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Our limited knowledge of the marine mammal fauna in northernmost Greenland and Canada, specifically north of 80°N, relies largely on opportunistic observations collected during expeditions with different objectives. The narwhal (Monodon monoceros) migrates long distances in response to ice formation and decay and is notoriously skittish, avoiding areas with ice breakers. Scattered observations from the past 20 years, assessed together with historical observations after 1881, suggest that there is a population of narwhals that uses Hall Basin and its adjacent fjord systems—for example, Nares Strait—as a summer ground. Dating the tusks and bones that have been found shows that narwhals were present in this area as far back as nearly 7000 years ago. The wintering locations of these narwhals remain unknown, highlighting the need to investigate whether they are vulnerable to hunting activities in north-west Greenland. By gaining a better understanding of the narwhals’ winter behaviour and potential hunting risks, we can develop more informed conservation and management strategies for this population.

Abstract Predator–prey interactions in marine ecosystems are dynamically influenced by light, as demonstrated by diel vertical migrations of low-trophic level organisms. At high latitudes, the long winter nights can provide foraging opportunities for marine predators targeting vertically migrating prey closer to the surface at night. However, there is limited documentation of such diel patterns in marine predators under extreme light regimes. To address this, we recorded the diving behavior of 17 harbour porpoises just south of the Arctic circle in West Greenland, from summer to winter. Unlike classical diel vertical migration, the porpoises dove 24–37% deeper at night and the frequency of deep dives (> 100 m) increased tenfold as they entered the darkest months. The daily mean depth was negatively correlated with daylength, suggesting an increased diving activity when approaching the polar night. Our findings suggest a light-mediated strategy in which harbour porpoises would either target (i) benthic prey, (ii) pelagic prey migrating seasonally towards the seafloor, or (iii) vertically migrating prey that may be otherwise inaccessible in deeper waters at night, therefore maximizing feeding activity during extended periods of darkness. Extreme light regimes observed at high latitudes are therefore critical in structuring pelagic communities and food webs.

Highly polymorphic single tandem repeat loci (STR, also known as microsatellite loci) remain a familiar, cost efficient class of genetic markers in genetic studies in ecology, behavior and conservation. Here we characterize a new, universal set of ten STR loci in seven species of baleen whales, optimized for PCR amplification in two multiplex reactions along with a Y chromosome marker for sex determination. The optimized, universal set of STR loci provides a convenient starting point for new genetic studies in baleen whales aimed at identifying individuals and populations. Data from the new STR loci were combined with genotypes from previously published STR loci to assess the power to assign parentage using paternity exclusion in four species: fin whale (Balaenoptera physalus), humpback whale (Megaptera novaeangliae), blue whale (B. musculus) and bowhead whale (Balaena mysticetus). Our results suggest that parentage studies should always be accompanied by a power analysis in order to ascertain that each individual specific study is based upon data with sufficient power to assign parentage with statistical rigor.

Inferring animal behavior from irregular tracking data is a challenging area of research. It is particularly difficult to determine if whales, who intermittently explore different depths while staying in the same acoustic medium, synchronize their days with their prey and each other over kilometer-scale distances. Here, we aim to better understand the diving behavior of bowhead whales (Balaena mysticetus) in Disko Bay, West Greenland, using the largest high-frequency dive-depth dataset to date (144 days at 1 Hz from 12 different whales) and nonlinear dynamics, whereby we consider the whales to be chaotic (aperiodic) oscillators. We find that foraging whales dive deeper during the daytime in spring, with this diving behavior being in apparent synchrony with their vertically migrating prey. Furthermore, we demonstrate that bowhead whales can synchronize their behavior with each other for up to a week while staying within a range of up to ∼100km. This discovery agrees with the acoustic herd theory of long-range signaling in baleen whales. On the other hand, the synchrony might emerge when animals experience similar ecological conditions, which are, however, difficult to name because targeted depths and locations were separated for hundreds of meters and tens of kilometers, respectively. In this paper, we identify a framework for studying the sociality and behavior of such chaotically moving, unrestrained marine animals and call for more simultaneous tagging campaigns.

Abstract Background Seasonal long-distance movements are a common feature in many taxa allowing animals to deal with seasonal habitats and life-history demands. Many species use different strategies to prioritize time- or energy-minimization, sometimes employing stop-over behaviours to offset the physiological burden of the directed movement associated with migratory behaviour. Migratory strategies are often limited by life-history and environmental constraints, but can also be modulated by the predictability of resources en route. While theory on population-wide strategies (e.g. energy-minimization) are well studied, there are increasing evidence for individual-level variation in movement patterns indicative of finer scale differences in migration strategies. Methods We aimed to explore sources of individual variation in migration strategies for long-distance migrators using satellite telemetry location data from 41 narwhal spanning a 21-year period. Specifically, we aimed to determine and define the long-distance movement strategies adopted and how environmental variables may modulate these movements. Fine-scale movement behaviours were characterized using move-persistence models, where changes in move-persistence, highlighting autocorrelation in a movement trajectory, were evaluated against potential modulating environmental covariates. Areas of low move-persistence, indicative of area-restricted search-type behaviours, were deemed to indicate evidence of stop-overs along the migratory route. Results Here, we demonstrate two divergent migratory tactics to maintain a similar overall energy-minimization strategy within a single population of narwhal. Narwhal migrating offshore exhibited more tortuous movement trajectories overall with no evidence of spatially-consistent stop-over locations across individuals. Nearshore migrating narwhal undertook more directed routes, contrasted by spatially-explicit stop-over behaviour in highly-productive fjord and canyon systems along the coast of Baffin Island for periods of several days to several weeks. Conclusions Within a single population, divergent migratory tactics can achieve a similar overall energy-minimizing strategy within a species as a response to differing trade-offs between predictable and unpredictable resources. Our methodological approach, which revealed the modulators of fine-scale migratory movements and predicted regional stop-over sites, is widely applicable to a variety of other aquatic and terrestrial species. Quantifying marine migration strategies will be key for adaptive conservation in the face of climate change and ever increasing human pressures.

