Song sharing and diversity in the Bering‐Chukchi‐Beaufort population of bowhead whales (Balaena mysticetus), spring 2011

Bowhead whales (Balaena mysticetus) of the Bering‐Chukchi‐Beaufort population migrate in nearshore leads through the Chukchi Sea each spring to summering grounds in the Beaufort Sea. As part of a population abundance study, hydrophones were deployed in the Chukchi Sea off Point Barrow, (12 April to 27 May 2011) and in the Beaufort Sea (12 April to 30 June 2011). Data from these sites were analyzed for the presence of bowhead whale song. We identified 12 unique song types sung by at least 32 individuals during ~95 h of recordings off Point Barrow. Six of these songs were detected at the Beaufort MARU site as well as six additional song types that were not analyzed. These results suggest a shared song repertoire among some individuals. This report represents the greatest number of songs to date during the spring migration for this population. We attribute this greater variety to population growth over the 30 yr since acoustic monitoring began in the early 1980s. Singing during early to mid‐spring is consistent with the hypothesis that song is a reproductive display, but further research is necessary to understand the exact function of this complex vocal behavior.     

A bowhead whale in the eastern North Pacific

Bowhead whales occur in the Arctic year‐round. Their movements are largely correlated with seasonal expansions and reductions of sea ice, but a few recent extralimital sightings have occurred in the eastern and western North Atlantic and one was also documented in the western North Pacific over 50 years ago. Here we present details of a juvenile bowhead whale that was photographed and filmed from above and below the water while it was skim‐feeding in Caamaño Sound, BC, Canada on May 31, 2016. This sighting occurred over 2000 km southeast from the nearest known range for this species in the Bering Sea at a time that most bowhead whales in that region would have been migrating northeast. This sighting represents the first and only documentation of a bowhead whale in the eastern North Pacific to date.     

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.     

A Swedish subfossil find of a bowhead whale from the late Pleistocene: shore displacement,paleoecology in south-west Sweden and the identity of the Swedenborg whale (Balaena swedenborgii Liljeborg, 1867)

The Swedenborg whale Balaena swedenborgii Liljeborg, 1867, is a baleen whale species believed to have existed in the North Sea from the period when the inland ice melted around 13,000 before present (BP) until about 8000 years ago. The first bones attributed to this species were found in Sweden in 1705. When whale remains were discovered on the Swedish west coast during the extension work of a motorway extension, it was speculated that this could be a specimen of the extinct Swedenborg whale. The bones were found 72 m above the present-day sea level embedded in glacial mud. Shelly remains of marine organisms were present in the deposit surrounding the whale-fall, and sediments with the associated specimens were therefore collected for further analyses. We applied radiocarbon dating, thin sectioning, morphological analyses, ancient DNA typing and analyses of the associated shelly assemblage in an interdisciplinary effort to understand the circumstances of this fossil whale-fall. Our results show that the whale is not the putative species B. swedenborgii, but a bowhead whale Balaena mysticetus. The results also indicate that the whale must have been rapidly covered by glacial sediments, highlighting the speed of the deglacial process in the area.     

Examination of ten thousand years of mitochondrial DNA diversity and population demographics in bowhead whales (Balaena mysticetus) of the Central Canadian Arctic

Mitochondrial DNA (mtDNA) sequences were analyzed from 106 bowhead whale (Balaena mysticetus) specimens dating 471 ± 44 ¹⁴C b.p. –10,290 ± 150 ¹⁴C b.p. to evaluate whether historical changes in distribution and connectivity were detectable in levels of diversity and population structuring in the Central Canadian Arctic. The species has maintained levels of mtDNA diversity over 10,000 yr comparable to other nonbottlenecked large whale species. When compared to data from the Holocene East Greenland/Spitsbergen and contemporary Bering‐Chuckchi‐Beaufort populations, differentiation was low (F_ST≤ 0.005, Φ_ST≤ 0.003) and no temporal or geographical genetic structuring was evident. A combination of analyses suggests that the population has expanded over the past 30,000 ¹⁴C yr. This genetic signature of expansion could result from population growth, admixture of multiple gene pools, or a combination of both scenarios. Despite known climatic change that altered bowhead distribution and led to isolation of populations, there is no detectable population structuring or change in genetic diversity during the Holocene. This may be due to long generation time, occasional population connectivity and a historically large global population. These characteristics warrant caution when interpreting contemporary bowhead whale DNA data, as it is unlikely that any population will be in mutation‐drift equilibrium.     

