Spatial and temporal patterns in the calling behavior of beluga whales,Delphinapterus leucas,in Cook Inlet,Alaska

Cook Inlet beluga whales (CIBs) are an endangered population residing in Cook Inlet, Alaska. We characterized the calling behavior of CIBs to improve our understanding of sounds produced by this population. Bottom‐moored hydrophones were deployed at Eagle Bay in summer 2009 and at Trading Bay in summer and winter 2009. CIB sounds were qualitatively analyzed and categorized as a whistle, pulsed call, or click train. A total of 4,097 calls were analyzed, and 66 unique whistle contours were identified. Whistles were quantitatively analyzed using a custom Matlab program. A chi‐square test showed the call category usage at Eagle Bay during summer 2009 and those at Trading Bay during summer 2009 and winter 2009–2010 differed significantly (P < 0.001). Pulsed calls were more common during summer months, and click trains within the frequency band (12.5 kHz) were more common in Eagle Bay. The variation in calling behavior suggests differences in habitat usage or in the surrounding environment, including background noise. With the proposed development projects in Cook Inlet and the potential increase in ambient noise level due to ocean acidification, it is important to understand how this endangered population uses sound, and what anthropogenic factors may influence that use.     

Intraspecific variability in the “vowel”‐like sounds of beluga whales (Delphinapterus leucas): Intra‐ and interpopulation comparisons

The beluga whale (Delphinapterus leucas) has a rich and complicated vocal repertoire. However, different populations use similar and common types of signals. We studied physical features of one of these types, “vowels,” in three Russian populations: the White Sea population (European North), the Chukotka population (the Bering Sea, Chukotka), and the Okhotsk Sea population (Russian Far East) as well as in four summer aggregations of the White Sea belugas over several years in duration. The pulse repetition rate (PRR) at half of the duration of the signal was measured. We found that the PRR of “vowels” collected in the same summer aggregation during different years is stable in time but varies between locations. The degree of variation corresponds with the geographic distance between different locations. Significant differences were discovered between populations separated by thousands of kilometers, and to a lesser extent, between summer aggregations inhabiting different bays of the White Sea. The variation in PRR between the locations can be caused by the divergence of signals owing to the accumulation of random errors during transmission of these signals from generation to generation, which progressed independently in different summer aggregations and populations.     

Mating ecology of beluga (Delphinapterus leucas) and narwhal (Monodon monoceros) as estimated by reproductive tract metrics

Narwhal and beluga whales are important species to Arctic ecosystems, including subsistence hunting by Inuit, and little is understood about their mating ecology. Reproductive tract metrics vary across species in relation to mating strategy, and have been used to infer mating ecology. Reproductive tracts from beluga and narwhal were collected between 1997 and 2008 from five beluga stocks and two narwhal stocks across the Canadian Arctic. Tract length for males and females, relative testes mass for males, and tusk length for male narwhal were measured. We assessed variation relative to species, body size, stock, maturity, and season. Significant variation was found in testes mass across month and stock for beluga, and no significant difference between stock or date of harvest for narwhal. Beluga had significantly larger testes relative to body size than narwhal, suggesting they were more promiscuous than narwhal. A significant relationship was found between narwhal tusk length and testes mass, indicating the tusk may be important in female mate choice. No significant differences were found between narwhal and beluga reproductive tract length for males or females. The mating systems suggested for narwhal and belugas by our results mean the two species may respond differently to climate change.     

Assessing the abundance of Bristol Bay belugas with genetic mark‐recapture methods

The Bristol Bay stock of beluga whales (Delphinapterus leucas) is genetically distinct and resides in Bristol Bay year‐round. We estimated the abundance of this population using genetic mark‐recapture, whereby genetic markers from skin biopsies, collected between 2002 and 2011, were used to identify individuals. We identified 516 individual belugas in two inner bays, 468 from Kvichak Bay and 48 from Nushagak Bay, and recaptured 75 belugas in separate years. Using a POPAN Jolly‐Seber model, abundance was estimated at 1,928 belugas (95% CI = 1,611–2,337), not including calves, which were not sampled. Most belugas were sampled in Kvichak Bay at a time when belugas are also known to occur in Nushagak Bay. The pattern of genetic recaptures and data from belugas with satellite transmitters suggested that belugas in the two bays regularly mix. Hence, the estimate of abundance likely applies to all belugas within Bristol Bay. Simulations suggested that POPAN estimates of abundance are robust to most forms of emigration, but that emigration causes negative bias in both capture and survival probabilities. Because it is likely that some belugas do not enter the sampling area during sampling, our estimate of abundance is best considered a minimum population size.     

