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As a precautionary measure to protect the culturally-significant bowhead whale ( Balaena mysticetus ) from vessel strike, this paper explores the potential for having the International Maritime Organization endorse ships׳ routeing measures in the Bering Strait region. The confined nature of the Bering Strait forces both vessel traffic and the migrating Bering–Chukchi–Beaufort stock of bowhead whales to occupy the same narrow space, thus putting individual animals at risk of vessel strike. The potential for vessel strike may become exacerbated as the reduction of Arctic sea ice makes the Arctic increasingly accessible, allowing vessels to transit through the Strait in greater densities. In drawing from lessons learned in the successful reduction of vessel strike to the North Atlantic right whale ( Eubalaena glacialis ), the author provides broad recommendations for implementing ships׳ routeing measures in the Bering Strait region based on findings by Citta et al. (2012). It is recommended that both the western side of Big Diomede Island and the western side of St. Lawrence Island be designated as seasonal Areas To Be Avoided (ATBA) during fall months when bowheads are known to inhabit the area. Alternatively, it is proposed that Traffic Separation Schemes (TSSs) be implemented so that vessels must keep to the eastern side of both Big Diomede Island and St. Lawrence Island during this time. Both measures would reduce the amount of overlap between migrating bowhead whales and transiting vessel traffic, effectively reducing the potential for vessel strike.

Mammals vary dramatically in lifespan, by at least two-orders of magnitude, but the molecular basis for this difference remains largely unknown. The bowhead whale Balaena mysticetus is the longest-lived mammal known, with an estimated maximal lifespan in excess of two hundred years. It is also one of the two largest animals and the most cold-adapted baleen whale species. Here, we report the first genome-wide gene expression analyses of the bowhead whale, based on the de novo assembly of its transcriptome. Bowhead whale or cetacean-specific changes in gene expression were identified in the liver, kidney and heart, and complemented with analyses of positively selected genes. Changes associated with altered insulin signaling and other gene expression patterns could help explain the remarkable longevity of bowhead whales as well as their adaptation to a lipid-rich diet. The data also reveal parallels in candidate longevity adaptations of the bowhead whale, naked mole rat and Brandt's bat. The bowhead whale transcriptome is a valuable resource for the study of this remarkable animal, including the evolution of longevity and its important correlates such as resistance to cancer and other diseases.

The Spitsbergen stock of bowhead whales (Balaena mysticetus) is considered Critically Endangered by IUCN. Over recent decades, there have been only a few sightings, and very few biological samples are available for molecular analyses. Hence, genetic diversity of the extant Spitsbergen stock is unknown. Here, we present mitochondrial genomes from eight skin biopsy samples that were collected in 2006 and 2010. There were three different haplotypes, two of which have not previously been detected. Six samples shared the same haplotype, indicating that they were obtained from closely related whales, or possibly the same individual. Average nucleotide diversity was π = 0.0037, with a total of 93 variable positions among the haplotypes.

In 1941, Abel establishedBalaena belgicabased on a series of fused cervical vertebrae and citing other cranial fragments from the late Neogene of the Antwerp harbor (northern Belgium). Later, Plisnier-Ladame & Quinet (1969) added a neurocranium and other skeletal remains from the same area to this species. Recently, the neurocranium was re-assigned to the genusEubalaenathanks to newer phylogenetic analyses. Here, a new description is provided of materials previously assigned to “Balaena”belgicatogether with taxonomic revisions. Our work suggests that the cervical complex originally designated as the type of “Balaena”belgicais too poorly preserved to be used as such and is assigned to Balaenidae gen. et sp. indet., thus making “Balaena”belgicaa nomen dubium. In addition to the neurocranium, the other remains consist in a fragment of maxilla assigned to Balaenidae gen. et sp. indet. and in a humerus assigned toEubalaenasp. Discovered in the Kruisschans Sands Member of the Lillo Formation (3.2–2.8 Ma, Piacenzian, Late Pliocene), the neurocranium is designated as the holotype of the new speciesEubalaena ianitrix. Our phylogenetic analysis supports a sister-group relationship ofEubalaena ianitrixandEubalaena glacialis, and helps constraining the ages of origin for balaenid clades. Ecological and phylogenetic data suggest thatEubalaena ianitrixmay represent the direct ancestor ofEubalaena glacialis, the latter having evolved through phyletic transformation including body size increase during the temperature decline of the Late Pliocene.

During summer 2012 Shell performed exploratory drilling at Sivulliq, a lease holding located in the autumn migration corridor of bowhead whales (Balaena mysticetus), northwest of Camden Bay in the Beaufort Sea. The drilling operation involved a number of vessels performing various activities, such as towing the drill rig, anchor handling, and drilling. Acoustic data were collected with six arrays of directional recorders (DASARs) deployed on the seafloor over ~7 weeks in Aug-Oct. Whale calls produced within 2 km of each DASAR were identified and localized using triangulation. A "tone index" was defined to quantify the presence and amplitude of tonal sounds from industrial machinery. The presence of airgun pulses originating from distant seismic operations was also quantified. For each 10-min period at each of the 40 recorders, the number of whale calls localized was matched with the "dose" of industrial sound received, and the relationship between calling rates and industrial sound was modeled using negative binomial regression. The analysis showed that with increasing tone levels, bowhead whale calling rates initially increased, peaked, and then decreased. This dual behavioral response is similar to that described for bowhead whales and airgun pulses in earlier work. Increasing call repetition rates can be a viable strategy for combating decreased detectability of signals arising from moderate increases in background noise. Meanwhile, as noise increases, the benefits of calling may decrease because information transfer becomes increasingly error-prone, and at some point calling may no longer be worth the effort.

The expansion of hydrocarbon exploration in northwest Greenland has made it increasingly important to understand the occurrence of marine mammals in the region. We describe the seasonal occurrence of marine mammals and the spatial distribution of their calls in Baffin Bay and Melville Bay. Four Autonomous Multichannel Acoustic Recorders (AMARs) were deployed during summer 2012 (late July to early October), five recorders during September 2013, and two recorders from late September 2013 to early September 2014. The call presence of several species was analyzed using automatic call detection and manual verification analysis methods. A novel approach to discern narwhal (Monodon monoceros) clicks from beluga (Delphinapterus leucas) clicks was implemented during the verification process. Narwhal calls were detected in spring and fall, showing a south-to-north migration pattern in spring and a north-to-south migration pattern in fall. Few beluga whales were detected during fall 2013 and spring 2014. Bearded seal (Erignathus barbatus) calls were detected mainly during spring (mating period). A small number of bowhead whale calls (Balaena mysticetus) were detected during fall 2013 and spring and summer 2014. For the first time at this latitude in Baffin Bay, long-finned pilot whales (Globicephala melas) and sperm whales (Physeter macrocephalus) were detected during summer and fall. Our results suggest that the presence of marine mammals in Baffin Bay and Melville Bay is governed mainly by the annual cycle of sea ice formation and decay.