Behaviour and kinematics of continuous ram filtration in bowhead whales (Balaena mysticetus)

Balaenid whales perform long breath-hold foraging dives despite a high drag from their ram filtration of zooplankton. To maximize the volume of prey acquired in a dive with limited oxygen supplies, balaenids must either filter feed only occasionally when prey density is particularly high, or they must swim at slow speeds while filtering to reduce drag and oxygen consumption. Using digital tags with three-axis accelerometers, we studied bowhead whales feeding off West Greenland and present here, to our knowledge, the first detailed data on the kinematics and swimming behaviour of a balaenid whale filter feeding at depth. Bowhead whales employ a continuous fluking gait throughout the bottom phase of foraging dives, moving at very slow speeds (less than 1 m s⁻¹), allowing them to filter feed continuously at depth. Despite the slow speeds, the large mouth aperture provides a water filtration rate of approximately 3 m³ s⁻¹, amounting to some 2000 tonnes of water and prey filtered per dive. We conclude that a food niche of dense, slow-moving zooplankton prey has led balaenids to evolve locomotor and filtering systems adapted to work against a high drag at swimming speeds of less than 0.07 body length s⁻¹ using a continuous fluking gait very different from that of nekton-feeding, aquatic predators.     

Bowhead whales Balaena mysticetus in the Okhotsk Sea

• 1 Little is known about the endangered population of bowhead whales Balaena mysticetus in the Okhotsk Sea (OS). Here, we review existing information about this stock, including much material published in Russian.
• 2 Whaling for OS bowheads began around 1846, was pursued intensively for two decades and continued sporadically until about 1913. Beginning in 1967, whalers from the USSR killed bowheads illegally, although the number of whales taken remains unknown. Estimates of the pre‐exploitation population size have ranged from 3000 to 20000 whales, but all such estimates are based upon untested assumptions and incomplete data.
• 3 Information on historical and current distribution of bowheads comes from whaling records (notably Townsend 1935 ) and from modern (notably Russian/Soviet) marine mammal surveys. Little is known about winter distribution. During spring and summer, known bowhead concentrations occur in Shelikhov Bay and at Shantar. Although historical whaling data show bowheads in Shelikhov Bay during summer and early autumn, there have been no recent sightings later than June. However, extensive 19th century catches were made over much of the northern OS, and the present range and habitat use of the population is probably broader than existing data suggest. There is evidence for age or maturational class segregation between Shantar and Shelikhov Bay; the former hosts immature whales and lactating females, and the latter hosts adults.
• 4 Genetic data indicate that the OS bowhead stock is separate from the Bering‐Chukchi‐Beaufort population, but that the two populations share a common ancestry. There is no evidence that bowheads ever leave the OS.
• 5 Russian observers have put the current size of the OS stock in the low hundreds, but this is not based on quantitative analysis. Overall, the OS bowhead population is very likely to be relatively small; it did not recover from the intensive whaling in the 19th century, and the illegal Soviet catches of the 1960s have further set back its recovery. Dedicated surveys and other research are required to assess the status and conservation needs of the population.

     

Allometric patterns of fetal head growth in mysticetes and odontocetes: Comparison of Balaena mysticetus and Stenella attenuata

Unlike other mammals, odontocetes and mysticetes have highly derived craniofacial bones. A growth process referred to as “telescoping” is partly responsible for this morphology. Here, we explore how changes in facial morphology during fetal growth relate to differences in telescoping between the adult odontocete Stenella attenuata and the mysticete Balaena mysticetus. We conclude that in both Stenella and Balaena head size increases allometrically. Similarly, odontocete nasal length and mysticete mouth size have strong positive allometry compared to total body length. However, the differences between odontocetes and mysticetes in telescoping are not directly associated with their fetal growth patterns. Our results suggest that cranial changes related to echolocation and feeding between odontocetes and mysticetes, respectively, begin during ontogeny before telescoping is initiated.     

SURFACE MORPHOLOGY OF THE FOREBRAIN OF THE BOWHEAD WHALE, BALAENA MYSTICETUS

Observations were made on 11 brains from bowhead whales subsistence-harvested by Alaskan Eskimos under International Whaling Commission guidelines. This study is part of a larger project to determine the basic morphology of this endangered species. The bowhead brain is similar to other cetacean brains, particularly that of the southern right whale. Long olfactory peduncles are reflected upon the rostrodorsal surface of the cerebral hemispheres. Olfactory bulbs have not been recovered but are presumed to exist since nerve fibers have been identified histologically in the olfactory peduncles. The induseum griseum is evident on the corpus callosum. The hippocampus proper is small but protrudes into the lateral ventricle. The cruciate sulcus runs diagonally across the rostral surface, limiting a small frontal lobe. The structure of the floor of the sylvian fissure varies from a few short gyri radiating toward the circular sulcus to a more extensive and complex two-tiered arrangement including numerous gyri perpendicular to the gyrus bordering the paleocortex. Pineal-body-like tissue was present in one specimen. There is no interthalamic adhesion. The lateral geniculate body is elevated but smaller than the large medial geniculate body. The neurohypophysis was adherent on most brain specimens received.     