Abstract Background Due to their Arctic habitat and elusive nature, little is known about the narwhal (Monodon monoceros) and its foraging behaviour. Understanding its ability to catch prey is essential for understanding its ecological role, but also to assess its ability to withstand climate changes and anthropogenic activities. Narwhals produce echolocation clicks and buzzing sounds as part of their foraging behaviour and these can be used as indicators of prey capture attempts. However, acoustic data are expensive to store on the tagging devices and require complicated post-processing. The main goal of this paper is to predict prey capture attempts directly from acceleration and depth data. The aim is to apply broadly used statistical models with interpretable parameters. The ultimate goal is to be able to estimate prey consumption without the more demanding acoustic data. Results We predict narwhal buzzing activity using mixed-effects logistic regression models with 83 features extracted from acceleration and depth data as explanatory variables. The features encompass both instantaneous values as well as delayed values to capture behavioural patterns lasting several seconds. The data correlations were not strong enough to predict the exact timing of the buzzes, but were reliably able to detect buzzes within a few seconds. Most of the of the buzz predictions were within 2 s of an observed buzz (68%), increasing to 94% within 30 s. Conversely, 46% of the observed buzzes were within 2 s of a predicted buzz, increasing to 82% within 30 s. Additionally, the model performed well, although with a tendency towards underestimation of the number of buzzes per dive. In total, we predicted 17, 557 buzzes versus 25, 543 observed across data from 10 narwhals. Classifying foraging and non-foraging dives yielded a precision of 86% and a recall of 91%. Conclusion We conclude that narwhal foraging estimation through acceleration and depth data is a valid alternative or supplement to buzz recordings, even when using somewhat simple statistical methods, such as logistic regression. The methods in this paper can be extended to foraging detection in similar marine species and can aid instrument development.

The Eastern Canada-West Greenland (EC-WG) bowhead whale (Balaena mysticetus) population is widely distributed across the eastern Canadian Arctic and across Baffin Bay to the western coast of Greenland. Their vast distribution makes obtaining population estimates via aerial surveys difficult, as coverage over their entire range is not possible. Genetic mark recapture analyses address this issue, as biopsy samples can be collected at various locales across the EC-WG bowhead whale population’s distribution and microsatellites can be analyzed to identify unique individuals. EC-WG bowhead whales were subject to intense commercial whaling pressure between the early 1700 s and early 1900 s, after which a moratorium on commercial whaling was put in place in 1915. We used available genetic samples from EC-WG bowhead whales in mark recapture models to estimate population abundance from 2012 to 2021 to gain insight on population dynamics ∼100 years post commercial whaling. The preferred model, using a Jolly-Seber structure, estimated the total abundance as 5173 individuals (CI: 3436–7788). Since the cessation of commercial whaling, the population has been thought to be rebounding, which is reflected by gradually increasing abundance estimates, from the low hundreds in the 1970 s and 1980 s, to ∼6000 in the early 2000 s, but our present estimate suggests population abundance may be plateauing well below the pre-commercial whaling carrying capacity estimate. This population estimate for EC-WG bowhead whales is required to update the population dynamics for conservation efforts.

The Spitsbergen population of bowhead whales (Balaena mysticetus) was harvested to near extinction during the whaling era. Here, we show possible signs of recovery of the population by reporting observations of two calves in the Scoresby Sund polynya in East Greenland on 2 and 4 May 2022, and it is the first observation of this kind in this area since the early 1900s.

The elongated, spiraled tusk of male narwhals (Monodon monoceros) grows continuously throughout the life of the whale and is most likely a secondary sexual trait used in male–male hierarchical competition and possibly in female mate choice. Sex determination in narwhals is typically based on the presence (male) or absence (female) of an erupted tusk, but anomalies such as females with tusks, tuskless males or double-tusked whales occur, although infrequently. In this study, we collected reproductive data and recorded the presence or absence of a tusk in narwhals from the Inuit hunt in Greenland (1993 and 2010–19) with the purpose of estimating the frequency of tusk anomalies. We found that of the 173 whales, 2.9% displayed tusk anomalies. Tusked females constituted 1.5% of sampled females, tuskless males 2.8% of sampled males and double-tusked males 0.9% of sampled males. Biological information on a tusked female, a tuskless male and a double-tusked male was collected and is presented here. The tusked female was sexually mature, and 18 ovarian scars (indicating pregnancies) documented a long reproductive lifespan. The complete female tusk was estimated to be between 146 and 151 cm in length. The tuskless male was sexually maturing, as indicated by body dimensions, and the double-tusked male was sexually immature, with the two tusks measuring

A decreasing trend in narwhal (Monodon monoceros Linnaeus, 1758) abundance has been identified in a small population in Scoresby Sound, Southeast Greenland. We hypothesize that excessive hunting has affected life history and population dynamics of this population. Biological information and samples collected from the Inuit hunt, from satellite-tagged narwhals and from official hunters’ reports, were used to estimate age, growth, and reproduction. During 2007 through 2019, a decreasing proportion of young and increasing proportion of older whales were harvested. Male and female body length and male tusk length increased significantly, while body mass of both sexes showed a nonsignificant increase. The probability of catching a female decreased significantly, while a nonsignificant decline of catching a pregnant female was observed in both biological samples and hunters’ reports. Narwhal swimming speeds correlated with fluke widths indicated that larger whales attain greater speeds. The decline in juveniles and females is probably due to an opportunistic hunting practice targeting the easiest-to-catch whales, where bigger whales are faster and more difficult to catch. The cumulative effect of overharvest with a declining proportion of females, an overrepresentation of large males, and a lack of calves and juveniles has detrimental implications for this small narwhal population.

