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.     

Entanglement‐scar acquisition rates and scar frequency for Bering‐Chukchi‐Beaufort Seas bowhead whales using aerial photography

The Bering‐Chukchi‐Beaufort Seas (BCBS) bowhead whale (Balaena mysticetus) has been considered at low‐risk for entanglement injuries and ship strikes because their range is mainly north of commercial fisheries; nevertheless, changes to their arctic habitat, including a longer open water period and declining sea ice, have resulted in increasing commercial activity and concern about fisheries interactions. We examined interyear matches (between 1985 and 2011) from a photo identification project and identified whales that had acquired entanglement injuries. We estimated the probability of a bowhead acquiring an entanglement injury using two statistical methods: interval censored survival analysis and a simple binomial model. Both methods give similar results, suggesting a 2.2% (95% CI: 1.1%–3.3%) annual probability of acquiring a scar. We also include an entanglement scar frequency analysis of aerial photographs from the 2011 spring and fall surveys near Point Barrow, Alaska, which suggest 12.4% of live bowheads show evidence of entanglement scarring. Entanglement rates for the BCBS bowhead stock are lower than many other large whale stocks, and abundance has increased over the past 35 yr; however, our findings indicate that fishing gear entanglement is a more serious concern for the BCBS bowhead whale population than previously thought.     

Non-lethal entanglement of humpback whales (Megaptera novaeangliae) in fishing gear in northern Southeast Alaska

Aim Entanglement in fishing gear is recognized as a potentially significant source of serious injury and mortality for humpback whales (Megaptera novaeangliae) in some parts of their range. In recent years, the number of humpback whales reported to have been entangled in Alaska has increased. In 2003–04 we quantified the prevalence of non‐lethal entanglements of humpback whales in northern Southeast Alaska (SEAK) with the ultimate goal of informing management discussions of the entanglement issue for the Central North Pacific stock of humpback whales. Location The near‐shore waters of northern Southeast Alaska. Methods We photographed individual humpback whales’ caudal peduncles as they dived and then examined the photographs for scars indicative of a previous entanglement. Results The percentage of whales assessed to have been non‐lethally entangled at some time in their lives ranged from 52% (minimal estimate) to 71% (conditional estimate) to 78% (maximal estimate). Of these, the conditional estimate is recommended because it is based solely on unambiguous scars. Eight per cent of the whales in one portion of the study area (Glacier Bay/Icy Strait) acquired new entanglement scars between 2003 and 2004, although the sample size was small. Calves were less likely than older whales to have entanglement scars, and males may be at higher risk than females. Whales with more photographs and/or photographic coverage may be more likely to be assessed as having been entangled than whales with fewer photographs and/or coverage. Main conclusions Caudal peduncle scars reveal that the majority of humpback whales in northern SEAK have been entangled. Comparison with statistics on reported entanglements suggests that most whales apparently shed the gear on their own, unless humans are disentangling whales much more often than is reported. While cumulative estimates of the percentage of whales with entanglement scars (e.g. the conditional estimate) provide useful baseline information, future efforts should focus on monitoring the annual rate of entanglement scar acquisition as a more powerful measure of contemporary entanglement rates. Our findings indicate that entanglement of humpback whales in fishing gear in SEAK is a management issue warranting increased attention. A proactive approach is needed to address the problem and to identify and implement preventive measures.     

Presence and behavior of bowhead whales (Balaena mysticetus) in the Alaskan Beaufort Sea in July 2011

The Western Arctic bowhead whale (Balaena mysticetus) is highly adapted to sea ice and annually migrates through the Bering, Chukchi, and Beaufort seas. While the overall distribution and seasonal movements of bowhead whales are mostly understood, information about their distribution in the Alaskan Beaufort Sea in early to mid-summer has not been well documented. In July 2011, we conducted an exploratory flight in the Alaskan Beaufort Sea, north of Camden Bay (71°N 144°W), near the location of a single satellite-tagged bowhead whale. Eighteen bowhead whales were observed, and behavior consistent with feeding was documented. To our knowledge, this is the first documentation of behavior consistent with feeding north of Camden Bay in mid-July. Few studies have focused on bowhead whale distribution in the Alaskan Beaufort Sea in early to mid-summer, and no long-term, region-wide surveys have been conducted during summer. Bowhead whales are already exposed to anthropogenic disturbance in the Canadian Beaufort Sea in summer, the Alaskan Beaufort Sea in fall, and the Chukchi and Bering seas from fall through spring. The presence of bowhead whale aggregations in the Alaskan Beaufort Sea in summer should be considered when assessing the cumulative effects of human-related activities.     

