The nightscape of the Arctic winter shapes the diving behavior of a marine predator.

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.

Stuck in a corner: Anthropogenic noise threatens narwhals in their once pristine Arctic habitat.

Niche-conservative species are especially susceptible to changes in their environment, and detecting the negative effects of new stressors in their habitats is vital for safeguarding of these species. In the Arctic, human disturbance including marine traffic and exploration of resources is increasing rapidly due to climate change-induced reduction of sea ice. Here, we show that the narwhal, Monodon monoceros, is extremely sensitive to human-made noise. Narwhals avoided deep diving (> 350 m) with simultaneous reduction of foraging and increased shallow diving activity as a response to either ship sound alone or ship sound with concurrent seismic airgun pulses. Normal behavior decreased by 50 to 75% at distances where received sound levels were below background noise. Narwhals were equally responsive to both disturbance types, hence demonstrating their acute sensitivity to ship sound. This sensitivity coupled with their special behavioral-ecological strategy including a narrow ecological niche and high site fidelity makes them thus especially vulnerable to human impacts in the Arctic.

Using quantile regression and relative entropy to assess the period of anomalous behavior of marine mammals following tagging.

Tagging of animals induces a variable stress response which following release will obscure natural behavior. It is of scientific relevance to establish methods that assess recovery from such behavioral perturbation and generalize well to a broad range of animals, while maintaining model transparency. We propose two methods that allow for subdivision of animals based on covariates, and illustrate their use on N = 20 narwhals (Monodon monoceros) and N = 4 bowhead whales (Balaena mysticetus), captured and instrumented with Acousonde™ behavioral tags, but with a framework that easily generalizes to other marine animals and sampling units. The narwhals were divided into two groups based on handling time, short ( t < 58 min) and long ( t ≥ 58 min), to measure the effect on recovery. Proxies for energy expenditure (VeDBA) and rapid movement (jerk) were derived from accelerometer data. Diving profiles were characterized using two metrics (target depth and dive duration) derived from depth data. For accelerometer data, recovery was estimated using quantile regression (QR) on the log-transformed response, whereas depth data were addressed using relative entropy (RE) between hourly distributions of dive duration (partitioned into three target depth ranges) and the long-term average distribution. Quantile regression was used to address location-based behavior to accommodate distributional shifts anticipated in aquatic locomotion. For all narwhals, we found fast recovery in the tail of the distribution (<3 h) compared with a variable recovery at the median (∼1-10 h) and with a significant difference between groups separated by handling time. Estimates of bowhead whale recovery times showed fast median recovery (<3 h) and slow recovery at the tail (>6 h), but were affected by substantial uncertainty. For the diving profiles, as characterized by the component pair (target depth, dive duration), the recovery was slower (narwhals-long: t < 16 h; narwhals-short: t < 10 h; bowhead whales: <9 h) and with a difference between narwhals with short vs long handling times. Using simple statistical concepts, we have presented two transparent and general methods for analyzing high-resolution time series data from marine animals, addressing energy expenditure, activity, and diving behavior, and which allows for comparison between groups of animals based on well-defined covariates.

Narwhals react to ship noise and airgun pulses embedded in background noise.

Anthropogenic activities are increasing in the Arctic, posing a threat to niche-conservative species with high seasonal site fidelity, such as the narwhal Monodon monoceros. In this controlled sound exposure study, six narwhals were live-captured and instrumented with animal-borne tags providing movement and behavioural data, and exposed to concurrent ship noise and airgun pulses. All narwhals reacted to sound exposure with reduced buzzing rates, where the response was dependent on the magnitude of exposure defined as 1/distance to ship. Buzzing rate was halved at 12 km from the ship, and whales ceased foraging at 7-8 km. Effects of exposure could be detected at distances > 40 km from the ship.At only a few kilometres from the ship, the received high-frequency cetacean weighted sound exposure levels were below background noise indicating extreme sensitivity of narwhals towards sound disturbance and demonstrating their ability to detect signals embedded in background noise. The narwhal's reactions to sustained disturbance may have a plethora of consequences both at individual and population levels. The observed reactions of the whales demonstrate their auditory sensitivity but also emphasize, that anthropogenic activities in pristine narwhal habitats needs to be managed carefully if healthy narwhal populations are to be maintained.