Abstract The East Greenland-Svalbard-Barents Sea (EGSB) bowhead whale stock (Balaena mysticetus) was hunted to near extinction and remains Endangered on the International Union of Conservation of Nature Red List. The intense, temporally extensive hunting pressure may have left the population vulnerable to other perturbations, such as environmental change. However, the lack of genomic baseline data renders it difficult to evaluate the impacts of various potential stressors on this stock. Twelve EGSB bowhead whales sampled in 2017/2018 were re-sequenced and mapped to a previously published draft genome. All individuals were unrelated and void of significant signs of inbreeding, with similar observed and expected homo- and heterozygosity levels. Despite the small population size, mean autosome-wide heterozygosity was 0.00102, which is higher than that of most mammals for which comparable estimates are calculated using the same parameters, and three times higher than a conspecific individual from the Eastern-Canada-West-Greenland bowhead whale stock. Demographic history analyses indicated a continual decrease of N e from ca. 1.5 million to ca. 250,000 years ago, followed by a slight increase until ca. 100,000 years ago, followed by a rapid decrease in N e between 50,000 and 10,000 years ago. These estimates are lower than previously suggested based on mitochondrial DNA, but suggested demographic patterns over time are similar.

Abundance estimation of narwhals is usually done with either visual or photographic aerial surveys. The basic estimation for both methods is detection of whales at the surface, and to obtain fully corrected abundance estimates, the at-surface detections need to be corrected for the proportion of whales that, at any given time, is available to be detected at the surface. The surfacing time or ‘availability correction factor’ is obtained from whales instrumented with dive recorders, that either relay concatenated information on the proportion of time spent at different depth intervals to satellites, or from recovered instruments that collect complete dive profiles, measured at high frequency. Concatenated data binned in depth histograms from Satellite-Linked-Time-Depth-Recorders (SLTDR) falls in two categories, where those that correct the zero depth values with information from the saltwater switch provide larger and apparently more accurate surfacing times, than those collected from instruments that does not correct the zero depth readings. The erroneous detection of near-surface pressure values is likely due to slow response of pressure transducers made from temperature sensitive materials. The high frequency sampling from AcousondeTM recorders documents erroneous surface detections, and adjustments of the dive profiles are needed to obtain realistic near-surface values. Any reconstruction of dive profiles and near-surface values apparently involves some level of corrections and it is recommended, for development of availability correction factors for aerial surveys, that data from zero-adjusted SLTDRs or TDR instruments are used. The mean estimate of surface time from 7 SLTDRs was 29% (CV=0.05). One SLTDR, with steel pressure transducer and zero-adjustments, that was retrieved from the whale, provided a particular long-record (83 days) of reliable high-resolution data. The surface time for this sample was 31%, when calculated as the sum of all depth readings at or above 2 m. The mean of 144 hourly depth readings during 06:00-18:00, and including dives above 3m, was 27.36% (CV=0.8) for 12 days overlapping with the usual timing of aerial surveys. Accurate estimation of smaller depth bins (e.g. 0-1 m) should, even with high resolution instruments, be used with caution when estimating availability bias.

Detecting structures within the continuous diving behavior of marine animals is challenging, and no universal framework is available. We captured such diverse structures using chaos theory. By applying time-delay embedding to exceptionally long dive records (83 d) from the narwhal, we reconstructed the state-space portrait. Using measures of chaos, we detected a diurnal pattern and its seasonal modulation, classified data, and found how sea-ice appearance shifts time budgets. There is more near-surface rest but deeper dives at solar noon, and more intense diving during twilight and at night but to shallower depths (likely following squid); sea-ice appearance reduces rest. The introduced geometrical approach is simple to implement and potentially helpful for mapping and labeling long-term behavioral data, identifying differences between individual animals and species, and detecting perturbations.

[This corrects the article DOI: 10.1371/journal.pcbi.1006425.].

Abstract Implementation of effective conservation planning relies on a robust understanding of the spatiotemporal distribution of the target species. In the marine realm, this is even more challenging for species rarely seen at the sea surface due to their extreme diving behavior like the sperm whales. Our study aims at (a) investigating the seasonal movements, (b) predicting the potential distribution, and (c) assessing the diel vertical behavior of this species in the Mascarene Archipelago in the south‐west Indian Ocean. Using 21 satellite tracks of sperm whales and eight environmental predictors, 14 supervised machine learning algorithms were tested and compared to predict the whales' potential distribution during the wet and dry season, separately. Fourteen of the whales remained in close proximity to Mauritius, while a migratory pattern was evidenced with a synchronized departure for eight females that headed towards Rodrigues Island. The best performing algorithm was the random forest, showing a strong affinity of the whales for sea surface height during the wet season and for bottom temperature during the dry season. A more dispersed distribution was predicted during the wet season, whereas a more restricted distribution to Mauritius and Reunion waters was found during the dry season, probably related to the breeding period. A diel pattern was observed in the diving behavior, likely following the vertical migration of squids. The results of our study fill a knowledge gap regarding seasonal movements and habitat affinities of this vulnerable species, for which a regional IUCN assessment is still missing in the Indian Ocean. Our findings also confirm the great potential of machine learning algorithms in conservation planning and provide highly reproductible tools to support dynamic ocean management.