Photo‐identification matches of humpback whales (Megaptera novaeangliae) from feeding areas in Russian Far East seas and breeding grounds in the North Pacific

Humpback whales migrate seasonally from high latitude feeding areas to lower latitude breeding areas for mating and calving. In 2004–2006, a North Pacific basin‐wide study called SPLASH was conducted as an international collaboration among various groups of researchers. The Russian Far East consists of multiple high latitude feeding areas and during SPLASH, 102 whales were identified and compared to catalogs from breeding areas. Our goal in this study was to further investigate the migratory destinations of whales from the Russian Far East using a total of 1,459 photographs of whales identified from 2004 to 2014. We compared the latest Russian catalog with the SPLASH catalog from wintering areas (2004–2006) and with two additional regional catalogs from Okinawa (1989–2006) and the northern Philippines (2000–2006). We found a total of 152 matches: 106 with Asian, 35 with Hawaiian, and 11 with Mexican breeding grounds. The match rate was higher in mainland Kamchatka and consisted mostly of whales from the Asian breeding ground. In the Commander Islands, the proportion of whales from Asia was twice that of Hawaii and six times higher than Mexico. The total match rate was low, supporting the hypothesis of some undiscovered humpback whale breeding location in the North Pacific.     

Two historical weapon fragments as an aid to estimating the longevity and movements of bowhead whales

The age of bowhead whales captured by Native Alaskan hunters in the Bering, Chukchi and Beaufort Seas has been estimated via chemical analyses of the eye lenses, and other techniques. The racemization-age estimates indicate that bowhead whales (Balaena mysticetus) have a lifespan of more than a century. Stone and ivory weapon fragments recovered from bowhead whales hunted in Wainwright and Barrow (Alaska) in 1981, 1992, 1993 and 1997, provided rough but independent assessments of the whales’ longevity; however, their date of manufacture was unknown. Adding further confirmation of these age estimates, this note describes bomb lance fragments recovered recently (2007) and about 30 years ago (1980) from bowhead whales harvested by Eskimo hunters that were “dateable” and likely manufactured between 1879 and 1885.     

APPARENT LATERALIZED BEHAVIOR IN GRAY WHALES FEEDING OFF THE CENTRAL BRITISH COLUMBIA COAST

A digital acoustic recording tag was used to examine the 3‐D orientation of gray whales feeding along the central British Columbia coast. A total of 96 feeding dives were recorded from six different whales. More than half (53.1%) of the whales' bottom time was spent rolled at an angle greater than 45°. Whales rolled an average of 2.9 times per feeding dive, and rolling behavior was often accompanied by a negative pitch angle. Out of 282 recorded rolls, 274 (97.2%) were to the right. Likewise, 98.5% of the total time spent rolled at an angle greater than 45° was spent rolled to the right. The gray whales in this study showed a significant right‐side bias on both an individual (P≤ 0.009) and group level (P < 0.001). Based on the findings of this study and previous reports of uneven baleen wear ( Kasuya and Rice 1970 ), it is proposed that gray whales exhibit a population‐wide right‐side rolling bias similar in character to the 90/10 split of right handedness in humans.     