We analyzed scarring data for Bering-Chukchi-Beaufort (BCB) Seas bowhead whales (Balaena mysticetus) harvested by Alaska Native hunters to quantify the frequency of line entanglement, ship strikes, and killer whale-inflicted injuries. We had 904 records in our database for whales landed between 1990 and 2012, and after data quality screening, we found 521 records containing information on scarring. Logistic regression was used to evaluate different combinations of explanatory variables (i.e., body length, sex, year, year-group) to develop a prediction model for each scar type. We also list bowhead whales that were harvested, found dead, or observed alive entangled in commercial line/fishing gear. Our findings suggest that about 12% of harvested bowheads show entanglement scars. Their frequency is highly correlated with body length and sex: about 50% of very large bowheads (> 17 m) show such scars, while whales under 9 m rarely do, and males show a significantly higher rate than females. Scars associated with ship strikes are infrequent and occur on ~2% of all harvested whales; body length, sex, and year were not significant factors. Scarring from attempted killer whale predation was evident on ~8% of landed whales. As with entanglement injuries, the frequency of killer whale scars was much higher (> 40%) on whales more than 16 m in length and statistically more frequent in the second half of the study (2002 – 12). Increased killer whale injuries in the recent decade are consistent with studies conducted on bowheads of the Eastern Canada-West Greenland population. The findings presented here reflect the most thorough analysis of injury rates from entanglement, ships, and killer whales for the BCB bowheads conducted to date. They indicate that (1) entanglement rates primarily from pot fishing gear (crab or cod or both) are relatively high for very large and presumably older bowheads, (2) collisions with ships are infrequent at present, and (3) scarring from killer whales is frequent on very large adult whales (> 17 m). Considering that bowhead habitat is changing rapidly (e.g., sea ice reduction), industrial ship traffic in the Arctic is increasing, and commercial fishing operations are expanding to the north, we strongly recommend that monitoring of scarring and injuries on harvested bowheads continue into the future as a means of documenting change.

Reports of bowhead whales (Balaena mysticetus) outside the Arctic Circle are scarce. On 15 May 2016, a juvenile bowhead whale was recorded in shallow water in Mount's Bay (Cornwall, UK) much further south than the species' normal dis- tribution. Fifteen months earlier, another such sighting was made involving a juvenile bowhead whale in the shallow water of an offshore island, St Martin's (Isles of Scilly, Cornwall), only 60 km from Mount's Bay. Other observations of suspected/confirmed immature bowhead whales include (1) an unconfirmed sighting involving a whale without a dorsal fin off Cornwall (13 May 2016), (2) a confirmed sighting off France (10 May 2016), and (3) a confirmed sighting in Carlingford Lough (border of Northern Ireland/ Republic of Ireland; 29 May 2016). These extra- limital encounters present the first confirmed sightings of bowhead whales within temperate European waters. Apart from the whale sighted in 2015, the sightings in 2016 probably all involve the same juvenile whale. Indeed, photographs depicting natural scars/marks suggest that the Carlingford Lough bowhead whale is the same individual as the Mount's Bay. The whale was displaying behaviour indicative of ram feeding in a sheltered, shallow bay (< 10 m water depth). Do these records present vagrant animals from nearby endangered Arctic stocks? Whatever trig- gered these immature bowhead whales to venture well outside the Arctic Circle remains unknown, yet these observations suggest an unexpected adaptability to foraging in temperate shallow waters.

Bowhead whales (Balaena mysticetus) have a nearly circumpolar distribution, and occasionally occupy warmer shallow coastal areas during summertime that may facilitate molting. However, relatively little is known about the occurrence of molting and associated behaviors in bowhead whales. We opportunistically observed whales in Cumberland Sound, Nunavut, Canada with skin irregularities consistent with molting during August 2014, and collected a skin sample from a biopsied whale that revealed loose epidermis and sloughing. During August 2016, we flew a small unmanned aerial system (sUAS) over whales to take video and still images to: 1) determine unique individuals; 2) estimate the proportion of the body of unique individuals that exhibited sloughing skin; 3) determine the presence or absence of superficial lines representative of rock-rubbing behavior; and 4) measure body lengths to infer age-class. The still images revealed that all individuals (n = 81 whales) were sloughing skin, and that nearly 40% of them had mottled skin over more than two-thirds of their bodies. The video images captured bowhead whales rubbing on large rocks in shallow, coastal areas-likely to facilitate molting. Molting and rock rubbing appears to be pervasive during late summer for whales in the eastern Canadian Arctic.

We contrast two methods for estimating the trends of bowhead whales (Balaena mysticetus) in West Greenland: (1) double platform visual aerial survey, corrected for missed sightings and the time the whales are available at the surface; and (2) a genetic capture-recapture approach based on a 14-yr-long biopsy sampling program in Disko Bay. The aerial survey covered 39,000 km2 and resulted in 58 sightings, yielding an abundance estimate of 744 whales (CV = 0.34, 95% CI: 357–1,461). The genetic method relied on determining sex, mitochondrial haplotypes and genotypes of nine microsatellite markers. Based on samples from a total of 427 individuals, with 11 recaptures from previous years in 2013, this resulted in an estimate of 1,538 whales (CV = 0.24, 95% CI: 827–2,249). While the aerial survey is considered a snapshot of the local spring aggregation in Disko Bay, the genetic approach estimates the abundance of the source of this aggregation. As the whales in Disko Bay primarily are adult females that do not visit the bay annually, the genetic method would presumably yield higher estimates. The studies indicate that an increase in abundance observed between 1998 and 2006 has leveled off.

Despite being a signatory to the International Convention for the Regulation of Whaling in 1946, the U.S.S.R. conducted a 30-yr campaign of illegal whaling which arguably represents one of the greatest failures of management in the history of the industry. Here, using a variety of sources including published literature, formerly secret Soviet industry reports, and interviews with former biologists and whalers, we provide an overview of the history, scope, and economic origins of Soviet whaling and examine the domestic and international political context in which it was set. At various times from 1933 into the 1970’s, the U.S.S.R. operated a total of seven whaling factory fl eets and several shore whaling stations. We estimate that 534,119 whales were killed, of which 178,726 were not reported to the International Whaling Commission (IWC). In the Southern Hemisphere, the greatest impact of these catches was on humpback whales, Megaptera novaeangliae, where (mostly illegal) takes of more than 48,000 whales precipitated a population crash and closure of shore whaling stations in Australia and New Zealand. The Southern Hemisphere also saw large illegal catches of southern right whales, Eubalaena australis. In the North Pacifi c, the greatest impacts were on sperm whales, Physeter macrocephalus (where data on sex and length were routinely misreported together with falsifi ed total catches), as well as on the two already-small populations of right whales, Eubalaena japonica, across the North Pacifi c, and bowhead whales, Balaena mysticetus, in the Okhotsk Sea. Soviet whaling was driven by the state industrial planning system, which frequently set high production targets without regard to the ability of the resource to sustain the Introduction In this article, we trace the history and details of what might be called one of the 20th century’s more notorious environmental crimes: the global campaign of illegal whaling conducted by the U.S.S.R. between 1948 and 19721, a campaign that, together with the poorly managed “legal” whaling of other nations, devastated many whale populations. It is a story of the sprawling Soviet planning system’s obsession with attaining production goals despite limited and diminishing resources, and of the U.S.S.R.’s desire to do every1In 1972 an International Observer Scheme was introduced and large-scale Soviet unrestricted catches ceased, but some falsifi cations of catch data continued on at least some Soviet fl eets for a few more years (Mikhalev et al., 2009). thing bigger and better than other nations, especially those in the capitalist world. More than anything, this is a cautionary tale of the failure of other whaling nations and the International Whaling Commission (IWC) to react to mounting evidence of declining whale stocks, and to adequately monitor adherence to regulations and catch limits2 as set through international