Abstract Environmental changes are prominent in Arctic ecosystems, where the distribution, abundance, life history, and health of marine organisms such as the bowhead whale (Balaena mysticetus) are tightly connected to sea ice and sea temperature. However, due to logistical and other challenges of data collection in the Arctic, appropriate assessments of past, present and future effects of climate change and human activities are lacking for many Arctic species. Environmental DNA (eDNA) is emerging as a noninvasive and cost‐effective way of obtaining genetic material from the environment and has the potential to complement traditional methods for biodiversity and genetic monitoring. In this study, we investigate whether eDNA isolated from seawater samples has the capacity to capture the genetic diversity of bowhead whales in Disko Bay, West Greenland, for the implementation of long‐term genetic monitoring programs of key Arctic marine species. A total of 41 eDNA “footprint” samples were obtained from the water surface after a whale had dived and an additional 54 eDNA samples were collected along transect lines. Samples were screened for bowhead DNA using a species‐specific qPCR primer and probe assay, and a subset of 30 samples were successfully Sanger‐sequenced to generate individual mitochondrial control region haplotypes. Moreover, by shotgun sequencing ten footprint samples on an Illumina NovaSeq platform we show that footprints generally contain less than 1% endogenous DNA, resulting in partial mitochondrial genomes in four samples out of ten samples. Our findings suggest that sampling in the footprint or wake of traveling animals is a promising method for capturing the genetic diversity of bowhead whales and other marine megafauna. With optimization of sampling and target DNA sequencing for higher endogenous DNA yield, seawater eDNA samples have a large potential for implementation in the long‐term population genetic monitoring of marine megafauna in the Arctic and elsewhere.

Abstract The narwhal (Monodon monoceros) is a high‐Arctic species inhabiting areas that are experiencing increases in sea temperatures, which together with reduction in sea ice are expected to modify the niches of several Arctic marine apex predators. The Scoresby Sound fjord complex in East Greenland is the summer residence for an isolated population of narwhals. The movements of 12 whales instrumented with Fastloc‐GPS transmitters were studied during summer in Scoresby Sound and at their offshore winter ground in 2017–2019. An additional four narwhals provided detailed hydrographic profiles on both summer and winter grounds. Data on diving of the whales were obtained from 20 satellite‐linked time‐depth recorders and 16 Acousonde™ recorders that also provided information on the temperature and depth of buzzes. In summer, the foraging whales targeted depths between 300 and 850 m where the preferred areas visited by the whales had temperatures ranging between 0.6 and 1.5°C (mean = 1.1°C, SD = 0.22). The highest probability of buzzing activity during summer was at a temperature of 0.7°C and at depths > 300 m. The whales targeted similar depths at their offshore winter ground where the temperature was slightly higher (range: 0.7–1.7°C, mean = 1.3°C, SD = 0.29). Both the probability of buzzing events and the spatial distribution of the whales in both seasons demonstrated a preferential selection of cold water. This was particularly pronounced in winter where cold coastal water was selected and warm Atlantic water farther offshore was avoided. It is unknown if the small temperature niche of whales while feeding is because prey is concentrated at these temperature gradients and is easier to capture at low temperatures, or because there are limitations in the thermoregulation of the whales. In any case, the small niche requirements together with their strong site fidelity emphasize the sensitivity of narwhals to changes in the thermal characteristics of their habitats.

One of the last pristine marine soundscapes, the Arctic, is exposed to increasing anthropogenic activities due to climate-induced decrease in sea ice coverage. In this study, we combined movement and behavioral data from animal-borne tags in a controlled sound exposure study to describe the reactions of narwhals, Monodon monoceros, to airgun pulses and ship noise. Sixteen narwhals were live captured and instrumented with satellite tags and Acousonde acoustic-behavioral recorders, and 11 of them were exposed to airgun pulses and vessel sounds. The sound exposure levels (SELs) of pulses from a small airgun (3.4 L) used in 2017 and a larger one (17.0 L) used in 2018 were measured using drifting recorders. The experiment was divided into trials with airgun and ship-noise exposure, intertrials with only ship-noise, and pre- and postexposure periods. Both trials and intertrials lasted ∼4 h on average per individual. Depending on the location of the whales, the number of separate exposures ranged between one and eight trials or intertrials. Received pulse SELs dropped below 130 dB re 1 μPa2 s by 2.5 km for the small airgun and 4–9 km for the larger airgun, and background noise levels were reached at distances of ∼3 and 8–10.5 km, respectively, for the small and big airguns. Avoidance reactions of the whales could be detected at distances >5 km in 2017 and >11 km in 2018 when in line of sight of the seismic vessel. Meanwhile, a ∼30% increase in horizontal travel speed could be detected up to 2 h before the seismic vessel was in line of sight. Applying line of sight as the criterion for exposure thus excludes some potential pre-response effects, and our estimates of effects must therefore be considered conservative. The whales reacted by changing their swimming speed and direction at distances between 5 and 24 km depending on topographical surroundings where the exposure occurred. The propensity of the whales to move towards the shore increased with increasing exposure (i.e., shorter distance to vessels) and was highest with the large airgun used in 2018, where the whales moved towards the shore at distances of 10–15 km. No long-term effects of the response study could be detected.

Summary: The narwhal (Monodon monoceros) is a highly specialized endemic Arctic cetacean, restricted to the Arctic seas bordering the North Atlantic. Low levels of genetic diversity have been observed across several narwhal populations using mitochondrial DNA and microsatellites. Despite this, the global abundance of narwhals was recently estimated at ∼170,000 individuals. However, the species is still considered vulnerable to changing climates due to its high specialization and restricted Arctic distribution. We assembled and annotated a genome from a narwhal from West Greenland. We find relatively low diversity at the genomic scale and show that this did not arise by recent inbreeding, but rather has been stable over an extended evolutionary timescale. We also find that the current large global abundance most likely reflects a recent rapid expansion from a much smaller founding population. : Biological Sciences; Genetics; Evolutionary Biology Subject Areas: Biological Sciences, Genetics, Evolutionary Biology

Intraspecific variation in resource use by individuals of different age, sex or size may reflect differing energetic requirements and physiological constraints. Males and females often show differences in diet owing to sexual size dimorphism, different life histories and/or habitat use. Here, we investigate how sex and size influence the long-term foraging ecology of belugas and narwhals in Greenland, using stable isotopes of carbon and nitrogen from bone collagen. We show that males have a higher trophic level and a larger ecological niche than females in West Greenland belugas and in East Greenland narwhals. In addition, for these two populations, we find that δ15N increases with size, particularly in males. We hypothesize that sexual size dimorphism together with strong maternal investment drive these differences. By contrast, we find no differences in foraging ecology between sexes in West Greenland narwhals and observe no influence of size on trophic level. This may reflect the influence of interspecific competition in West Greenland, where the distributions of belugas and narwhals overlap, and/or geographical resource partitioning among different summer aggregations of narwhals. Our results suggest that sex and size variations in diet are population dependent, and probably the result of varying ecological interactions.

