. Pliocene taxa, Balaenula astensis and Balaenula balaenopsis, form a clade that is the sister group to the Japanese Pliocene Balaenula sp. Eubalaena glacialis and Pliocene Eubalaena belgica, are in an unresolved polytomy with a clade including E. japonica and E. australis. Extant and fossil species of Balaena form a monophyletic group that is sister group to the Dutch Pliocene Balaenella, although phylogenetic relationships within Balaena remain unresolved.     

Whale tear glands in the bowhead and the beluga whales: Source and function

Orbital glands are found in many tetrapod vertebrates, and are usually separate structures, consisting of individual glands lying in the eyelids and both canthi of the orbit. In cetaceans, however, the orbital glandular units are less distinct and have been described by numerous authors as a single, periorbital mass. There are few histochemical and immunhistochemical studies to date of these structures. In this study, we examined the orbital glandular region of both the bowhead whale (Balaena mysticetus: Mysticeti) and the beluga whale (Delphinapterus leucas: Odontoceti) using histological, histochemical, and immunohistochemical techniques. Histologically, in the bowhead, three glandular areas were noted (circumorbital, including Harderian and lacrimal poles), palpebral (midway in the lower eyelid), and rim (near the edge of the eyelid). In the beluga, there was only a large, continuous mass within the eyelid itself. Histochemical investigation suggests neither sexual dimorphism nor age-related differences, but both whales had two cell types freely intermingling with each other in all glandular masses. Large cells (cell type 1) were distended by four histochemically distinct intracellular secretory granules. Smaller cells (cell type 2) were not distended (fewer granules) and had two to three histochemically distinct intracellular secretory granules. The beluga orbital glands had additional lipid granules in cell type 1. Counterintuitively, both lipocalin and transferrin were localized to cell type 2 only. This intermingling of cell types is unusual for vertebrates in whom individual orbital glands usually have one cell type with one to two different secretory granules present. The heterogeneity of the orbital fluid produced by cetacean orbital glands implies a complex function, or series of functions, for these orbital glands and their role in producing the tear fluid.     

KILLER WHALES, ORCINUS ORCA, IN THE SOUTHEASTERN BERING SEA: RECENT SIGHTINGS andPREDATION ON OTHER MARINE MAMMALS

An unusual number of killer whales appeared in inshore waters of the southeastern Bering Sea in summer 1989 and 1990. Multiple sightings occurred in Bristol and Kuskokwim bays where killer whales had been seen only rarely in previous years. Three animals became stranded on mud flats in Kuskokwim Bay but were able to free themselves on a high tide. Killer whales were observed interacting with salmon, harbor seals, Steller sea lions, walruses, and beluga whales. Detailed observations were made of killer whales attacking belugas in the Naknek River. Local conditions and behavioral adaptations may reduce the susceptibility of belugas to killer whale predation. Continued killer whale activity in this area would be unlikely to affect fish resources, but might have some influence on beluga whales.     

Past and present distribution, densities and movements of blue whales Balaenoptera musculus in the Southern Hemisphere and northern Indian Ocean