Radiocarbon-dated marine mammal remains from emergent Arctic coastlines have frequently been used to reconstruct Holocene sea-ice histories. The use of such reconstructions has hitherto been complicated by uncertain marine reservoir corrections precluding meaningful intercomparisons with data reported in calibrated or sidereal years. Based on an exhaustive compilation of previously published marine mammal radiocarbon dates (both live-harvested materials and subfossils) from the Canadian Arctic Archipelago (CAA), new, statistically-derived δ13C and ΔR values are provided. Average δ13C values are: −16.1 ± 1.1‰ (bone collagen; n = 193) for bowhead (Balaena mysticetus); −14.4 ± 0.5‰ (n = 44; dentine) for beluga (Delphinapterus leucas); −14.8 ± 1.9‰ (teeth and tusks; n = 18) and −18.0 ± 4.7‰ (n = 9; bone collagen) for walrus (Odobenus rosmarus). ΔR values are 170 ± 95 14C years for bowhead (n = 23) and 240 ± 60 14C years for beluga (n = 12). Scarce data preclude calculation of meaningful, statistically robust walrus ΔR. Using the new ΔR values, an expanded and revised database of calibrated bowhead dates (651 dates; many used in previous CAA sea-ice reconstructions) shows pronounced late Quaternary spatio-temporal fluctuations in bone abundance. Though broadly resembling earlier bowhead subfossil frequency data, analysis of the new expanded database suggests early- and mid-Holocene increases in whale abundance to be of longer duration and lower amplitude than previously considered. A more even and persistent spread of infrequent low-abundance remains during “whale free” intervals is also seen. The dominance of three eastern regions (Prince Regent Inlet & Gulf of Boothia; Admiralty Inlet; Berlinguet Inlet/Bernier Bay) in the CAA data, collectively contributing up to 88% of all subfossil remains in the mid-Holocene, is notable. An analysis of calibrated regional sea-level index points suggests that severance of the Admiralty Inlet-Gulf of Boothia marine channel due to isostatically-driven regression may have played a significant role in enhanced whale mortality during this interval. Comparisons between the newly calibrated bowhead data and other regional sea-ice proxy data further highlight spatial and temporal discrepancies, potentially due to regional asynchronicities and variable sensitivities in proxy response to climate and oceanographic forcing. However, the limited number of deglacial–postglacial marine records continues to hamper extensive intercomparisons between marine mammal and other proxy datasets. Nevertheless, an examination of assumptions inherent in linking bowhead subfossil frequencies, population densities, and sea-ice thickness and distribution, shows that such relationships are highly complex. Factors such as broad sea-ice preferences, variable mortality rates and causes, long distance carcass transport, variable coastline and basin/channel geometries, and changing emergence rates all complicate the correlation of whale bone abundance to sea-ice histories.

This study describes the morphology of the laryngeal apparatus in bowhead whales (Balaena mysticetus) with respect to respiration, deglutition, and vocalization. We also examined the intrinsic cricoarytenothyroid muscle (Musculus (M.) diverticuli laryngei) which forms the laryngeal diverticulum, to ascertain its interactions with the laryngeal cartilages during respiration and sound production. Five fetal larynges and four from adult whales were studied using noninvasive imaging, as well as macroscopic and microscopic techniques. The larynx extends from the skull base into the thoracic inlet. The dorsally curved laryngeal stalk, supported by epiglottis and the corniculate processes of arytenoid cartilages, is situated within the nasopharynx. The epiglottic cartilage exhibits a prominent medial ridge. The arytenoid cartilages are rod-shaped, and extend through the laryngeal cavity. The thyroid cartilage possesses a prominent caudal horn with a fibrous articulation to the ventrally incomplete cricoid cartilage. The M. thyroepiglotticus forms the connection between epiglottic and thyroid cartilages. The M. cricothyroideus lateralis connects the caudal horn of the thyroid cartilage with the cricoid cartilage and the M. cricothyroideus medialis connects the cricoid and thyroid cartilage. An extensive laryngeal diverticulum (Diverticulum laryngis), formed by the laryngeal mucosa and M. diverticuli laryngei, is positioned caudo-ventral to the laryngeal vestibule. The mucosa thickens into a fold medial to the vocal processes of the arytenoid cartilages. Experiments with airflow combined with histological and anatomical evidence strongly suggest a sound producing function for these (vocal) folds. This analysis provides the first account of sound producing structures and function in bowhead whales.

In this paper we studied three related aspects of the ontogeny of the vertebral centrum of cetaceans and terrestrial mammals in an evolutionary context. We determined patterns of ontogenetic fusion of the vertebral epiphyses in bowhead whale (Balaena mysticetus) and beluga whale (Delphinapterus leucas), comparing those to terrestrial mammals and Eocene cetaceans. We found that epiphyseal fusion is initiated in the neck and the sacral region of terrestrial mammals, while in recent aquatic mammals epiphyseal fusion is initiated in the neck and caudal regions, suggesting locomotor pattern and environment affect fusion pattern. We also studied bony fusion of the sacrum and evaluated criteria used to homologize cetacean vertebrae with the fused sacrum of terrestrial mammals. We found that the initial ossification of the vertebral pedicles in the fetus may be a reliable indicator of sacral homology in modern cetaceans. Finally, we also studied fusion of the centra of cervical vertebrae in B. mysticetus and found that it is not completed until after sexual maturity, and after 20 years of age.

The Arctic is one of the fastest-changing parts of the planet. Global climate change is already having major impacts on Arctic ecosystems. Increasing temperatures and reductions in sea ice are particular conservation concerns for ice-associated species, including three endemic cetaceans that have evolved in or joined the Arctic sympagic community over the last 5 M years. Sea ice losses are also a major stimulant to increased industrial interest in the Arctic in previously ice-covered areas. The impacts of climate change are expected to continue and will likely intensify in coming decades. This paper summarizes information on the distribution and movement patterns of the three ice-associated cetacean species that reside year-round in the Arctic, the narwhal ( Monodon monoceros ), beluga (white whale, Delphinapterus leucas ), and bowhead whale ( Balaena mysticetus ). It maps their current distribution and identifies areas of seasonal aggregation, particularly focussing on high-density occurrences during the summer. Sites of oil and gas exploration and development and routes used for commercial shipping in the Arctic are compared with the distribution patterns of the whales, with the aim of highlighting areas of special concern for conservation. Measures that should be considered to mitigate the impacts of human activities on these Arctic whales and the aboriginal people who depend on them for subsistence include: careful planning of ship traffic lanes (re-routing if necessary) and ship speed restrictions; temporal or spatial closures of specified areas (e.g. where critical processes for whales such as calving, calf rearing, resting, or intense feeding take place) to specific types of industrial activity; strict regulation of seismic surveys and other sources of loud underwater noise; and close and sustained monitoring of whale populations in order to track their responses to environmental disturbance.