Deep diving air-breathing species by necessity must balance submergence time and level of exercise during breath-holding: a low activity level preserves oxygen stores and allows longer duration submergence whereas high activity levels consume oxygen quickly and shorten submergence time. In this study, we combined high-resolution multi sensor animal-borne tag data to investigate diving behavior and locomotion styles of the narwhal (Monodon monoceros) (n = 13, mean record length 91 h)–a deep diving Arctic species. Narwhals in this study dove down to >800 m but despite the deep diving abilities, one-third of the dives (33%) were shallow (>100 m) and short in duration (

Narwhals (Monodon monoceros) are gregarious toothed whales that strictly reside in the high Arctic. They produce a broad range of signal types; however, studies of narwhal vocalizations have been mostly descriptive of the sounds available in the species' overall repertoire. Little is known regarding the functions of highly stereotyped mixed calls (i.e., biphonations with both sound elements produced simultaneously), although preliminary evidence has suggested that such vocalizations are individually distinctive and function as contact calls. Here we provide evidence that supports this notion in narwhal mother-calf communication. A female narwhal was tagged as part of larger studies on the life history and acoustic behavior of narwhals. At the time of tagging, it became apparent that the female had a calf, which remained close by during the tagging event. We found that the narwhal mother produced a distinct, highly stereotyped mixed call when separated from her calf and immediately after release from capture, which we interpret as preliminary evidence for contact call use between the mother and her calf. The mother's mixed call production occurred continually over the 4.2 day recording period in addition to a second prominent but different stereotyped mixed call which we believe belonged to the narwhal calf. Thus, narwhal mothers produce highly stereotyped contact calls when separated from their calves, and it appears that narwhal calves similarly produce distinct, stereotyped mixed calls which we hypothesize also contribute to maintaining mother-calf contact. We compared this behavior to the acoustic behavior of two other adult females without calves, but also each with a unique, stereotyped call type. While we provide additional support for individual distinctiveness across narwhal contact calls, more research is necessary to determine whether these calls are vocal signatures which broadcast identity.

RNA editing is a post-transcriptional process in which nucleotide changes are introduced into an RNA sequence, many of which can contribute to proteomic sequence variation. The most common type of RNA editing, contributing to nearly 99% of all editing events in RNA, is A-to-I (adenosine-to-inosine) editing mediated by double-stranded RNA-specific adenosine deaminase (ADAR) enzymes. A-to-I editing at ‘recoding’ sites results in non-synonymous substitutions in protein-coding sequences. Here, we present studies of the conservation of A-to-I editing in selected mRNAs between pigs, bowhead whales, humans and two shark species. All examined mRNAs–NEIL1, COG3, GRIA2, FLNA, FLNB, IGFBP7, AZIN1, BLCAP, GLI1, SON, HTR2C and ADAR2 –showed conservation of A-to-I editing of recoding sites. In addition, novel editing sites were identified in NEIL1 and GLI1 in bowhead whales. The A-to-I editing site of human NEIL1 in position 242 was conserved in the bowhead and porcine homologues. A novel editing site was discovered in Tyr244. Differential editing was detected at the two adenosines in the NEIL1 242 codon in both pig and bowhead NEIL1 mRNAs in various tissues and organs. No conservation of editing of KCNB1 and EEF1A mRNAs was seen in bowhead whales. In silico analyses revealed conservation of five adenosines in ADAR2, some of which are subject to A-to-I editing in bowheads and pigs, and conservation of a regulatory sequence in GRIA2 mRNA that is responsible for recognition of the ADAR editing enzyme.

Estimating abundance is one of the most fundamental and important aspects of population biology, with major implications on how the status of a population is perceived and thus on conservation and management efforts. Although typically based on one of two methods (distance sampling or mark-recapture), there are many individual identification methods that can be used for mark-recapture purposes. In recent years, the use of genetic data for individual identification and abundance estimation through mark-recapture analyses have increased, and in some situations such genetic identifications are more efficient than their field-based counterparts for population monitoring. One issue with mark-recapture analyses, regardless of which method of individual identification is used, is that the study area must provide adequate opportunities for “capturing” all individuals within a population. However, many populations are unevenly and widely distributed, making it unfeasible to adequately sample all necessary areas. Here we develop an analytical technique that accounts for unsampled locations, and provides a means to infer “missing” individuals from unsampled locations, and therefore obtain more accurate abundance estimates when it is not possible to sample all sites. This method is validated using simulations and is used to estimate abundance of the Eastern Canada-West Greenland (EC-WG) bowhead whale population. Based on these analyses, the estimated size of this population is 11,747 individuals during the sampling period, with a 95% highest density interval of 8,169–20,043.