Age estimates based on aspartic acid racemization for bowhead whales (Balaena mysticetus) harvested in 1998–2000 and the relationship between racemization rate and body temperature

Fifty‐two eyes were collected and analyzed to estimate ages of 42 bowhead whales using the aspartic acid racemization aging technique. Between‐eye and within‐eye variance components for the ratio of the D and L optical isomers (D/L ratio) were estimated via analysis of variance using multiple measurements from nine whales with both eyes sampled and analyzed. For whales with more than one (D/L)_act value, an inverse variance weighted average of the values was used as (D/L)_act for the whale. Racemization rate (k_Asp) and D/L ratio at birth (D/L)₀ were estimated using (D/L)_act from 27 bowhead whales with age estimates based on baleen or ovarian corpora data and two term fetuses. The estimates were k_Asp = 0.977 × 10^?3/yr and (D/L)₀ = 0.0250. The nonlinear least squares analysis that produced these estimates also estimated female age at sexual maturity as ASM = 25.86 yr. SE(age) was estimated via a bootstrap that took into account the SE of (D/L)_act and the variances and covariance of k_Asp and (D/L)₀. One male exceeded 100 yr of age; the oldest female was 88. A strong linear relationship between k_Asp and body temperature was estimated by combining bowhead data with independent data from studies of humans and fin whales. Using this relationship, we estimated k_Asp and ASM for North Atlantic minke whales.     

Detecting genetic structure in migrating bowhead whales off the coast of Barrow, Alaska

We develop a general framework for analysing and testing genetic structure within a migratory assemblage that is based on measures of genetic differences between individuals. We demonstrate this method using microsatellite DNA data from the Bering-Chukchi-Beaufort stock of bowhead whales (Balaena mysticetus), sampled via Inuit hunting during the spring and autumn migration off Barrow, Alaska. This study includes a number of covariates such as whale ages and the time separation between captures. Applying the method to a sample of 117 bowhead whales, we use permutation methods to test for temporal trends in genetic differences that can be ascribed to age-related effects or to timing of catches during the seasons. The results reveal a pattern with elevated genetic differences among whales caught about a week apart, and are statistically significant for the autumn migration. In contrast, we find no effects of time of birth or age-difference on genetic differences. We discuss possible explanations for the results, including population substructuring, demographic consequences of historical overexploitation, and social structuring during migration.     

Vitamin A and E tissue distribution with comparisons to organochlorine concentrations in the serum, blubber and liver of the bowhead whale (Balaena mysticetus)

Vitamin A and E concentrations were determined in liver (n = 51), blubber (n = 23) and serum (n = 53) of subsistence-hunted bowhead whales (Balaena mysticetus), between 1998 and 2001. Retinol and alpha-tocopherol were the major forms of vitamins A and E detected, respectively. Liver contained the highest mean concentrations of vitamin A, followed by epidermis, blubber, and serum. Liver also contained the highest mean concentration of vitamin E, followed by serum, epidermis, and blubber. Stratification of retinol and tocopherol was examined throughout the blubber cores collected. Retinol concentrations were significantly higher in the epidermis than in the deeper blubber layers. Tocopherol concentrations were similar for epidermis and the intermediate layer of blubber. Both the epidermis and the intermediate layer of blubber had significantly higher tocopherol concentrations than the innermost and outermost blubber layers. Vitamin A and E concentrations were investigated with respect to gender and reproductive status of females (males, non-pregnant females, pregnant females), age groups and season of harvest. Certain persistent organic contaminants are known to have a negative effect on retinol concentration in serum of pinnipeds and cetaceans. Bowhead whales have relatively low concentrations of organochlorines (OCs) in comparison to other mysticete species. The relationships between serum, liver and blubber retinol and serum and blubber OC concentrations were examined with no significant correlations noted.     

Microflora associated with the skin of the bowhead whale (Balaena mysticetus)

A study of the microbiological flora isolated from cultures of normal and lesional skin tissue samples collected from 19 bowhead whales (Balaena mysticetus) over a 4 yr period is presented. These cultures were obtained from 30 tissue samples (17 normal, 13 lesion) and 248 swab samples (157 normal, 91 lesion). Seven hundred-thirty bacterial and yeast isolations were made (285 normal, 445 lesion). Distribution revealed that 56% of the gram positive bacterial isolates, 75% of the gram negative bacterial isolates and 64% of the yeast isolates recovered were associated with lesional skin. It was found that 80% of one group of Corynebacterium sp. isolates, 90% of the Acinetobacter sp. isolates and 94% of the Moraxella sp. isolates were associated with lesional skin. Although the primary yeasts recovered were Candida spp., they were found on both normal and lesional skin. Enzymatic assays of isolates from normal and lesional skin demonstrated production of enzymes capable of causing necrosis. The majority of the microorganisms recovered were facultative anaerobes and many of them could be considered potential pathogens of mammalian hosts.     