• 1 Blue whale locations in the Southern Hemisphere and northern Indian Ocean were obtained from catches (303 239), sightings (4383 records of ≥8058 whales), strandings (103), Discovery marks (2191) and recoveries (95), and acoustic recordings.
• 2 Sighting surveys included 7 480 450 km of effort plus 14 676 days with unmeasured effort. Groups usually consisted of solitary whales (65.2%) or pairs (24.6%); larger feeding aggregations of unassociated individuals were only rarely observed. Sighting rates (groups per 1000 km from many platform types) varied by four orders of magnitude and were lowest in the waters of Brazil, South Africa, the eastern tropical Pacific, Antarctica and South Georgia; higher in the Subantarctic and Peru; and highest around Indonesia, Sri Lanka, Chile, southern Australia and south of Madagascar.
• 3 Blue whales avoid the oligotrophic central gyres of the Indian, Pacific and Atlantic Oceans, but are more common where phytoplankton densities are high, and where there are dynamic oceanographic processes like upwelling and frontal meandering.
• 4 Compared with historical catches, the Antarctic (‘true’) subspecies is exceedingly rare and usually concentrated closer to the summer pack ice. In summer they are found throughout the Antarctic; in winter they migrate to southern Africa (although recent sightings there are rare) and to other northerly locations (based on acoustics), although some overwinter in the Antarctic.
• 5 Pygmy blue whales are found around the Indian Ocean and from southern Australia to New Zealand. At least four groupings are evident: northern Indian Ocean, from Madagascar to the Subantarctic, Indonesia to western and southern Australia, and from New Zealand northwards to the equator. Sighting rates are typically much higher than for Antarctic blue whales.
• 6 South‐east Pacific blue whales have a discrete distribution and high sighting rates compared with the Antarctic. Further work is needed to clarify their subspecific status given their distinctive genetics, acoustics and length frequencies.
• 7 Antarctic blue whales numbered 1700 (95% Bayesian interval 860–2900) in 1996 (less than 1% of original levels), but are increasing at 7.3% per annum (95% Bayesian interval 1.4–11.6%). The status of other populations in the Southern Hemisphere and northern Indian Ocean is unknown because few abundance estimates are available, but higher recent sighting rates suggest that they are less depleted than Antarctic blue whales.

     

Antarctic killer whales make rapid,round-trip movements to subtropical waters: evidence for physiological maintenance migrations?

Killer whales (Orcinus orca) are important predators in high latitudes, where their ecological impact is mediated through their movements. We used satellite telemetry to provide the first evidence of migration for killer whales, characterized by fast (more than 12 km h(-1), 6.5 knots) and direct movements away from Antarctic waters by six of 12 type B killer whales tagged when foraging near the Antarctic Peninsula, including all tags transmitting for more than three weeks. Tags on five of these whales revealed consistent movements to subtropical waters (30-37° S) off Uruguay and Brazil, in surface water temperatures ranging from -1.9°C to 24.2°C; one 109 day track documented a non-stop round trip of almost 9400 km (5075 nmi) in just 42 days. Although whales travelled slower in the warmest waters, there was no obvious interruption in swim speed or direction to indicate calving or prolonged feeding. Furthermore, these movements were aseasonal, initiating over 80 days between February and April; one whale returned to within 40 km of the tagging site at the onset of the austral winter in June. We suggest that these movements may represent periodic maintenance migrations, with warmer waters allowing skin regeneration without the high cost of heat loss: a physiological constraint that may also affect other whales.     

Characterization of 25 microsatellite loci in bowhead whales (Balaena mysticetus)

Bowhead whales (Balaena mysticetus) experienced a severe demographic population bottleneck caused by commercial whaling that ceased in 1914. Aboriginal subsistence whale harvests have continued and are managed by the International Whaling Commission. In an effort to provide management advice for bowhead whales, 25 microsatellite loci were isolated from genomic DNA libraries. This panel of markers will be utilized to analyse stock structure hypotheses of current bowhead whale populations.     

Seasonal and diel changes in cetacean vocalizations monitored by passive acoustic methods in Nemuro Strait adjacent to the Shiretoko World Natural Heritage Site

UNESCO World Natural Heritage sites are established to ensure the long-term conservation of natural areas. Nemuro Strait in northern Japan is adjacent to the Shiretoko World Natural Heritage Site, and attracts various trophic levels of marine species, including marine mammals. Although the coexistence of humans and marine mammals is an important issue in this area, the temporal habitat use of cetaceans in this area is unknown. Here, we document seasonal and diel changes in cetacean vocalizations collected using passive acoustic recording devices during November 2012–March 2014. Killer whale calls occurred in spring and summer, and sperm whale clicks were detected in summer. Pacific white-sided dolphin calls were recorded in summer and late fall. No cetaceans were recorded during the sea ice period in February and March. The dolphin calls and unknown click trains were significantly more frequent at night. In contrast, marginal diel changes in killer whale calls were detected. Our results suggest that the majority of cetaceans utilize Nemuro Strait at night during the ice-free period, and we provide new insights into the habitat use and diversity of marine mammals in the Strait.     