Stable carbon and nitrogen isotope ratios in muscle and epidermis of arctic whales

Collection of minimally invasive biopsy samples has become an important method to establish normal stable isotopes reference ranges in various wildlife species. Baseline data enhance the understanding of feeding ecology, habitat use, and potential food limitation in apparently healthy, free‐ranging cetaceans. Epidermis and muscle were collected from subsistence‐hunted northern Alaskan bowhead (n= 133 epidermis/134 muscle) and beluga whales (n= 42/49) and subsistence‐hunted Russian gray whales (n= 25/17). Additional samples were obtained from gray whales stranded in California (n= 18/11) during mortality events (1999, 2000). Both δ¹⁵N and δ¹³C are trophic position and benthic/pelagic feeding indicators, respectively, in muscle and epidermis. Epidermis is generally enriched in ¹⁵N over muscle, while epidermal ¹³C is more depleted. Lipid extraction does not alter δ¹⁵N in either tissue, but affects epidermal δ¹³C. Nitrogen‐15 is enriched in muscle, but not epidermis of stranded compared to subsistence‐hunted gray whales, indicating probable protein catabolism and nutritional stress in stranded whales. Similarly, epidermal δ¹³C of harvested whales is lower than in stranded whales, suggesting depleted lipid stores and/or food limitation in stranded animals. Epidermal isotope signatures are similar in both present‐day bowheads and in an ancient sample from the Northern Bering Sea region. Although only one specimen, this suggests trophic level of the ancient whale compares to modern bowheads after a millennium.     

Blubber fatty acid composition of bowhead whales, Balaena mysticetus: Implications for diet assessment and ecosystem monitoring

Fatty acids (FA) have a diversity of structures that are transferred with little modification through food webs, making them valuable in assessing diets of animals that cannot be directly observed feeding. Before using FA to estimate diets, it is necessary to evaluate variation in FA signatures within and among individuals of a given species. To begin assessing diets and foraging of western Arctic bowhead whales (Balaena mysticetus), we examined the FA in blubber of 64 bowheads taken in the spring and fall subsistence hunts in 1997–2002 at Barrow and Kaktovik, Alaska. We found no significant differences in FA characteristics of inner blubber layers taken from either duplicate samples on the dorsal surface, or between dorsal and ventral sites. Significant differences were found in the FA composition between inner and outer layers of blubber at the same body site. We also found age, season and year to have significant effects on FA composition; however, gender was not found to be significant. While the importance of the Beaufort Sea as a feeding ground of bowhead whales remains uncertain, our results indicate that adults and sub-adults foraged to some extent on different prey and that both age classes consumed copepods there in summer at sufficient levels to significantly alter their blubber FA profiles. Both of these findings correspond with dietary conclusions reached from the analysis of stomach contents. Furthermore, we found compelling evidence that yearly variation in bowhead FA reflect changes in FA compositions of phytoplankton at the base of the food web, probably in response to climate variation. Variability in phytoplankton-derived FA in blubber was correlated significantly with yearly mean values of the Pacific Decadal Oscillation. FA in bowhead whale blubber, therefore, might be used to monitor effects of climate change on lower trophic levels and production processes in the western Arctic.     

A simple photograph‐based approach for discriminating between free‐ranging long‐finned (Globicephala melas) and short‐finned (G. macrorhynchus) pilot whales off the east coast of the United States

Line transect based abundance estimation is complicated for long‐finned (LFPW, Globicephala melas) and short‐finned (SFPW, G. macrorhynchus) pilot whales because of their similarity in appearance and their overlapping summertime range in some areas. We developed a photograph‐based approach to distinguish between species of free‐ranging pilot whales in the northwest Atlantic. We collected skin samples and photographs during the summers of 2004–2007 and used skin samples to distinguish species based on mitochondrial DNA. Relative morphometric measurements from photographs were examined using mixed‐effect models and logistic regression. The best model among 94 candidate models had an overall classification error rate of 2.5%. We tested the presence/absence of pigmentation in four regions of the dorsal body (melon, eye, cape, and saddle) for differences. Pigmentation was present in all four regions in 100% of the SFPWs sampled. Melon patch, blaze, and saddle patch pigmentation were present in 6%, 68%, and 50%, respectively, of the LFPWs, but the cape was completely absent. Both types of analyses provided positive species discrimination of free‐ranging animals. We created a cost‐effective, simple tool which could ultimately assist in providing appropriate management, mitigation, and conservation strategies for both northwest Atlantic species of pilot whales.     