Passive acoustic monitoring has become a standard method for detecting bowhead whale (Balaena mysticetus) activity in Arctic waters. Between 2007 and 2014, over 40 autonomous vector sensors, known as DASARs, were deployed in the Beaufort Sea during the bowhead whale migration season. Individual DASARs can estimate azimuth, allowing calls to be localized by triangulation using multiple DASARs. However, these bearings are subject to calibration biases, and individual sensors were not precisely time-synchronized, making relative time-of-arrival information unreliable for standard localization purposes. Double-difference methods have previously been applied in seismology to obtain high-precision relative positions of earthquakes by measuring changes in relative travel-times between multiple events over widely distributed seismic sensors. This same concept has also been used to track fin whales on a seafloor seismic network. Here, the double-difference method is applied to previously localized bowhead whale calls in order to improve their relative positions. The approach uses changes in both relative call travel-times and bearings, detected at multiple DASARs, to determine high-precision relative locations of these calls despite the presence of systemic timing and bearing errors in the measurements. The resulting positions may allow tracking of individual whales, which would provide insight into the function of these calls.

Remains of bowhead whales (Balaena mysticetus) harvested by Inupiat whalers are deposited in bone piles along the coast of Alaska and have become persistent and reliable food sources for polar bears (Ursus maritimus). The importance of bone piles to individuals and the population, the patterns of use, and the number, sex, and age of bears using these resources are poorly understood. We implemented barbed-wire hair snaring to obtain genetic identities from bears using the Point Barrow bone pile in winter 2010–11. Eighty-three percent of genotyped samples produced individual and sex identification. We identified 97 bears from 200 samples. Using genetic mark–recapture techniques, we estimated that 228 bears used the bone pile during November to February, which would represent approximately 15% of the Southern Beaufort Sea polar bear subpopulation, if all bears were from this subpopulation. We found that polar bears of all age and sex classes simultaneously used the bone pile. More males than females...

Autonomous passive acoustic recorders were deployed to record sounds of bowhead whales (Balaena mysticetus) in the southeast Beaufort Sea for periods of 30-55 days during the late summer, open-water seasons of 2008-2010. Recordings were made in three areas licensed for hydrocarbon exploration, spanning the continental slope and adjacent outer shelf, and in a shallow inner-shelf area where bowheads have been observed congregating to feed in recent decades. Bowhead sounds were counted in samples comprising 10% of each recorded hour. In mid-August and September in all 3 years, the rate of bowhead calling at outer shelf sites exceeded that at adjacent continental slope sites by one to two orders of magnitude. Higher rates of calling occurred on the slope in late July and early August than at later dates. Calling rates varied by an order of magnitude between years in the one area that was monitored in different years. The highest rates of calling occurred on the inner shelf, offshore of the northern Tuktoyaktuk Peninsula. These trends are consistent with patterns of habitat use previously reported from aerial surveys in this and nearby areas of the Beaufort Sea and with the results of satellite tagging studies.

Bowhead whale Balaena mysticetus progesterone concentrations were measured in different sample matrices (serum, blubber, and urine) to investigate (1) concordance among sam- ple type and (2) variation among life-history class. Samples were collected from subsistence- hunted whales (n = 86) taken from 1999 to 2009. In general, irrespective of sample matrix, preg- nant females had the highest concentrations by orders of magnitude, followed by mature animals of both sexes, and subadults had the lowest concentrations. Subadult males and females had sim- ilar progesterone concentrations in all sample matrices measured. When pregnant animals were included in our analyses, permuted regression models indicated a strong positive relationship between serum and blubber progesterone levels (r 2 = 0.894, p = 0.0002). When pregnant animals were not included, we found no significant relationship between serum and blubber levels (r 2 = 0.025, p = 0.224). These results suggest that progesterone concentrations are mirrored in these sample types over longer periods (i.e. on the order of weeks to months, time frame of repro- ductive changes) but not shorter periods (i.e. on the order of hours to days, time frame of daily fluc- tuations). This conclusion is consistent even for progesterone concentrations measured in females that had recently changed pregnancy states (either new mothers or newly pregnant animals), for which blubber progesterone levels seem to lag those in the serum. Finally, urine progesterone had statistically significant positive relationships with serum (r 2 = 0.136, p = 0.0460) and blubber pro - gesterone (r 2 = 0.150, p = 0.0421). Our results suggest that progesterone concentrations first peak in the serum, then in the urine, and finally in the blubber.

Several cetacean and pinniped species use the northeastern Chukchi Sea as seasonal or year-round habitat. This area has experienced pronounced reduction in the extent of summer sea ice over the last decade, as well as increased anthropogenic activity, particularly in the form of oil and gas exploration. The effects of these changes on marine mammal species are presently unknown. Autonomous passive acoustic recorders were deployed over a wide area of the northeastern Chukchi Sea off the coast of Alaska from Cape Lisburne to Barrow, at distances from 8 km to 200 km from shore: up to 44 each summer and up to 8 each winter. Acoustic data were acquired at 16 kHz continuously during summer and on a duty cycle of 40 or 48 min within each 4-h period during winter. Recordings were analyzed manually and using automated detection and classification systems to identify calls. Bowhead ( Balaena mysticetus ) and beluga ( Delphinapterus leucas ) whale calls were detected primarily from April through June and from September to December during their migrations between the Bering and Beaufort seas. Summer detections were rare and usually concentrated off Wainwright and Barrow, Alaska. Gray ( Eschrichtius robustus ) whale calls were detected between July and October, their occurrence decreasing with increasing distance from shore. Fin ( Balaenoptera physalus ), killer ( Orcinus orca ), minke ( Balaenoptera acutorostrata ), and humpback ( Megaptera novaeangliae ) whales were detected sporadically in summer and early fall. Walrus ( Odobenus rosmarus ) was the most commonly detected species between June and October, primarily occupying the southern edge of Hanna Shoal and haul-outs near coastal recording stations off Wainwright and Point Lay. Ringed ( Pusa hispida ) and bearded ( Erignathus barbatus ) seals occur year-round in the Chukchi Sea. Ringed seal acoustic detections occurred throughout the year but detection numbers were low, likely due to low vocalization rates. Bearded seal acoustic detections peaked in April and May during their breeding season, with much lower detection numbers in July and August, likely as a result of reduced calling rates after breeding season. Ribbon seals ( Histriophoca fasciata ) were only detected in the fall as they migrated south through the study area toward the Bering Sea. These results suggest a regular presence of marine mammals in the Chukchi Sea year-round, with species-dependent seasonal and spatial density variations.

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.