Allele-sharing methods for estimation of population size

Genetic data are becoming increasingly important in ecology and conservation biology. This article presents a novel method for estimating population size from DNA profiles obtained from a random sample of individuals. The underlying idea is that the degree of biological relationship between individuals in the sample reflects the size of the population and that DNA profiles provide information about relatedness. A pseudolikelihood approach is taken, involving pairwise comparison of individuals. The main field of applications is seen to be catch data, and as an example, the method is applied to DNA profiles (10 microsatellite loci) from 334 North Atlantic minke whales. It is concluded that the sample size is too small for the method to give useful results. The question about the required sample size is investigated by simulation.     

ANALYSIS OF MITOCHONDRIAL DNA DIVERSITY WITHIN AND BETWEEN NORTH AND SOUTH ATLANTIC RIGHT WHALES

DNA sequences of the mitochondrial control region of 180 North Atlantic right whales ( Euhalaena glacialis ) and 16 South Atlantic right whales ( E. australis ) have been determined using a combination of direct DNA sequencing and single stranded conformation polymorphism (SSCP) analysis. Five haplotypes were found in E. glacialis , and 10 in E. australis , but none were shared, supporting the reproductive isolation and separate species status of the North and South Atlantic right whales. One haplotype in E, glacialis was found in only three males born before 1982 and this matriline will likely be lost soon. The nucleotide diversity estimates for the five North Atlantic right whale haplotypes was 0.6% and 2.0% for the 10 haplotypes found in the South Atlantic right whales. The average haplotypic diversity was 0.87 in E. glacialis and 0.96 in E. australis , which is consistent with other studies showing a lower level of genetic variation in the North Atlantic right whale. Phylogenetic analysis identified two major assemblages of haplotypes in E. australis from the samples collected from Peninsula Valdes, suggesting a mixing of two historically divergent populations. Using genetic distance measurements with a divergence rate of 0.5%–1.0%/myr, we estimate E. glacialis diverged from E. australis 3–12.5 mya.     

The effects of prey demography on humpback whale (Megaptera novaeangliae) abundance around Anvers Island, Antarctica

Baleen whales and Adelie penguins in the near-shore waters around the Antarctic Peninsula forage principally on Antarctic krill. Given the spatial overlap in the distribution of these krill predators (particularly humpback whales) and their dependence on krill, the goals of this paper are to determine if the inter-annual community structure and relative abundance of baleen whales around Anvers Island is related to krill demography and abundance, and if the potential exists for inter-specific interactions between Adelie penguins and baleen. We use whale sightings and prey data from both net tows and Adelie penguin stomach samples to correlate the abundance of humpback whales with krill demography and abundance from 1993 to 2001. We find significant relationships between whale abundance and the size–frequency distribution of krill targeted by Adelie penguins, as well as the foraging success of Adelie penguins. These findings suggest both krill predators share common prey preferences in the upper portions of the water column around Anvers Island. These findings highlight the need for better knowledge of baleen whale foraging ecology and inter-specific interactions with penguins, as sea ice and krill populations around the Antarctic Peninsula are affected by rapid changes in climate.     

The origins of the killer whale ecomorph

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Review of the distribution of Mesoplodon species (order Cetacea, family Ziphiidae) in the North Atlantic

Four species of beaked whales of the genus Mesoplodon regularly occur in the North Atlantic. These are Sowerby's Beaked Whale Mesoplodon bidens, Blainville's Beaked Whale Mesoplodon densirostris, Gervais Beaked Whale Mesoplodon europaeus and True's Beaked Whale Mesoplodon mirus. These four species differ in their distribution within this area. M. bidens is the most northerly recorded species followed by M. mirus. M. densirostris and M. europaeus both generally occur further south and their distributions cross the equator. The distribution of Mesoplodon species may relate to variations in water temperature.     