Population trends for humpback whales (Megaptera novaeangliae) foraging in the Francisco Coloane Coastal‐Marine Protected Area,Magellan Strait,Chile

In 2003 a feeding aggregation of southeastern Pacific humpback whales (Megaptera novaeangliae) was reported in the Magellan Strait. While Chile established its first marine national park in the Strait to protect humpback whale habitat, fatal ship strikes remain a concern because of overlap with a busy shipping lane. To better understand population risk, we estimated abundance and survival for this population using Bayesian robust‐design mark‐recapture models fit to photographic data from 2004 to 2016. Overall, the model estimated a total of 204 whales (95% CI: 199–210) during the last 12 yr, and 93 (95% CI: 86–100) in the 2016/2017 austral summer. The population grew at 2.3% (CI: 2.1%–3.1%), an annual increase of two whales. Annual survival (including calves) was estimated at 0.892 (CI: 0.871–0.910). Our results corroborate a persistent feeding population, but one that is increasing relatively slowly. Owing to its vulnerability stemming from its small size, coupled with significant overlap with a busy shipping lane, we argue this subpopulation is at significant risk from ship strikes and may be one of the few populations where anthropogenic mortalities could regulate population dynamics. We therefore encourage continued monitoring via photographic mark‐resighting surveys, and analyses explicitly investigating potential population‐level ship strike effects.     

Rorqual whale (Balaenopteridae) surface lunge‐feeding behaviors: Standardized classification,repertoire diversity,and evolutionary analyses

Rorqual whales (Family: Balaenopteridae) are the world's largest predators and sometimes feed near or at the sea surface on small schooling prey. Most rorquals capture prey using a behavioral process known as lunge‐feeding that, when occurring at the surface, often exposes the mouth and head above the water. New technology has recently improved historical misconceptions about the natural variation in rorqual lunge‐feeding behavior yet missing from the literature is a dedicated study of the identification, use, and evolution of these behaviors when used to capture prey at the surface. Here we present results from a long‐term investigation of three rorqual whale species (minke whale, Balaenoptera acutorostrata; fin whale, B. physalus; and blue whale, B. musculus) that helped us develop a standardized classification system of surface lunge‐feeding (SLF) behaviors. We then tested for differences in frequency of these behaviors among the three species and across all rorqual species. Our results: (1) propose a unified classification system of six homologous SLF behaviors used by all living rorqual whale species; (2) demonstrate statistically significant differences in the frequency of each behavior by minke, fin, and blue whales; and (3) provide new information regarding the evolution of lunge‐feeding behaviors among rorqual whales.     

Inferring trackline detection probabilities,g(0), for cetaceans from apparent densities in different survey conditions

Visual line‐transect surveys are commonly used to estimate cetacean abundance. A key parameter in such studies is g(0), the probability of detecting an animal that is directly on the transect line. This is typically considered to be constant for a species across survey conditions. A method is developed to estimate the relative values of g(0) in different survey conditions (Beaufort state) by comparing Beaufort‐specific density estimates. The approach is based on fitting generalized additive models, with the presence of a sighting on a survey segment as the dependent variable, Beaufort state as the key explanatory variable, and year, latitude, and longitude as nuisance variables to control for real differences in density over time and space. Values of relative g(0) are estimated for 20 cetacean taxa using 175,000 km of line‐transect survey data from the eastern and central Pacific Ocean from 1986 to 2010. Results show that g(0) decreases as Beaufort state increases, even for visually conspicuous species. This effect is greatest for the least conspicuous species (rough‐toothed dolphins, beaked whales, minke whales, and dwarf and pygmy sperm whales). Ignoring these large effects results in a nontrivial bias in cetacean abundance estimates.     