Matriarchal genetic population structure of North American beluga whales Delphinapterus leucas (Cetacea: Monodontidae)

The North American beluga whale Delphinapterus leucas population has been divided into a number of putative geographical stocks based upon migration routes and areas of summer concentration. Nucleotide sequences of the mitochondrial DNA (mtDNA) control region were used to assess whether these geographical stocks are genetically distinct. Beluga whale samples from 25 sites were collected primarily from aboriginal subsistence hunts across North America from 1984 to 1994. Thirty-nine mtDNA haplotypes were identified in 628 beluga samples. No differences were found in the distribution of haplotypes between male and female beluga whales at any sampling site. These haplotypes segregated into two distinct assemblages in both a haplotype network and a neighbour-joining tree. The haplotype assemblages had a geographically disjunct distribution that suggests postglacial recolonization of the North American Arctic from two different refugia.
An analysis of molecular variance based on haplotype relationships and frequency indicated genetic heterogeneity among beluga whale summering groups ( P ≤ 0.001). Sequence divergence estimates between sampling sites also indicated geographical differentiation, particularly between samples taken at east Hudson Bay or St Lawrence River and the western or central Arctic. The results of this study show a high degree of philopatry to specific summering areas by this highly mobile animal.     

Seasonal hematology and serum chemistry of wild beluga whales (Delphinapterus leucas) in Bristol Bay, Alaska, USA

We collected blood from 18 beluga whales (Delphinapterus leucas), live-captured in Bristol Bay, Alaska, USA, in May and September 2008, to establish baseline hematologic and serum chemistry values and to determine whether there were significant differences in hematologic values by sex, season, size/age, or time during the capture period. Whole blood was collected within an average of 19 min (range=11-30 min) after the net was set for capture, and for eight animals, blood collection was repeated in a later season after between 80-100 min; all blood was processed within 12 hr. Mean hematocrit, chloride, creatinine, total protein, albumin, and alkaline phosphatase were significantly lower in May than they were in September, whereas mean corpuscular hemoglobin concentration, monocytes, phosphorous, magnesium, blood urea nitrogen, alanine aminotransferase, aspartate aminotransferase, γ-glutamyltranspeptidase, and creatinine kinase were significantly higher. Mean total protein, white blood cell count, neutrophils, and lymphocytes were significantly higher early in the capture period than they were later. No significant differences in blood analyte values were noted between males and females. Using overall body length as a proxy for age, larger (older) belugas had lower white blood cell, lymphocyte, and eosinophil counts as well as lower sodium, potassium, and calcium levels but higher creatinine levels than smaller belugas. These data provide values for hematology and serum chemistry for comparisons with other wild belugas.     

A risk‐management approach to a mycotic disease potential in captive beluga whales (Delphinapterus leucas)

In response to the isolation of Aspergillus fumigatus and other potentially pathogenic fungi in expired air from four of five captive beluga whales (Delphinapterus leucas), a systematic assessment of the magnitude and nature of the risks associated with this finding was conducted. Ten genera of ubiquitous fungi, most of which were potential pathogens, were isolated from the whales over a 16‐month period. During this same period, the whales did not exhibit any behavioral, physical, hematological, or biochemical indications of disease. All the genera of fungi isolated from the whales were also found in air and water samples taken throughout the facility. Opportunities for contamination of samples of expired air were identified. A decision tree, which incorporated subjective probability estimates, was created. The final risk‐management plan involved: 1) alterations to the management of the pool to reduce environmental contamination and opportunities for exposure to fungi, 2) more rigorous quality control for sample collection, and 3) an increased frequency of blowhole cultures. The animals at the facility, including the newborn calf, remained healthy during the 3 years after the risk‐management program. Zoo Biol 18:5–16, 1999. © 1999 Wiley‐Liss, Inc.     