Evidence of stereotyped contact call use in narwhal (Monodon monoceros) mother-calf communication.

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.

Some like it cold: Temperature-dependent habitat selection by narwhals.

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.

Sea surface temperature predicts the movements of an Arctic cetacean: the bowhead whale.

The effects of climate change constitute a major concern in Arctic waters due to the rapid decline of sea ice, which may strongly alter the movements and habitat availability of Arctic marine mammals. We tracked 98 bowhead whales by satellite over an 11-year period (2001-2011) in Baffin Bay - West Greenland to investigate the environmental drivers (specifically sea surface temperature and sea ice) involved in bowhead whale's movements. Movement patterns differed according to season, with aggregations of whales found at higher latitudes during spring and summer likely in response to sea-ice retreat and increasing sea temperature (SST) facilitated by the warm West Greenland Current. In contrast, the whales moved further south in response to sea temperature decrease during autumn and winter. Statistical models indicated that the whales targeted a narrow range of SSTs from -0.5 to 2 °C. Sea surface temperatures are predicted to undergo a marked increase in the Arctic, which could expose bowhead whales to both thermal stress and altered stratification and vertical transport of water masses. With such profound changes, bowhead whales may face extensive habitat loss. Our results highlight the need for closer investigation and monitoring in order to predict the extent of future distribution changes.

Spatial and temporal patterns of sound production in East Greenland narwhals.

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 (<100 m), with more than half of these calls occurring near the surface (<7 m), where the whales also spent more than half of their time. A period of silence following release, present in all subjects, was attributed to the capture and tagging operations, emphasizing the importance of longer (multi-day) records. This study provides basic life-history information on a poorly known species-and therefore control data in ongoing or future sound-effect studies.

High source levels and small active space of high-pitched song in bowhead whales (Balaena mysticetus).

The low-frequency, powerful vocalizations of blue and fin whales may potentially be detected by conspecifics across entire ocean basins. In contrast, humpback and bowhead whales produce equally powerful, but more complex broadband vocalizations composed of higher frequencies that suffer from higher attenuation. Here we evaluate the active space of high frequency song notes of bowhead whales (Balaena mysticetus) in Western Greenland using measurements of song source levels and ambient noise. Four independent, GPS-synchronized hydrophones were deployed through holes in the ice to localize vocalizing bowhead whales, estimate source levels and measure ambient noise. The song had a mean apparent source level of 185±2 dB rms re 1 µPa @ 1 m and a high mean centroid frequency of 444±48 Hz. Using measured ambient noise levels in the area and Arctic sound spreading models, the estimated active space of these song notes is between 40 and 130 km, an order of magnitude smaller than the estimated active space of low frequency blue and fin whale songs produced at similar source levels and for similar noise conditions. We propose that bowhead whales spatially compensate for their smaller communication range through mating aggregations that co-evolved with broadband song to form a complex and dynamic acoustically mediated sexual display.

Evidence for simultaneous sound production in the bowhead whale (Balaena mysticetus).

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.

Seasonal changes in the vocal behavior of bowhead whales (Balaena mysticetus) in Disko Bay, Western-Greenland.

Singing behavior has been described from bowhead whales in the Bering Sea during their annual spring migration and from Davis Strait during their spring feeding season. It has been suggested that this spring singing behavior is a remnant of the singing during the winter breeding season, though no winter recordings are available. In this study, the authors describe recordings made during the winter and spring months of bowhead whales in Disko Bay, Western-Greenland. A total of 7091 bowhead whale sounds were analyzed to describe the vocal repertoire, the singing behavior, and the changes in vocal behavior from February to May. The vocal signals could be divided into simple (frequency-modulated) calls (n=483), complex (amplitude-modulated) calls (n=635), and song notes (n=5973). Recordings from the end of February to middle of March were characterized by higher call rates with a greater diversity of call types than recordings made later in the season. This study is the first description of bowhead song from the stock in Western-Greenland during both the winter and spring months, and provides support for the hypothesis that song during the winter months contains more song notes than song from the spring making the winter song more variable.