Diving behaviour of narwhals is still largely unknown. We use Hidden Markov models (HMMs) to describe the diving behaviour of a narwhal and fit the models to a three-dimensional response vector of maximum dive depth, duration of dives and post-dive surface time of 8,609 dives measured in East Greenland over 83 days, an extraordinarily long and rich data set. Narwhal diving patterns have not been analysed like this before, but in studies of other whale species, response variables have been assumed independent. We extend the existing models to allow for dependence between state distributions, and show that the dependence has an impact on the conclusions drawn about the diving behaviour. We try several HMMs with 2, 3 or 4 states, and with independent and dependent log-normal and gamma distributions, respectively, and different covariates to characterize dive patterns. In particular, diurnal patterns in diving behaviour is inferred, by using periodic B-splines with boundary knots in 0 and 24 hours.

An aerial line transect survey of whales in West and East Greenland was conducted in August-September 2015. The survey covered the area between the coast of West Greenland and offshore (up to 100 km) to the shelf break. In East Greenland, the survey lines covered the area from the coast up to 50 km offshore crossing the shelf break. A total of 423 sightings of 12 cetacean species were obtained and abundance estimates were developed for common minke whale, (Balaenoptera acutorostrata) (32 sightings), fin whale (Balaenoptera physalus) (129 sightings), humpback whale (Megaptera novaeangliae) (84 sightings), harbour porpoise (Phocoena phocoena) (55 sightings), long-finned pilot whale, (Globicephala melas) (42 sightings) and white-beaked dolphin (Lagenorhynchus albirostri) (50 sightings). The developed at-surface abundance estimates were corrected for both perception bias and availability bias if possible. Data on surface corrections for minke whales and harbour porpoises were collected from whales instrumented with satellite-linked time-depth-recorders. Options for estimation methods are presented and the preferred estimates are: minke whales: 5,095 (95% CI: 2,171-11,961) in West Greenland and 2,762 (95% CI: 1,160-6,574) in East Greenland, fin whales: 2,215 (95% CI: 1,017-4,823) in West Greenland and 6,440 (95% CI: 3,901-10,632) in East Greenland, humpback whales: 993 (95% CI: 434-2,272) in West Greenland and 4,223 (95% CI: 1,845-9,666) in East Greenland, harbour porpoises: 83,321 (95% CI: 43,377-160,047) in West Greenland and 1,642 (95% CI: 319-8,464) in East Greenland, pilot whales: 9,190 (95% CI: 3,635-23,234) in West Greenland and 258 (95% CI: 50-1,354) in East Greenland, white-beaked dolphins 15,261 (95% CI: 7,048-33,046) in West Greenland and 11,889 (95% CI: 4,710-30,008) in East Greenland. The abundance of cetaceans in coastal areas of East Greenland has not been estimated before, but the limited historical information from the area indicates that the achieved abundance estimates were remarkably high. When comparing the abundance estimates from 2015 in West Greenland with a similar survey conducted in 2007, there is a clear trend towards lower densities in 2015 for the three baleen whale species and white-beaked dolphins. Harbour porpoises and pilot whales, however, did not show a similar decline. The decline in baleen whale and white-beaked dolphin abundance is likely due to emigration to the East Greenland shelf areas where recent climate driven changes in pelagic productivity may have accelerated favourable conditions for these species.

The aspartic acid racemization (AAR) technique has been applied for age estimation of humans and other mammals for more than four decades. In this study, eye lenses from 124 animals representing 25 mammalian species were collected and D/L ratios obtained using the AAR technique. The animals were either of known age or had the age estimated by other methods. The purpose of the study was to: a) estimate the accuracy of the AAR technique, and b) examine the effect of body temperature on racemization rates. Samples from four of the 25 species covered the range of ages that is needed to estimate species-specific racemization rates. The sample size from a single species of known age, the pygmy goat (Capra hircus, n = 35), was also large enough to investigate the accuracy of ages obtained using the AAR technique. The 35 goats were divided into three datasets: all goats (n = 35), goats >0.5 yrs old (n = 26) and goats >2 yrs old (n = 19). Leave-one-out analyses were performed on the three sets of data. Normalized root mean squared errors for the group of goats >0.5 yrs old were found to be the smallest. The higher variation in D/L measurements found for young goats 0.5 yrs old was for three age groups of the goats: 0.934 yrs for young goats 8 yrs (n = 4). Thus, the age of an adult or an old animal can be predicted with approximately 10% accuracy, whereas the age of a young animal is difficult to predict. A goat specific racemization rate, as a 2kAsp value, was estimated to 0.0107 ± 3.8 x 10-4 SE (n = 26). The 2kAsp values from 12 species, four estimated in this study and another eight published, were used to examine the effect of core body temperature on the rate of racemization. A positive relationship between AAR and temperature was found (r2 = 0.321) but results also suggest that other factors besides temperature are involved in the racemization process in living animals. Based on our results we emphasize that non-species-specific racemization rates should be used with care in AAR age estimation studies and that the period of postnatal growth of the eye lens be considered when estimating species-specific D/L0 values and ages of young individuals.

Changes in climate are rapidly modifying the Arctic environment. As a result, human activities-and the sounds they produce-are predicted to increase in remote areas of Greenland, such as those inhabited by the narwhals (Monodon monoceros) of East Greenland. Meanwhile, nothing is known about these whales' acoustic behavior or their reactions to anthropogenic sounds. This lack of knowledge was addressed by instrumenting six narwhals in Scoresby Sound (Aug 2013-2016) with Acousonde™ acoustic tags and satellite tags. Continuous recordings over up to seven days were used to describe the acoustic behavior of the whales, in particular their use of three types of sounds serving two different purposes: echolocation clicks and buzzes, which serve feeding, and calls, presumably used for social communication. Logistic regression models were used to assess the effects of location in time and space on buzzing and calling rates. Buzzes were mostly produced at depths of 350-650 m and buzzing rates were higher in one particular fjord, likely a preferred feeding area. Calls generally occurred at shallower depths (

The bowhead whale (Balaena mysticetus) is estimated to live over 200 years and is possibly the longest-living mammal. These animals should possess protective molecular adaptations relevant to age-related diseases, particularly cancer. Here, we report the sequencing and comparative analysis of the bowhead whale genome and two transcriptomes from different populations. Our analysis identifies genes under positive selection and bowhead-specific mutations in genes linked to cancer and aging. In addition, we identify gene gain and loss involving genes associated with DNA repair, cell-cycle regulation, cancer, and aging. Our results expand our understanding of the evolution of mammalian longevity and suggest possible players involved in adaptive genetic changes conferring cancer resistance. We also found potentially relevant changes in genes related to additional processes, including thermoregulation, sensory perception, dietary adaptations, and immune response. Our data are made available online (http://www.bowhead-whale.org) to facilitate research in this long-lived species.