The surfacing, respiration and diving (SRD) behavior of bowhead whales Balaena mysticetus changes upon exposure to seismic operations. However, it is unknown whether these changes differ by season, reproductive status (calves, mothers, and non-calves), and whale activity (traveling, foraging, or socializing). Such SRD behavioral responses to seismic operations might influence the detectability of whales during aerial surveys. We addressed these questions by applying non-parametric univariate tests and linear mixed models to behavioral data collected by aerial observation of bowheads in the Beaufort Sea from 1980 to 2000. Durations of surfacings decreased upon exposure to seismic operations, especially for traveling or socializing non-calf whales. The mixed models also indicated that dive durations were affected by the presence of seismic operations, but the effects depended on other variables such as season and whale activity. Overall, our results suggest that changes in the behavior exhibited by bowhead whales exposed to seismic operations are context-dependent (i.e. responses to seismic operations depend on both the circumstance and activity of the whale). The level of perceived threat may also be important based on similarities with behavioral changes observed in other air-breathing aquatic foragers facing dangers. We conclude that seismic-induced changes in bowhead SRD behaviors may affect the availability of bowhead whales for visual detection in some circumstances. This in turn means that estimates of abundance and distribution of bowhead whales near seismic surveys should be context-sensitive and incorporate correction factors that account for sound exposure, season, reproductive status, and whale activity.

Bowhead whales (Balaena mysticetus) of the western Arctic stock winter in icecovered continental shelf regions of the Bering Sea, where pot fisheries for crabs (Paralithodes and Chionoecetes spp.) and Pacific cod (Gadus macrocephalus) pose a risk of entanglement. In the winter of 2008–2009 and 2009–2010 the spatial distribution of 21 satellite tagged bowhead whales partially overlapped areas in which pot fisheries for cod and blue king crab (Paralithodes platypus) occurred. However, these fisheries ended before whales entered the fishing areas, thus avoiding temporal overlap. A fishery for snow crab (Chionoecetes opilio) typically runs from January to May and provides the greatest potential for bowhead whales to encounter active pot gear. Tagged whales did not enter the area of the snow crab fishery during this study and generally remained in areas with >90% sea ice concentration, which is too concentrated for crab boats to penetrate. Pack ice sometimes overruns active fishing areas, resulting in lost gear, which is the most likely source of entanglement. The western Arctic stock of bowhead whales was increasing as of 2004; as such, incidental mortality from commercial pot fisheries is probably negligible at this time. Regardless, entanglement may increase over time and should be monitored.

Zooarchaeologists continue to experience difficulty defining the importance of bowhead (Balaena mysticetus) whaling in Neoeskimo coastal deposits. The large size of bowhead bones, combined with their use as structural elements in Neoeskimo architecture, creates a suite of taphonomic issues that tend to obscure their usefulness as a measure of relative abundance, and thus as an overall economic indicator. Here we present a regional approach that focuses on contrasts in relative taxonomic abundance between sites with diverse economic signatures, supported by related differences in element frequencies, site locations, features, artefact frequencies, and manufacturing detritus. Using this approach, a generalized picture of the relative importance of bowhead whales in Neoeskimo subsistence economies can be assembled. Such an analysis, applied to the archaeological record of the Mackenzie Inuit, or Siglit, reveals the role that bowhead whaling played in subsistence economies from the 15 th to 19 th centuries AD. Specifically, the archaeological record indicates that the prehistoric Qikiqtaryungmiut and Nuvugarmiut practiced specialized bowhead whaling at coastal promontories, though the seasonal scheduling and success rate of these hunts contrasted considerably.

Killer whales (Orcinus orca) are increasing in occurrence and residence time in the eastern Canadian Arctic (ECA) in part due to a decrease in sea ice associated with global climate change. Killer whales prey on bowhead whales (Balaena mysticetus) of the Eastern Canada-West Greenland (EC-WG) population, but their patterns of predation pressure and effect on the EC-WG population’s ability to recover from historical whaling remain unknown. We analyzed photographs of individual bowhead whale flukes from five regions within the EC-WG population’s geographic range (Cumberland Sound, Foxe Basin, Isabella Bay, Repulse Bay and Disko Bay), taken during 1986 and from 2007 to 2012, to estimate the occurrence of rake marks (parallel scars caused by killer whale teeth). Of 598 identified whales, 10.2 % bore rake marks from killer whales. A higher occurrence of rake marks was found in Repulse and Disko Bays, where primarily adult bowhead whales occur seasonally, than in Foxe Basin, where juveniles and females with calves occur. Older bowheads, which have had greater exposure time to killer whales due to their age, had higher occurrences of rake marks than juveniles and calves, which may indicate that younger whales do not survive killer whale attacks. A high proportion of adult females also had rake marks, perhaps due to protecting their calves from killer whale predation. In order to quantify the effect of killer whales on EC-WG population recovery, further research is needed on the relationship between the occurrence of rake marks and bowhead adult, calf, and juvenile mortality in the ECA, as well as more information about Arctic killer whale ecology.

The novel observation of a palatal retial organ in the bowhead whale (Balaena mysticetus) is reported, with characterization of its form and function. This bulbous ridge of highly vascularized tissue, here designated the corpus cavernosum maxillaris, runs along the center of the hard palate, expanding cranially to form two large lobes that terminate under the tip of the rostral palate, with another enlarged node at the caudal terminus. Gross anatomical and microscopic observation of tissue sections discloses a web-like internal mass with a large blood volume. Histological examination reveals large numbers of blood vessels and vascular as well as extravascular spaces resembling a blood-filled, erectile sponge. These spaces, as well as accompanying blood vessels, extend to the base of the epithelium. We contend that this organ provides a thermoregulatory adaptation by which bowhead whales (1) control heat loss by transferring internal, metabolically generated body heat to cold seawater and (2) protect the brain from hyperthermia. We postulate that this organ may play additional roles in baleen growth and in detecting prey, and that its ability to dissipate heat might maintain proper operating temperature for palatal mechanoreceptors or chemoreceptors to detect the presence and density of intraoral prey. Anat Rec, 296:701–708, 2013. © 2013 Wiley Periodicals, Inc.

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 w...

Patterns of genetic variation observed within species reflect evolutionary histories that include signatures of past demography. Understanding the demographic component of species' history is fundamental to informed management because changes in effective population size affect response to environmental change and evolvability, the strength of genetic drift, and maintenance of genetic variability. Species experiencing anthropogenic population reductions provide valuable case studies for understanding the genetic response to demographic change because historic changes in the census size are often well documented. A classic example is the bowhead whale, Balaena mysticetus, which experienced dramatic population depletion due to commercial whaling in the late 19th and early 20th centuries. Consequently, we analyzed a large multi-marker dataset of bowhead whales using a variety of analytical methods, including extended Bayesian skyline analysis and approximate Bayesian computation, to characterize genetic signatures of both ancient and contemporary demographic histories. No genetic signature of recent population depletion was recovered through any analysis incorporating realistic mutation assumptions, probably due to the combined influences of long generation time, short bottleneck duration, and the magnitude of population depletion. In contrast, a robust signal of population expansion was detected around 70,000 years ago, followed by a population decline around 15,000 years ago. The timing of these events coincides to a historic glacial period and the onset of warming at the end of the last glacial maximum, respectively. By implication, climate driven long-term variation in Arctic Ocean productivity, rather than recent anthropogenic disturbance, appears to have been the primary driver of historic bowhead whale demography.