Measurement of cortisol in blow samples collected from free-swimming beluga whales (Delphinapterus leucas)

Blow or respiratory vapor collection is a noninvasive technique for monitoring the physiology of cetaceans and although this technique shows promise, a major challenge of blow collection is difficulty quantifying samples due to variable amounts of seawater contamination. Here, we aimed to (1) determine whether blow samples could be collected from free-swimming belugas, (2) assess whether urea could be used as a dilution marker to normalize blow samples, and (3) determine which factors influenced cortisol and urea concentrations. We collected a total of 252 blow samples from free-swimming belugas in the Churchill River in Manitoba, Canada. Cortisol, an indicator of individual stress and health, was detected in 65.87% of samples with concentrations ranging from 6.73 to 963.17 pg/ml of extract volume, while urea was detected in 90.48% of samples with concentrations ranging from 0.21 to 20.30 mg/L. We were unable to verify the efficacy of urea as a dilution marker to normalize blow samples from free-swimming belugas; however, absolute cortisol concentrations varied based on sample device and quantity rating. Although we demonstrated that blow can be successfully collected from free-swimming belugas, further refinement of this technique is needed before it can be used as a reliable method for physiological assessments.     

Allelic and haplotype variation of major histocompatibility complex class II DRB1 and DQB loci in the St Lawrence beluga (Delphinapterus leucas)

In order to assess levels of major histocompatibility complex (Mhc) variation within the St Lawrence beluga (Delphinapterus leucas) the variation at the beluga Mhc DRB1 class II locus was assessed by single-strand conformation polymorphism (SSCP) analysis of the peptide-binding region for 313 whales collected from 13 sampling locations across North America. In addition, samples from west Greenland and the St Lawrence were also typed at the DQB locus, allowing comparison to a previous study and assessment of linkage disequilibrium of alleles at the two loci. Comparisons of DRB1 and DQB allele frequencies among all sampling locations indicated genetic structure (α < 0.005). Most of this structure resulted from differences between the different wintering groups. Significant genetic structure (α = 0.05) exists among each pair of the following groups at both the DRB1 and DQB loci; St Lawrence, Hudson Strait, Bering Sea, Cunningham Inlet, and Davis Strait (minus Cunningham Inlet), except the St Lawrence and Hudson Strait for the DQB locus. In the St Lawrence population, six of the eight DRB1 alleles are present representing all five known allelic lineages. Evidence of linkage disequilibrium between the DRB1 and DQB is present in two sampling locations, the St Lawrence and Nuussuaq (α = 0.05). Analysis of probable DRB1–DQB haplotypes among groups of beluga suggests a haplotype reduction in the St Lawrence.     

Satellite telemetry reveals population specific winter ranges of beluga whales in the Bering Sea

At least five populations (stocks) of beluga whales (Delphinapterus leucas) are thought to winter in the Being Sea, including the Bristol Bay, Eastern Bering Sea (Norton Sound), Anadyr, Eastern Chukchi Sea, and Eastern Beaufort Sea (Mackenzie) populations. Belugas from each population have been tagged with satellite‐linked transmitters, allowing us to describe their winter (January–March) distribution. The objectives of this paper were to determine: (1) If each population winters in the Bering Sea, and if so, where? (2) Do populations return to the same area each year (i.e., are wintering areas traditional)? (3) To what extent do the winter ranges of different populations overlap? Tagged belugas from all five populations either remained in, or moved into, the Bering Sea and spent the winter there. Each population wintered in a different part of the Bering Sea and populations with multiple years of data (four of five) returned to the same regions in multiple years. When data were available from two populations that overlapped in the same year, they did not occupy the shared area at the same time. Although our sample sizes were small, the evidence suggests belugas from different populations have traditional winter ranges that are mostly exclusive to each population.     