Walruses (Odobenus rosmarus) are exploited for subsistence purposes in West Greenland. However, current information about the abundance of walruses subject to harvest in eastern Baffin Bay subject to harvest has been unavailable despite being critical for maintaining sustainable catch levels. Three visual aerial surveys were conducted in 2006 (21 March to 19 April 2006), 2008 (3 to 12 April) and 2012 (24 March to 14 April) to estimate the number of walruses on the wintering grounds in eastern Baffin Bay and Davis Strait. Data on the fraction of walruses that were submerged below a 2m detection threshold during the surveys were obtained from 24 walruses instrumented with satellite-linked-time-depth-recorders in northern Baffin Bay in May-June 2010-2012. An availability correction factor was estimated at 36.5% (cv=0.08) after filtering of data for an observed drift of the pressure transducer of more than 2.5 m. The surveys resulted in walrus abundance estimates that were corrected for walruses submerged below a detection threshold and for walruses that were missed by the observers. The estimates of abundance were 1,105 (cv=0.31, 95% CI 610-2,002) in 2006, 1,137 (0.48, 468-2,758) in 2008 and 1,408 (0.22, 922-2,150) in 2012.

Between 2005 and 2008, 31 walruses were tagged with Argos satellite transmitters at their wintering grounds at Store Hellefiske Banke, Central West Greenland (N=23), and at their summering grounds off the coast of Southeast Baffin Island, Canada (N=8). Two male walruses moved along the Greenland coast from Store Hellefiske Banke north to Disko Banke, where contact was lost. Two other males went further north, up to the Upernavik area. Contact was lost with one of these tags, but the other animal travelled southward again and went towards SE Baffin Island. Eight of the tagged walruses moved between West Greenland and Baffin Island, demonstrating a connection between walruses at these sites. Walruses left the Store Hellefiske Banke during the period 7 April to 17 May and on average used 7 days to swim the 400 km across Davis Strait. The migration routes were quite similar; they travelled over the shallowest areas at the narrowest part of the strait. The timing of the spring dispersal and migration towards Canada was closely linked to the extent and timing of the retreat of the pack ice edge. One flipper tagged male that was marked off South Baffin Island was recovered in a hunt on Store Hellefiske Banke, documenting that the reverse migration also occurs. Off West Greenland satellite tagged walruses spent a lot of time around the Store Hellefiske Banke (55.0o-56.5o W), using this shallow area as feeding grounds irrespective of the ice cover in this area. Partial sexual segregation was observed. Despite a tendency in West Greenland for males to occur farther offshore, farther from the ice edge and at greater depths, only their preference for denser ice cover (64% ice cover) differed significantly (P=0.019) from the habitat preferences of females (52% ice cover). Coast dispersal was more condensed and the segregation between males and females was more pronounced during autumn along the Southeast Baffin Island. Females remained farther north (P

Atlantic walruses (Odobenus rosmarus rosmarus) belonging to the Baffin Bay subpopulation occur year round in the North Water polynya (NOW) between NW Greenland and eastern Ellesmere Island (Canada). They are hunted for subsistence purposes by residents of the Qaanaaq area (NW Greenland) bordering the NOW to the east and by Canadian Inuit at the entrance to Jones Sound in Nunavut. During the open-water period NW Greenland is virtually devoid of walruses which concentrate along eastern and southern Ellesmere Island at this time of the year. To determine the abundance of walruses in the NOW area, aerial surveys were conducted in August of 1999, 2008, and 2009. In July 2009, nine satellite-linked transmitters were deployed in nearby Kane Basin. Surveys on 9 and 20 August 2009 along eastern Ellesmere Island were the most extensive and were augmented with concomitant data on haul-out and at water surface activity from three (1 F, 2 M) of the nine tags that were still functioning. We therefore focus on the 2009 surveys. Walruses were observed on the ice and in water primarily in Buchanan Bay and Princess Marie Bay where the remaining functional tags were located. The Minimum Counted population (MCP) was 571 on 20 August. Adjusting the MCP of walruses on ice for those not hauled out, the estimate of abundance of walruses in the Baffin Bay stock was 1,251(CV=1.00, 95% CI = 1,226) when adjusted by the proportion of tags ‘dry’ at the time of the survey and 1,249 (CV=1.12, 95% CI = 1,370) when adjusted by the average time tags were dry. The surveys did not cover all potential walrus summering habitat along eastern Ellesmere Island and are negatively biased to an unknown degree.