Serology and virology of the bowhead whale (Balaena mysticetus L.)

Sera from four bowhead whales (Balaena mysticetus L.) were examined for the presence of specific antibodies, and tissue and swab samples from six and four animals respectively were processed for isolation of viruses and for initiation of bowhead whale cell cultures. All sera were negative for antibodies to nine serovars of Leptospira interrogans and to 21 orthomyxovirus subtypes and a paramyxovirus (Newcastle disease virus). All sera were positive, however, for neutralizing antibodies to one or more calicivirus serotypes. Two untyped adenoviruses were isolated from colon samples of two different whales, but neutralizing antibodies to the agents could not be demonstrated in any sera. Three primary bowhead whale cell cultures were derived from kidney (two cultures) and testis (one culture), from three individual whales.     

Fractured mandible and associated oral lesions in a subsistence-harvested bowhead whale (Balaena mysticetus)

A fractured right mandible with midlength nonunion and oral lesions were noted in a subsistence-harvested female bowhead whale (Balaena mysticetus) near Wainwright, Alaska (USA). The cause of the fracture was not apparent. The fracture resulted in misalignment of the mandible. The abnormal mobility at the fracture site probably caused irregular baleen stowage within the oral cavity, leading to breakage of many baleen plates and extensive ulceration of the tongue and lips. Good body condition suggested the fracture was not debilitating.     

Recent sightings of the bowhead whale (Balaena mysticetus) in Northeast Greenland and the Greenland Sea

By the end of the 19th century, European whalers had brought the bowhead whale (Balaena mysticetus) in the “Spitsbergen stock” ranging the waters between eastern Greenland and the western Russian Arctic to the verge of extinction. This paper presents observations of this species in Northeast Greenland and in the Greenland Sea between 1940 and 2004. The number of observations has increased in Northeast Greenland since the mid-1980s. Only three observations are known for the period 1940–1979 but during the 1980s, the 1990s and between 2000 and 2004, six, six and eight observations of bowhead whales were made, involving an absolute minimum of three, five and eight to ten different individuals, respectively. It remains uncertain whether this represents an increase in survey effort, an immigration from other areas, a recent recovery of an eastern Greenland relict “tribe” belonging to the Spitsbergen stock, or a combination of these factors.     

Effects of airgun sounds on bowhead whale calling rates in the Alaskan Beaufort Sea

This study assesses effects of airgun sounds on bowhead calling behavior during the autumn migration. In August–October 2007, 35 directional acoustic recorders (DASARs) were deployed at five sites in the Alaskan Beaufort Sea. Location estimates were obtained for >137,500 individual calls; a subsample of locations with high detection probability was used in the analyses. Call localization rates (CLRs) were compared before, during, and after periods of airgun use between sites near seismic activities (median distance 41–45 km) and sites relatively distant from seismic activities (median distance >104 km). At the onset of airgun use, CLRs dropped significantly at sites near the airguns, where median received levels from airgun pulses (SPL) were 116–129 dB re 1 μPa (10–450 Hz). CLRs remained unchanged at sites distant from the airguns, where median received levels were 99–108 dB re 1 μPa. This drop could result from a cessation of calling, deflection of whales around seismic activities, or both combined, but call locations alone were insufficient to differentiate between these possibilities. Reverberation from airgun pulses could have masked a small number of calls near the airguns, but even if masking did take place, the analysis results remain unchanged.     

ELEMENTS IN LIVER TISSUES OF BOWHEAD WHALES (BALAENA MYSTICETUS)

Livers from 20 bowhead whales taken in subsistence hunts at Barrow, Alaska, were analyzed for essential and non-essential elements. Concentrations of essential elements were similar to those found in other cetaceans. Non-essential element concentrations were generally comparable to other baleen whales and very low compared to most odontocetes. One deviation from this pattern was a relatively high level of hepatic cadmium that may be related to specific dietary factors of this krill-dependent species. No sex-related differences were found in element concentrations. Hepatic cadmium and mercury increased with animal length, and significant positive interelement correlations were found between copper/zinc and mercury/selenium. We found a mercury-toselenium ratio much lower (1:40) than the frequently reported ratio of one to one. The data suggest that further studies of the postulated mercurydetoxifying role of selenium are warranted.