ABUNDANCE AND POPULATION TREND (1978-2001) OF WESTERN ARCTIC BOWHEAD WHALES SURVEYED NEAR BARROW, ALASKA

The 2001 survey of western Arctic (Bering, Chukchi, and Beaufort seas) bowhead whales was conducted from 5 April to 7 June near Barrow, Alaska. Visual observers recorded a total of 3,295 “new” (not seen before) and 532 “conditional” (possibly seen before) whales in 1,130 h of watch effort, including 121 new calves (3.7% of the new whales). Concurrent with the visual survey, passive acoustic surveillance was conducted almost continuously from 16 April to 31 May, resulting in 27,023 locations of vocalizing bowhead whales. The estimated number of whales within 4 km of the perch (N₄) was 7,025 (SE = 1,068). The estimated proportion of the whales within 4 km of the perch (P₄) was 0.862 (SE = 0.044, computed by a moving blocks bootstrap). Combining these, the abundance estimate (N₄/P₄) for 2001 is 10,470 (SE = 1, 351) with a 95% confidence interval of 8, 100–13, 500. The estimated annual rate of increase (ROI) of the population from 1978 to 2001 is 3.4% (95% CI 1.7%‐5%). Reports from hunters and results of an aerial survey in June 2001 indicate whales continued to pass Barrow after the survey had ended. In 2001 51% (572 h) of the watch was scored as occurring during “fair‐excellent” visibility conditions, somewhat lower than the average for all surveys since 1978. Sea ice in the leads and fog were the principle environmental factors affecting visibility for all years. The estimated rate of increase and the fact that the number of calves counted in 2001 is the highest ever recorded suggest a steady recovery of this population. Other populations of large balaenids, notably the North Atlantic right whale, have failed to recover despite 70 yr of protection. The recovery of the howhead whale is likely attributable to low anthropogenic mortality, a relatively pristine habitat, and a well‐managed subsistence hunt. Nonetheless, offshore oil development, increasing shipping traffic, changes in the Bering Sea ecosystem, sea ice retreat, and possibly killer whale predation within its range could impact this bowhead population and should be carefully monitored.     

Evidence of two subaggregations of humpback whales on the Kodiak,Alaska, feeding ground revealed from stable isotope analysis

Knowledge of humpback whale (Megaptera novaeangliae) foraging on feeding grounds is becoming increasingly important as the growing North Pacific population recovers from commercial whaling and consumes more prey, including economically important fishes. We explored spatial and temporal (interannual, within‐season) variability in summer foraging by humpback whales along the eastern side of the Kodiak Archipelago as described by stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope ratios of humpback whale skin (n = 118; 2004–2013). The trophic level (TL) of individual whales was calculated using basal food web δ¹⁵N values collected within the study area. We found evidence for the existence of two subaggregations of humpback whales (“North,” “South”) on the feeding ground that fed at different TLs throughout the study period. Linear mixed models suggest that within an average year, Kodiak humpback whales forage at a consistent TL during the feeding season. TL estimates support mixed consumption of fish and zooplankton species in the “North” (mean ± SE; 3.3 ± 0.1) and predominant foraging on zooplankton in the “South” (3.0 ± 0.1). This trend appears to reflect spatial differences in prey availability, and thus, our results suggest North Pacific humpback whales may segregate on feeding aggregations and target discrete prey species.     

FEEDING, SOCIAL AND MIGRATION BEHAVIOR OF BOWHEAD WHALES, BALAENA MYSTICETUS, IN BAFFIN BAY VS. THE BEAUFORT SEA—REGIONS WITH DIFFERENT AMOUNTS OF HUMAN ACTIVITY