Sexual differences in the foraging ecology of 19th century beluga whales (Delphinapterus leucas) from the Canadian High Arctic

Marine mammals often exhibit significant sexual segregation in their diet and habitat use but these differences have not been studied systematically in historic or ancient populations due to the difficulties associated with determining the sex of skeletal elements based on gross morphology. Using a combined ancient DNA and stable isotope approach, we document a sexual difference in the foraging ecology of late 19th century beluga whales (Delphinapterus leucas) from the Canadian High Arctic. Using two PCR assays that coamplify fragments of the Y-linked SRY and X-linked ZFX genes, we assigned reproducible sex identities to 35 beluga specimens. This provided a basis for investigating sex-specific differences in foraging ecology using stable carbon and nitrogen isotope analyses of bone collagen. These isotopic data demonstrate that although both males and females primarily consumed Arctic cod, males utilized a wider range of prey than females, feeding on high trophic level benthic prey (sculpins) to a greater extent. Because bone collagen integrates prey isotopic compositions over the course of several years these sex-based differences in beluga bone collagen isotopic compositions reflect long-term and sustained sexual differences in foraging.     

Immature euphausiids do not appear to be prey for humpback whales (Megaptera novaeangliae) during spring and summer in Southeast Alaska

Humpbacks whales (Megaptera novaeangliae) have shown a remarkable recovery in the North Pacific, raising concerns regarding their impact on marine communities. In Southeast Alaska, humpbacks feed heavily on euphausiids; however, it remains unclear whether they target immature individuals despite evidence that they do so elsewhere. I evaluate the hypothesis that humpbacks target immature euphausiids in late spring‐summer in Southeast Alaska. Plankton samples were collected at random sites (n = 44) and near whales (n = 53) between 8 June and 9 September 2008 in Frederick Sound and Stephens Passage. The proportion of samples containing immature euphausiids, and immature euphausiid abundance within those samples, were compared between the two sample types. Similar analyses were conducted for adult euphausiids (prey) and calanoid copepods (nonprey) for comparison. I found no statistical difference between the whale and random samples with respect to the occurrence or numerical density of immature euphausiids, which is consistent with the hypothesis that whales did not target them in 2008. Smaller size, insufficient numerical densities and lower energy density of immature euphausiids are suggested as possible reasons. These findings can assist in resolving regional humpback abundance and distribution patterns, and can contribute to an understanding of the trophic interactions characterizing the local ecosystem.     

Patterns and trends in the diet of long‐finned pilot whales (Globicephala melas) in the northeast Atlantic

There is little previous information on feeding habits of long‐finned pilot whales (Globicephala melas) in the northeast Atlantic. The present study analyzed stomach contents of pilot whales stranded in Portugal (n = 6), Galicia (northwest Spain) (n = 32), and Scotland (United Kingdom) (n = 10), from 1990 to 2011. These animals ranged from 213 to 555 cm in length (24 females, 19 males and 5 of unknown sex). The main prey identified were cephalopods of the families Octopodidae and Ommastrephidae, the former being numerically more important in Iberia (Portugal and Galicia) and the latter more important in Scotland, with Iberian whales also showing a more diverse diet. Multivariate analysis revealed evidence of geographical and seasonal variation in diet. Generalized Additive Modeling results indicated that more octopus (Eledone cirrhosa) were eaten in Iberia than in Scotland, more in the first half of the year, and more in larger whales. Numbers of ommastrephid squids in the stomach decreased over the study period and varied with season and whale length. This study confirms cephalopods as the main prey of pilot whales, as previously reported, although our results also suggest that, in the northeast Atlantic, ommastrephid squid are largely replaced as the main prey by octopods at lower latitudes.     