The eastern Canada-West Greenland (EC-WG) bowhead whale Balaena mysticetus population is slowly recovering from the intensive commercial whaling of the 18th and 20th cen- turies. However, climate change, through effects on ice conditions and prey availability, is one of several threats that might affect bowhead whale recovery. In this study, we exploited the variabil- ity observed in isotopic signatures of prey assemblages across the eastern Arctic to examine vari- ability in diet among bowhead whales (n = 202) and identify their potential foraging areas. We compared δ 13 C and δ 15 N isotope ratios of biopsied skin samples with those of potential zooplank- ton prey species collected across the Canadian eastern Arctic, and calculated the proportional contributions of various sources (zooplankton) to the diet of bowhead whales using a Bayesian sta- ble isotope mixing model. A cluster analysis indicated some variability in isotopic composition among groups of individuals, but not between males and females or age classes. The isotopic model discounted Davis Strait and Disko Bay as potential foraging areas for bowhead whales, at least in spring and summer. Lancaster Sound, Baffin Bay and the Gulf of Boothia were the 3 main areas likely used for summer feeding, where bowhead whales fed primarily on large Arctic calanoid copepods (Calanus hyperboreus, C. glacialis, Metridia longa, and Paraeuchaeta spp.), mysids and euphausiids. While some inter-individual variability in diet was observed, the strong dependence of this endemic Arctic species on Arctic zooplankton may make them vulnerable to the predicted latitudinal shift in prey species composition caused by ongoing warming.

William Spiers Bruce was a medical student who sailed as ship's surgeon on the Balaena, one of the ships of the Dundee Whaling Expedition to the Antarctic (1892–93). On this expedition he kept a diary listing his medical consultations and the treatments he administered. This paper considers the expedition and the diaries.

Bowhead whales (Balaena mysticetus) of the Bering-Chukchi-Beaufort population form annual late-summer feeding aggregations offshore of the eastern Tuktoyaktuk Peninsula and Cape Bathurst. This region is strongly influenced by episodic upwelling events. A systematic aerial survey (10% coverage) was flown over the eastern shelf on 2 August 2008, revealing large numbers of bowheads (est. 3500 individuals) in a clumped distribution over the continental shelf. Between 7 and 11 August 2008, zooplankton samples and hydrography were obtained from the vicinity of two bowhead feeding aggregations, one at the northeast edge of the continental shelf, and the second off the northeast shore of the Cape Bathurst. Dense aggregations of zooplankton were found in the lower part of the water column (below 40 m), on the shelf, in water with an upwelling signature. The zooplankton were largely resting phases of calanoid copepods (Calanus hyperboreus and C. glacialis) and collectively had twice the abundance, twice the biomass, three times the caloric density and thus six times the energy content of contemporaneous zooplankton samples from the western Canadian Beaufort Shelf. Although upwelling at Cape Bathurst is wind-driven and thus episodic in nature, its consistent delivery of zooplankton to the bowhead foraging area creates conditions attractive to bowhead whales, with an estimated 33% of the bowhead population present on the eastern portion of the shelf at the time of the August 2008 aerial survey.

Sea ice is believed to be a major factor shaping gene flow for polar marine organisms, but it remains unclear to what extent it represents a true barrier to dispersal for arctic cetaceans. Bowhead whales are highly adapted to polar sea ice and were targeted by commercial whalers throughout Arctic and subarctic seas for at least four centuries, resulting in severe reductions in most areas. Both changing ice conditions and reductions due to whaling may have affected geographic distribution and genetic diversity throughout their range, but little is known about range-wide genetic structure or whether it differed in the past. This study represents the first examination of genetic diversity and differentiation across all five putative stocks, including Baffin Bay-Davis Strait, Hudson Bay-Foxe Basin, Bering-Beaufort-Chukchi, Okhotsk, and Spitsbergen. We also utilized ancient specimens from Prince Regent Inlet (PRI) in the Canadian Arctic and compared them with modern stocks. Results from analysis of molecular variance and demographic simulations are consistent with recent and high gene flow between Atlantic and Pacific stocks in the recent past. Significant genetic differences between ancient and modern populations suggest PRI harbored unique maternal lineages in the past that have been recently lost, possibly due to loss of habitat during the Little Ice Age and/or whaling. Unexpectedly, samples from this location show a closer genetic relationship with modern Pacific stocks than Atlantic, supporting high gene flow between the central Canadian Arctic and Beaufort Sea over the past millennium despite extremely heavy ice cover over much of this period.

The processes of lipid deposition and utilization, via the gene leptin (Lep), are poorly understood in taxa with varying degrees of adipose storage. This study examines how these systems may have adapted in marine aquatic environments inhabited by cetaceans. Bowhead (Balaena mysticetus) and beluga whales (Delphinapterus leucas) are ideal study animals—they possess large subcutaneous adipose stores (blubber) and undergo bi-annual migrations concurrent with variations in food availability. To answer long-standing questions regarding how (or if) energy and lipid utilization adapted to aquatic stressors, we quantified variations in gene transcripts critical to lipid metabolism related to season, age, and blubber depth. We predicted leptin tertiary structure conservation and assessed inter-specific variations in Lep transcript numbers between bowheads and other mammals. Our study is the first to identify seasonal and age-related variations in Lep and lipolysis in these cetaceans. While Lep transcripts and protein oscillate with season in adult bowheads reminiscent of hibernating mammals, transcript levels reach up to 10 times higher in bowheads than any other mammal. Data from immature bowheads are consistent with the hypothesis that short baleen inhibits efficient feeding. Lipolysis transcripts also indicate young Fall bowheads and those sampled during Spring months limit energy utilization. These novel data from rarely examined species expand the existing knowledge and offer unique insight into how the regulation of Lep and lipolysis has adapted to permit seasonal deposition and maintain vital blubber stores.

There are no standards for assessment of the cumulative effects of underwater sound. Quantitative assessments typically consider a single source, whereas qualitative assessments may include multiple sources but rarely identify response variables. As a step toward understanding the cumulative effects of underwater sound, we assessed the aggregated sounds of multiple sources received by migrating bowhead whales (Balaena mysticetus). The quantitative method models the sound field from multiple sources and simulates movement of a population through it. The qualitative method uses experts to assess the responses of individuals and populations to sound sources and identify the potential mechanisms. These methods increase the transparency of assessments.

Management strategy evaluation (MSE) involves using simulation to compare the relative effectiveness for achieving management objectives of different combinations of data collection schemes, methods of analysis and subsequent processes leading to management actions. MSE can be used to identify a ‘best’ management strategy among a set of candidate strategies, or to determine how well an existing strategy performs. The ability of MSE to facilitate fisheries management achieving its aims depends on how well uncertainty is represented, and how effectively the results of simulations are summarized and presented to the decision-makers. Key challenges for effective use of MSE therefore include characterizing objectives and uncertainty, assigning plausibility ranks to the trials considered, and working with decision-makers to interpret and implement the results of the MSE. This paper explores how MSEs are conducted and characterizes current ‘best practice’ guidelines, while also indicating whether and how these best practices were applied to two case-studies: the Bering–Chukchi–Beaufort Seas bowhead whales (Balaena mysticetus; Balaenidae) and the northern subpopulation of Pacific sardine (Sardinops sagax caerulea; Clupeidae).