This paper compares the behavior of bowhead whales of the Davis Strait/Baffin Bay stock, as observed along the east coast of Baffin Island in 1979–1986, with behavior of the Bering/Chukchi/Beaufort Sea stock observed in the Beaufort Sea in 1980–1986. All data used here were collected during late summer and early autumn in the absence of acute human disturbance. The behavioral repertoires of the two populations were similar. However, quantitative differences were found for whales engaged in all three activities studied: (1) Bowheads feeding in deep water off Isabella Bay, Baffin Island, had longer dives and surfacings, on average, than noted for bowheads feeding in the Beaufort Sea. (2) Among whales socializing in shallow water, we saw sexual interactions more often at Isabella Bay than in the Beaufort Sea. Calls emitted by socializing whales off Baffin Island were similar to those heard in the Chukchi and Beaufort Seas. However, pulsed tonal calls were longer off Baffin Island, and previously undescribed mechanical "crunch" sounds were recorded there near socializing bowheads. (3) During autumn migration, "fluke-out" dives were less common, and dive durations were longer, in the Beaufort Sea than off Baffin Island (P<0.001). Multivariate and other analyses indicated that some but not all differences can be ascribed to regional differences in the natural environment or in whale activities, However, during 1974–1986, Bering/Chukchi/Beaufort bowheads were exposed to more industrial, hunting and other human activity than Davis Strait/Baffin Bay bowheads. The "inconspicuous" behavior during autumn migration in the Beaufort may have been attributable to human activities, but causative links cannot be isolated.     

Decadal shifts in autumn migration timing by Pacific Arctic beluga whales are related to delayed annual sea ice formation

Migrations are often influenced by seasonal environmental gradients that are increasingly being altered by climate change. The consequences of rapid changes in Arctic sea ice have the potential to affect migrations of a number of marine species whose timing is temporally matched to seasonal sea ice cover. This topic has not been investigated for Pacific Arctic beluga whales (Delphinapterus leucas) that follow matrilineally maintained autumn migrations in the waters around Alaska and Russia. For the sympatric Eastern Chukchi Sea (‘Chukchi’) and Eastern Beaufort Sea (‘Beaufort’) beluga populations, we examined changes in autumn migration timing as related to delayed regional sea ice freeze‐up since the 1990s, using two independent data sources (satellite telemetry data and passive acoustics) for both populations. We compared dates of migration between ‘early’ (1993–2002) and ‘late’ (2004–2012) tagging periods. During the late tagging period, Chukchi belugas had significantly delayed migrations (by 2 to >4 weeks, depending on location) from the Beaufort and Chukchi seas. Spatial analyses also revealed that departure from Beaufort Sea foraging regions by Chukchi whales was postponed in the late period. Chukchi beluga autumn migration timing occurred significantly later as regional sea ice freeze‐up timing became later in the Beaufort, Chukchi, and Bering seas. In contrast, Beaufort belugas did not shift migration timing between periods, nor was migration timing related to freeze‐up timing, other than for southward migration at the Bering Strait. Passive acoustic data from 2008 to 2014 provided independent and supplementary support for delayed migration from the Beaufort Sea (4 day yr^?1) by Chukchi belugas. Here, we report the first phenological study examining beluga whale migrations within the context of their rapidly transforming Pacific Arctic ecosystem, suggesting flexible responses that may enable their persistence yet also complicate predictions of how belugas may fare in the future.     

Shallow and deep lunge feeding of humpback whales in fjords of the West Antarctic Peninsula

Humpback whales (Megaptera novaeangliae) belong to the class of marine mammals known as rorquals that feed through extraordinarily energetic lunges during which they engulf large volumes of water equal to as much as 70% of their body mass. To understand the kinematics of humpback lunge feeding, we attached high‐resolution digital recording tags incorporating accelerometers, magnetometers, pressure and sound recording to whales feeding on euphausiids in fjords of the West Antarctic Peninsula. Instances of near vertical lunges gave us the unique opportunity to use the signal from the accelerometer to obtain a fine scale record of the body accelerations involved in lunging. We found that lunges contain extreme accelerations reaching 2.5 m/s² in certain instances, which are then followed by decelerations. When animals are intensively feeding the inter‐lunge interval is similar for both deep and shallow lunges suggesting a biomechanical constraint on lunges. However, the number of lunges per dive varies from one for shallow feeding (<25 m) to a median of six for deeper dives. Different feeding patterns were evident in the kinematic record, for deep and shallow feeding bouts with the much greater mean turn rates occurring in shallow feeding.