Southern right whales (Eubalaena australis) return to a former wintering calving ground: Fowlers Bay,South Australia

Southern right whales (SRW), Eubalaena australis, have reoccupied historically important winter habitat ranges (calving grounds) in recent years along the southern Australian coast. Here we present findings of increased abundance of SRW at Fowlers Bay, South Australia, a previous shore‐based whaling station. This study investigates: SRW inter‐ and intraseasonal trends in relative abundance; changes to the relative proportion of the southwestern subpopulation represented by SRW at Fowlers Bay; distribution; and occupancy. Sighting and photo identification data were collected during annual aerial (1993–2016) and vessel surveys (2014–2016). The total number of female and calf pairs was 3 during 1993–2003 and 63 during 2004–2014. Despite high variability in annual relative abundance, the rate of mean increase from 1993 to 2016 (29.0%/yr, 95% CI = 0, 54.2) exceeded the maximum biological rate for the species (6%–7%/yr). Peak relative abundance was recorded in July and August. SRW at Fowlers Bay represent an increasing proportion of the southwestern subpopulation (range = 0.9%–7.4%). Mean occupancy was 23 d (range = 1–75) for female and calf pairs and 2 d (range = 1–15) for unaccompanied adults. Reduced sightings in 2015 and 2016 demonstrate plasticity in SRW abundance at Fowlers Bay. Research into the movement and connectivity of SRW is needed to understand drivers of habitat dispersal in Australia.     

Assessment of wound healing of tagged gray (Eschrichtius robustus) and blue (Balaenoptera musculus) whales in the eastern North Pacific using long‐term series of photographs

Tags have been used to examine migration routes and habitat use of large whales for >40 yr, however, evaluation of tag wound healing has largely been short‐term, anecdotal or generalized. This study developed methods for systematic photographic assessment of long‐term external consequences of tag placement, to determine potential differences in wound healing between species and tag types and thus advise future tagging efforts to possibly minimize undesirable side effects. Tag site appearance and healing characteristics were evaluated by two reviewers and a time series evaluated by five veterinarians from photographs during 995 postdeployment encounters with 34 gray and 63 blue whales tagged in the North Pacific. Blue whale resightings were less frequent, but spanned a longer time period due to earlier tag deployments than the more frequent gray whale follow‐up observations. Swelling occurred in 74% of reencountered gray whales, with the highest frequency 6 mo postdeployment. Swellings were common in blue whales with early tag designs but rare with current models. Depressions occurred in 82% of gray and 71% of blue whales. This study demonstrates the value of follow‐up studies of tagged animals and systematic scoring of photographs to quantitatively compare tag response.     

Behavioral and acoustical characteristics of the reproductive gathering of beluga whales (Delphinapterus leucas) in the vicinity of Myagostrov, Golyi Sosnovets, and Roganka Islands (Onega Bay, the White Sea)

The structure of the summer reproductive gathering of beluga whales Delphinapterus leucas was studied in the vicinity of Myagostrov, Golyi Sosnovets, and Roganka islands (Onega Bay, the White Sea) in 2006 and 2008. The abundance, age and sex structure, behavior, and swimming and acoustic behavior were studied in detail.     

Development of predictive models for determining fetal age‐at‐length in belugas (Delphinapterus leucas) and their application toward in situ and ex situ population management

For wild belugas (Delphinapterus leucas), gestation length estimates based on fetal size have produced extreme ranges. Ex situ populations thereby provide unique opportunities to define this important life history event. Accordingly, research with ultrasound was conducted on six beluga whales over 11 gestations with known conception dates to serially measure fetal changes in biparietal diameter (BP), thoracic diameter (TD), thoracic circumference (TC), and total length (TL). Incremental polynomial regression analyses were performed on each fetal measurement to develop predictive models for determining age based on fetal size and days from parturition. Gestation length (n = 11) was a mean 467 ± 5.4 d with male calves (478 ± 8.6 d) experiencing a longer gestation (P = 0.04) than females (457 ± 3.9 d). Age at TL was best described using a 2nd order polynomial model, while linear relationships existed for BPD, TD, and TC. Accuracy was improved for predicting age (P = 0.001) or days prior to parturition (P = 0.038) using data from the first vs. the second half of gestation. The results provide accurate models for aging beluga fetuses based on size in both in situ and ex situ populations.