Dental care for a captive killer whale,Orcinus orca

The crowns of several teeth of a captive killer whale, particularly on the mandible, were worn to the level of the pulp cavities by biting a cement structure in the pool. Food plugging partially vacant pulp cavities created intense vascularization, inflammation, and eventually a systemic focus for infection. This trauma correlated with an elevated white blood cell count. Haematology was restored to normal following regular care for the worn teeth. Patent drainage of the pulp cavity was maintained through routine brushing with a large-scale toothbrush. Administration of antibiotics was not necessary in controlling the white blood cell count.     

Stable carbon isotope ratios for the gray whale (<Emphasis Type="Italic">Eschrichtius robustus</Emphasis>) in the breeding grounds of Baja California Sur,Mexico

This work investigated the feeding ecology and behaviour of gray whales in Bahía Magdalena. Underwater observations of bottom feeding were made (n=4). Skin biopsies of the gray whale had a carbon isotope value of –16.5 ± 0.1 (range from –16.4 to –16.7, n=7). Prey in Bahía Magdalena had a carbon isotope value of –18.4. Dietary enrichment from prey in Bahía Magdalena would correspond to 2 ± 0.1, whereas previously published results for prey in Alaska would result in an enrichment of 3, which suggests that whales were more likely feeding on prey from Bahía Magdalena. Carbon isotopic oscillation along the baleen plate of a stranded 1-year-old whale showed a variation in diet during the year, which suggests continual feeding during this time and corresponding to dietary sample measurements from Bahía Magdalena in winter and Alaska in summer.     

PREDICTING NUCLEAR GENE COALESCENCE FROM MITOCHONDRIAL DATA: THE THREE-TIMES RULE

Abstract.— Coalescence theory predicts when genetic drift at nuclear loci will result in fixation of sequence differences to produce monophyletic gene trees. However, the theory is difficult to apply to particular taxa because it hinges on genetically effective population size, which is generally unknown. Neutral theory also predicts that evolution of monophyly will be four times slower in nuclear than in mitochondrial genes primarily because genetic drift is slower at nuclear loci. Variation in mitochondrial DNA (mtDNA) within and between species has been studied extensively, but can these mtDNA data be used to predict coalescence in nuclear loci? Comparison of neutral theories of coalescence of mitochondrial and nuclear loci suggests a simple rule of thumb. The “three‐times rule” states that, on average, most nuclear loci will be monophyletic when the branch length leading to the mtDNA sequences of a species is three times longer than the average mtDNA sequence diversity observed within that species. A test using mitochondrial and nuclear intron data from seven species of whales and dolphins suggests general agreement with predictions of the three‐times rule. We define the coalescence ratio as the mitochondrial branch length for a species divided by intraspecific mtDNA diversity. We show that species with high coalescence ratios show nuclear monophyly, whereas species with low ratios have polyphyletic nuclear gene trees. As expected, species with intermediate coalescence ratios show a variety of patterns. Especially at very high or low coalescence ratios, the three‐times rule predicts nuclear gene patterns that can help detect the action of selection. The three‐times rule may be useful as an empirical benchmark for evaluating evolutionary processes occurring at multiple loci.     

A review of big data analysis methods for baleen whale passive acoustic monitoring

Many organizations collect large passive acoustic monitoring (PAM) data sets that need to be efficiently and reliably analyzed. To determine appropriate methods for effective analysis of big PAM data sets, we undertook a literature review of baleen whale PAM analysis methods. Methodologies from 166 studies (published between 2000–2019) were summarized, and a detailed review was performed on the 94 studies that recorded more than 1,000 hr of acoustic data (“big data”). Analysis techniques for extracting baleen whale information from PAM data sets varied depending on the research observed. A spectrum of methodologies was used and ranged from manual analysis of all acoustic data by human experts to completely automated techniques with no manual validation. Based on this assessment, recommendations are provided to encourage robust research methods that are comparable across studies and sectors, achievable across research groups, and consistent with previous work. These include using automated techniques when possible to increase efficiency and repeatability, supplementing automation with manual review to calculate automated detector performance, and increasing consistency in terminology and presentation of results. This work can be used to facilitate discussion for minimum standards and best practices to be implemented in the field of marine mammal PAM.