Fisheries sonar was used to determine the applicability of active acoustic monitoring (AAM) for marine mammal detection in the Canadian Beaufort Sea. During 170 h of simultaneous observation by marine mammal observers and active acoustic observation, 119 Balaena mysticetus (bowheads) and 4 Delphinapterus leucas (belugas) were visually sighted, while 59 acoustic signals of bowheads were detected by AAM operators. Observations and detection of seals were also recorded. Comparative results indicate that commercially available active acoustic systems can detect seals at distances up to 500 m and large baleen whales at distances up to 2 km.

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 (kAsp) 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 kAsp = 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 kAsp and (D/L)0. One male exceeded 100 yr of age; the oldest female was 88. A strong linear relationship between kAsp and body temperature was estimated by combining bowhead data with independent data from studies of humans and fin whales. Using this relationship, we estimated kAsp and ASM for North Atlantic minke whales.

Both bowhead Balaena mysticetus and North Pacific (NP) right whales Eubalaena japonica were reduced to low levels by historical whaling. Despite their protected status, it is known that the USSR illegally killed both species in the NP and Okhotsk Sea (OS). Here, we pro- vide revised Soviet catch totals, as well as other new information on the distribution and other details of these catches. Right whale catches were made from 1962 to 1968 in the eastern NP and in 1967 and 1968 in the OS. Our best estimate of total right whale catches is 661, consisting of 529 for the eastern NP (compared to the previously published figure of 372) and 152 for the OS (cf. a previous figure of 136). Catches were distributed in the Bering Sea (BS, 115), eastern Aleutian Islands (28), Gulf of Alaska (GOA, 366), OS (132), and other areas (20). Detailed information on catches of 112 right whales taken in May/June 1963 shows a broad distribution in offshore waters of the GOA, consistent with 19th century historical whaling records. Other major areas in which right whales were caught include south of Kodiak Island, western Bristol Bay (southeastern BS), and the central OS off eastern Sakhalin Island. The best estimate of bowhead whale catches in the OS in 1967 and 1968 is 145 animals, although this is contingent upon certain assumptions regard- ing species identity. Of these, 79 were killed in the Shantar Islands region and 66 in Shelikhov Bay. The catches of both species primarily involved large mature animals, thus greatly inhibiting recovery of the populations concerned.

We used baleen lengths and age estimates from 175 whales and body lengths and age estimates from 205 whales to test which of several single- and multi-stage growth models best characterized age-specific baleen and body lengths for bowhead whales ( Balaena mysticetus L., 1758) with the goal of determining which would be best for predicting whale age based on baleen or body length. Previous age estimates were compiled from several techniques, each of which is valid over a relatively limited set of physical characteristics. The best fitting single-stage growth model was a variation of the von Bertalanffy growth model for both baleen and body length data. Based on Bayesian information criterion, the two- and three-stage versions of the von Bertalanffy model fit the data better than did the single-stage models for both baleen and body length. The best baleen length models can be used to estimate expected ages for bowhead whales with up to 300–325 cm baleen, depending on sex, which correspond to age estimates approaching 60 years. The best body length models can be used to estimate expected ages for male bowhead whales up to 14 m, and female bowheads up to 15.5 m or ages up to approximately 40 years.

Bowhead whales Balaena mysticetus are long-lived cetaceans, uniquely adapted among the baleen whales to live year-round in the Arctic. All bowhead whale populations were greatly reduced by commercial whaling from the 1600s through the 1800s, with the largest, the Spitsber- gen population in the North Atlantic, depleted to the point of extinction. Recent sightings of bow- head whales west of Svalbard precipitated an effort to listen for their vocalizations via 2 re - corders deployed in 2008 on oceanographic moor- ings spaced 95 km apart at 78.8° N latitude in the Fram Strait. Year-round acoustic records were examined for the occurrence of bowhead whale sounds. Simple calls, call sequences, and complex songs were recorded. Repeated call sequences or bowhead whale songs were detected nearly every hour from early November 2008 through late April 2009 on the western Fram Strait recorder. More than 60 unique songs were recorded from October 2008 to April 2009. In contrast, simple calls and call sequences were the most common signals recorded on the central Fram Strait instrument. Peak levels of song production coincided with the period of lowest water temperature, dense ice con- centration, and almost complete darkness. Given the diversity, loudness, and period over which songs were recorded, western Fram Strait appears to be a wintering ground — and potentially a mat- ing area — for this Critically Endangered popula- tion of bowhead whales.

Laidre, K. L., and Heide-Jørgensen, M. P. 2012. Spring partitioning of Disko Bay, West Greenland, by Arctic and Subarctic baleen whales. – ICES Journal of Marine Science, 69: . Movements of co-occurring bowhead (Balaena mysticetus) and humpback (Megaptera novaeangliae) whales in Disko Bay, West Greenland, were examined using satellite telemetry. Data on movements, habitat use, and phenology were collected from tagged 49 bowheads and 44 humpbacks during the transition from sea-ice breakup to open water between 2008 and 2010. Bowhead whales began their northward spring migration around 27 May (median day-of-the-year departure date = 147, interquartile range 141–153) and were distributed broadly in northern and central Disko Bay in water depths between 100 and 400 m. Humpback whales arrived in Disko Bay no later than 2 June and were located in shallow water (

Bowhead whale Balaena mysticetus critical habitat was identified as a key informa- tion gap by the Eastern Arctic Bowhead Whale Recovery Team. To fill this gap, data on eastern Canadian Arctic (ECA) bowhead whales and their habitat were collected and analyzed. We selected governmental, private, and historical whaling bowhead location datasets which differed in temporal and spatial extent, sample size, and quality. Sufficient data were available only for the 'reduced-ice' period (June to October) and pooled by month. Data for 6 ecogeographical variables (EGVs) were integrated into a geographical information system (GIS): sea surface temperature, chlorophyll, ice, depth, slope, and distance to shore. A monthly ecological niche factor analysis was performed for each whale and EGV dataset to determine habitat suitability in the ECA. Eleven habitat suitability models were produced, and a composite map of predicted high suitabil- ity habitat, for all 5 months, was developed. Twenty-one areas within the ECA were identified as highly suitable habitat and ranked according to analytical confidence. Six critical habitats were identified and are supported by recent scientific evidence and Inuit knowledge. Recently, the pop- ulation estimate, conservation status, and management of the Eastern CanadaWest Greenland bowhead population have changed dramatically (bowhead whales of this population also inhabit the ECA). In parallel, evidence of ecological change from climate warming has increased and associated loss of sea ice is anticipated to increase interactions between bowheads and anthro- pogenic activity. As envisioned by the recovery team, this study provides resource managers with a timely tool for population recovery, conservation, and protection.

Mitochondrial DNA (mtDNA) sequences were analyzed from 106 bowhead whale (Balaena mysticetus) specimens dating 471 ± 44 14 C B.P.–10,290 ± 150 14 C 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 (FST ≤ 0.005, � 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 14 C 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.