The ages of three species of cetaceans were estimated by counting the growth layer groups (GLG) and measuring the aspartic acid racemization rate (kAsp) by what is referred to as the Aspartic Acid Racemization (AAR) technique. Data on kAsp and the D/L ratio of aspartic acid at birth [(D/L)0] in North Atlantic common minke whales (Balaenoptera acutorostrata) are presented along with data on fin whales (B. physalus) and harbour porpoises (Phocoena phocoena) already published by Nielsen et al. (2012). The kAsp specific for minke whales was 1.40 x 10-3 yr-1 (SE ± 0.00005) and the (D/L)0 was 0.0194 (SE ± 0.0012). The correlation of GLG age and D/L ratio for all three species was highly significant; however, the correlation coefficient varied greatly (fin whales: R2 = 0.59, p

Ages of marine mammals have traditionally been estimated by counting dentinal growth layers in teeth. However, this method is difficult to use on narwhals (Monodon monoceros) because of their special tooth structures. Alternative methods are therefore needed. The aspartic acid racemization (AAR) technique has been used in age estimation studies of cetaceans, including narwhals. The purpose of this study was to estimate a species-specific racemization rate for narwhals by regressing aspartic acid d/l ratios in eye lens nuclei against growth layer groups in tusks (n=9). Two racemization rates were estimated: one by linear regression (r2=0.98) based on the assumption that age was known without error, and one based on a bootstrap study, taking into account the uncertainty in the age estimation (r2 between 0.88 and 0.98). The two estimated 2kAsp values were identical up to two significant figures. The 2k Asp value from the bootstrap study was found to be 0.00229±0.000089 SE, which corresponds to a racemization rate of 0.00114−yr±0.000044 SE. The intercept of 0.0580±0.00185 SE corresponds to twice the (d/l)0 value, which is then 0.0290±0.00093 SE. We propose that this species-specific racemization rate and (d/l)0 value be used in future AAR age estimation studies of narwhals, but also recommend the collection of tusks and eyes of narwhals for further improving the (d/l)0 and 2kAsp estimates obtained in this study.

The beluga whale is one of three endemic Arctic whales. The species is philopatric, and its migration patterns are passed from mother to calf. Management of the species is informed by the levels of genetic structuring among summer aggregation sites based on mitochondrial D-Loop data. To assess the levels of differentiation across the entire mitochondrial genome within belugas, we present a comparison between the first two complete mitochondrial genomes from opposite sides of their distribution range: Baffin Bay and the Russian Far East. Our analyses reveal that additional phylogenetic insights can be gained from expanding the genetic region analyzed. Further, we estimate the divergence time between the two mitochondrial genomes to be 0.469 MYA.

We report the results of an analysis of the variation in the nucleotide sequence of the mitochondrial control region obtained in 218 samples collected from belugas, Delphinapterus leucas, around the Baffin Bay. We detected multiple instances of significant heterogeneity in the distribution of genetic variation among the analyzed mitochondrial control region sequences on a spatial as well as temporal scale indicating a high degree of maternal population structure. The detection of significant levels of heterogeneity between samples collected in different years but within the same area and season was unexpected. Re-examination of earlier results presented by Brown Gladden and coworkers also revealed temporal genetic heterogeneity within the one area where sufficient (n>15) samples were collected in multiple years. These findings suggest that non-random breeding and maternally directed site-fidelity are not the sole causes of genetic heterogeneity among belugas but that a matrilineal pod structure might cause significant levels of genetic heterogeneity as well, even within the same area. We propose that a maternal pod structure, which has been shown to be the cause of significant genetic heterogeneity in other odontocetes, may add to the overall level of heterogeneity in the maternally inherited DNA and hence that much of the spatial heterogeneity observed in this and previous studies might be attributed to pod rather than population structure. Our findings suggest that it is important to estimate the contribution of pod structure to overall heterogeneity before defining populations or management units in order to avoid interpreting heterogeneity due to sampling of different pods as different populations/management units.

Belugas (Delphinapterus leucas) caught by hunters from various hamlets in the Arctic differed in the concentrations of organochlorine contaminants in their blubber. By applying Canonical Discriminant Analysis (CDA) it was possible to separate all seven sampling locations from each other. Over 90% of the samples could be classified back to their landing location based on the data transformations developed by CDA. This analysis suggested that “stock” or management unit for belugas is best described by the culturally transmitted behaviour of their migration route. The analysis also provides evidence that most belugas caught by hunters from Grise Fiord are not the same as belugas caught while migrating along West Greenland; that some belugas caught in Sanikiluaq are not the same as beluga caught in the Nastapoka River estuary; and that the belugas caught in Kimmirut are not the same as belugas caught in Cumberland Sound. There is a need to redefine the stock descriptions of some belugas in Canada and Greenland.

Analysis of 454 stomachs of ringed seals (Phoca hispida) collected from the Inuit hunt in six municipalities in West Greenland and 30 alimentary tracts collected by scientists in East Greenland, showed seasonal and regional differences in the diet. In Northwest and East Greenland polar cod (Boreogadus saida) and Arctic cod (Arctogadus glacialis) were the most dominant prey items. In contrast, seals in central West Greenland mainly preyed upon amphipods (Parathemisto spp.), capelin (Mallotus villosus), redfish (Sebastes sp.) and squid (Gonatus sp.), while capelin was the most important prey item in Southwest Greenland. The material from Uummannaq demonstrates seasonal variations, as euphausiids were common in spring, and snailfish (Liparis spp.) dominated from October through January, where after snailfish disappeared with the formation of fast ice. No age related differences in the diet were found in stomach samples from Avanersuaq, but in Upemavik crustaceans were more abundant in stomach samples from younger immature seals. Greenland halibut (Reinhardtius hippoglossoides) was only present in samples from seals older than 4 years. In Upemavik Arctic cod was the dominant prey item for seals caught in May at the ice edge, whereas polar cod dominated the samples from seals caught in open water.

Seven studies of netting and tagging of ringed seals (Phoca hispida) in Greenland 1976-1997 are described. The areas of operation were: northern part of the Upernavik area (Northwest Greenland), Kong Oscars Fjord (Northeast Greenland), and Avanersuaq (Thule, North Greenland). Altogether, 135 seals were caught, of which 99 were tagged and released with Dalton Jumbo rototags. Recapture of 38 tagged seals has been reported to date. The recaptures demonstrate movements of ringed seals in Greenland of more than 100km within districts, as well as long-distance movements of more than 1000km from the site of release.