The objective of this paper was to investigate and illustrate how insights gained from experience managing human activities in order to protect North Atlantic right whales (Eubalaena glacialis) along the heavily industrialized east coast of North America might be applied in the Arctic, where bowhead whales (Balaena mysticetus) face some of the same risks as right whales. The reduced extent and thickness of sea ice and the resultant longer open-water season have major, complex implications for the Arctic marine ecosystem. Increased maritime ship traffic and commercial fishing in the Arctic are bound to affect bowheads and Native (indigenous) hunting communities who depend on whales for subsistence and cultural identity. Bowheads and right whales were greatly depleted by commercial whaling in the 19th and early 20th centuries. While the Western Arctic bowhead population has been recovering steadily in recent decades, North Atlantic right whales remain highly endangered because of persistent lethal and sublethal vessel strikes and frequent entanglement in commercial fishing gear. Entanglement can be transitory or persistent, with debilitation lasting for months before the animal finally succumbs. Vessel strike and fishing gear trauma has been documented in bowheads, but at a much lower rate than in right whales. Initiatives intended to mitigate the impacts of ship traffic on North Atlantic right whales have included speed limits and routing changes. Those meant to reduce the incidence and severity of entanglements include the modification of gear design and gear deployment practices. Management measures need to be considered in advance in the Arctic in order to minimize the risks to bowhead whales as shipping and industrial fishing expand in the Arctic with ice retreat.

The bowhead whale Balaena mysticetus is the only mysticete species endemic to the Arctic. The Eastern Arctic-West Greenland (EA-WG) population is considered of special concern in Canada as these whales remain at risk of becoming threatened or endangered due to their slow rate of growth and low fecundity, and ongoing environmental changes in the Arctic. In this context, we used satellite-linked time-depth recorders (SDR-T16) to investigate their movements and dive behaviour and identify plausible summer feeding areas in the Canadian Arctic. Seven individuals from the northern Foxe Basin (FB) (n = 4 in July 2003) and Cumberland Sound (CS) (n = 3 in July 2006) were tracked by satellite for 17 to 293 d. Movement patterns from 4 whales were analyzed using a hidden Markov model (HMM) to inform the probability of whales being in one of 2 behavioural modes: transient or resident. Comparing dive characteristics during transient and resident periods, we observed that the Gulf of Boothia (GB) with moderate ice coverage (54−62%) was used as a summer foraging area by all 4 whales even though they came from different regions. All animals transited rapidly through Fury and Hecla Strait, an area of heavy ice coverage (80−98%). Whales spent most of their time at shallow depths (8−16 m) regardless of time of day when in resident mode, likely feeding on near-surface aggregations of zooplankton. Considering the apparent importance of the GB as a feeding area for this population, every effort should be made to maintain the integrity of this ecosystem.

Simultaneous production of two harmonically independent sounds, the two-voice phenomenon, is a well-known feature in bird song. Some toothed whales can click and whistle simultaneously, and a few studies have also reported simultaneous sound production by baleen whales. The mechanism for sound production in toothed whales has been largely uncovered within the last three decades, whereas mechanism for sound production in baleen whales remains poorly understood. This study provides three lines of evidence from recordings made in 2008 and 2009 in Disko Bay, Western Greenland, strongly indicating that bowhead whales are capable of simultaneous dual frequency sound production. This capability may function to enable more complex singing in an acoustically mediated reproductive advertisement display, as has been suggested for songbirds, and/or have significance in individual recognition.

Multi-locus genotype and sex were used to identify individual bowhead whales Bal- aena mysticetus from 710 samples collected between 1995 and 2010 at 4 localities in eastern Canada (Foxe Basin, Pelly Bay, Repulse Bay, and Cumberland Sound) and at 1 locality in West Greenland (Disko Bay). In total, 29 recaptures of the same individuals were identified between years, of which 26 individuals were recaptured within the same locality. The remaining 3 were recaptured between sampling localities, from 2 putative International Whaling Commission (IWC) stocks: the Hudson BayFoxe Basin stock and the Baffin BayDavis Strait stock. These recaptures agree with satellite tracking results that demonstrate movement between IWC stocks and ques- tion whether these stocks are true biological entities. The intervals between capture and recap- ture of females in Disko Bay ranged from 1 to 8 yr. The observed number of multi-year recaptures compared to the expected numbers of recaptures did not indicate any clear cyclicity in the use of Disko Bay consistent with the notion that the migration pattern of females to this area might be tied to their reproductive cycles. A markrecapture estimate of whales identified in 2010 com- pared to all identifications between 2000 and 2009 resulted in an estimate of 1410 bowhead whales (SE = 320, 95% CI: 783�2038) constituting the spring aggregation in Disko Bay. The esti- mate for the female portion of the aggregation was 999 individuals (SE = 231, 95% CI: 546�1452). The multi-year cycle of appearance in Disko Bay emphasizes the necessity for identifying whales over multiple years for inclusion in markrecapture estimation.

During the International Polar Year (IPY), acoustic recorders were deployed on oceanographic moorings in Fram Strait and on the Chukchi Plateau, representing the first coordinated year-round sampling of underwater acoustic habitats at two sites in the High Arctic. Examination of species-specific marine mammal calls recorded from autumn 2008–2009 revealed distinctly different acoustic habitats at each site. Overall, the Fram Strait site was acoustically complex compared with the Chukchi Plateau site. In Fram Strait, calls from bowhead whales (Balaena mysticetus) and a variety of toothed whales (odontocetes) were recorded year-round, as were airgun pulses from seismic surveys. In addition, calls from blue whales (Balaenoptera musculus) and fin whales (B. physalus) were recorded from June to October and August to March, respectively. Conversely, at the Chukchi Plateau site, beluga (Delphinapterus leucas) and bowhead whale calls were recorded primarily from May to August, with airgun signals detected only in September–October. Ribbon seal (Phoca fasciata) calls were detected in October–November, with no marine mammals calls at all recorded from December to February. Of note, ice-adapted bearded seals (Erignathus barbatus) were recorded at both sites, primarily in spring and summer, corresponding with the mating season for that species. Differences in acoustic habitats between the two sites were related to contrasts in sea ice cover, temperature, patterns of ocean circulation and contributions from anthropogenic noise sources. These data provide a provisional baseline for the comparison of underwater acoustic habitats between Pacific and Atlantic sectors of the High Arctic.

Aerial photographs were analyzed to investigate the feeding habits of the Bering-Chukchi-Beaufort (BCB) population of bowhead whales (Balaena mysticetus), particularly epibenthic feeding near Barrow, Alaska. Evidence of epibenthic feeding was based on mud visible on the dorsal surface of whales, resulting from feeding near the seafloor. Other cues used to assess feeding were an open mouth or the presence of feces in photographs. Over 3,600 photographs were analyzed including photos from surveys in spring and late summer and in both the western and eastern Beaufort Sea. Of all the photographs analyzed, 64% were scored as definitively muddy. In spring, ratios ranged from a low of 27% in 2003 to a high of 76% in 2004. When all May sample sets off Barrow were combined (1985, 1986, 2003, 2004), there was a significant difference (t-test, P < 0.004) between the proportion of muddy juveniles to the proportion of muddy adults, with muddy adults being more common. The Barrow area was a commonly used feeding ground during migrations in both the spring (61% of the sample were feeding; 55% epibenthically) and autumn (99% of the sample; 97% epibenthically). Bowheads both migrate and feed through areas where petroleum extraction is underway and anticipated; hence, exposure to oil after a spill is of considerable concern to Native communities and management agencies.