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.

DNA methylation-based biomarkers for ageing long-lived cetaceans.

Epigenetic approaches for estimating the age of living organisms are revolutionizing studies of long-lived species. Molecular biomarkers that allow age estimates from small tissue biopsies promise to enhance studies of long-lived whales, addressing a fundamental and challenging parameter in wildlife management. DNA methylation (DNAm) can affect gene expression, and strong correlations between DNAm patterns and age have been documented in humans and nonhuman vertebrates and used to construct "epigenetic clocks". We present several epigenetic clocks for skin samples from two of the longest-lived cetaceans, killer whales and bowhead whales. Applying the mammalian methylation array to genomic DNA from skin samples we validate four different clocks with median errors of 2.3-3.7 years. These epigenetic clocks demonstrate the validity of using cytosine methylation data to estimate the age of long-lived cetaceans and have broad applications supporting the conservation and management of long-lived cetaceans using genomic DNA from remote tissue biopsies.

Combining δ13C and δ15N from bone and dentine in marine mammal palaeoecological research: insights from toothed whales.

Stable carbon (δ13C) and nitrogen (δ15N) isotopic compositions of bone and dentine collagen extracted from museum specimens have been widely used to study the paleoecology of past populations. Due to possible systematic differences in stable isotope values between bone and dentine, dentine values need to be transformed into bone-collagen equivalent using a correction factor to allow comparisons between the two collagen sources. Here, we provide correction factors to transform dentine δ13C and δ15N values into bone-collagen equivalent for two toothed whales: narwhal and beluga. We sampled bone and dentine from the skulls of 11 narwhals and 26 belugas. In narwhals, dentine was sampled from tusk and embedded tooth; in belugas, dentine was sampled from tooth. δ13C and δ15N were measured, and intra-individual bone and dentine isotopic compositions were used to calculate correction factors for each species. We detected differences in δ13C and δ15N. In both narwhals and belugas, we found lower average δ13C and δ15N in bone compared with dentine. The correction factors provided by the study enable the combined analysis of stable isotope data from bone and dentine in these species.

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.

Population-specific sex and size variation in long-term foraging ecology of belugas and narwhals.

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 δ 15N 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.

Analysis of narwhal tusks reveals lifelong feeding ecology and mercury exposure.

The ability of animals to respond to changes in their environment is critical to their persistence. In the Arctic, climate change and mercury exposure are two of the most important environmental threats for top predators.1-3 Rapid warming is causing precipitous sea-ice loss, with consequences on the distribution, composition, and dietary ecology of species4-7 and, thus, exposure to food-borne mercury.8 Current understanding of global change and pollution impacts on Arctic wildlife relies on single-time-point individual data representing a snapshot in time. These data often lack comprehensive temporal resolution and overlook the cumulative lifelong nature of stressors as well as individual variation. To overcome these challenges, we explore the unique capacity of narwhal tusks to characterize chronological lifetime biogeochemical profiles, allowing for investigations of climate-induced dietary changes and contaminant trends. Using temporal patterns of stable isotopes (δ13C and δ15N) and mercury concentrations in annually deposited dentine growth layer groups in 10 tusks from Northwest Greenland (1962-2010), we show surprising plasticity in narwhal feeding ecology likely resulting from climate-induced changes in sea-ice cover, biological communities, and narwhal migration. Dietary changes consequently impacted mercury exposure primarily through trophic magnification effects. Mercury increased log-linearly over the study period, albeit with an unexpected rise in recent years, likely caused by increased emissions and/or greater bioavailability in a warmer, ice-free Arctic. Our findings are consistent with an emerging pattern in the Arctic of reduced sea-ice leading to changes in the migration, habitat use, food web, and contaminant exposure in Arctic top predators.

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.

Influence of past climate change on phylogeography and demographic history of narwhals, Monodon monoceros.

The Arctic is warming at an unprecedented rate, with unknown consequences for endemic fauna. However, Earth has experienced severe climatic oscillations in the past, and understanding how species responded to them might provide insight into their resilience to near-future climatic predictions. Little is known about the responses of Arctic marine mammals to past climatic shifts, but narwhals (Monodon monoceros) are considered one of the endemic Arctic species most vulnerable to environmental change. Here, we analyse 121 complete mitochondrial genomes from narwhals sampled across their range and use them in combination with species distribution models to elucidate the influence of past and ongoing climatic shifts on their population structure and demographic history. We find low levels of genetic diversity and limited geographic structuring of genetic clades. We show that narwhals experienced a long-term low effective population size, which increased after the Last Glacial Maximum, when the amount of suitable habitat expanded. Similar post-glacial habitat release has been a key driver of population size expansion of other polar marine predators. Our analyses indicate that habitat availability has been critical to the success of narwhals, raising concerns for their fate in an increasingly warming Arctic.

The longer the better: evidence that narwhal tusks are sexually selected.

Once thought to be the magical horn of a unicorn, narwhal tusks are one of the most charismatic structures in biology. Despite years of speculation, little is known about the tusk's function, because narwhals spend most of their lives hidden underneath the Arctic ice. Some hypotheses propose that the tusk has sexual functions as a weapon or as a signal. By contrast, other hypotheses propose that the tusk functions as an environmental sensor. Since assessing the tusks function in nature is difficult, we can use the morphological relationships of tusk size with body size to understand this mysterious trait. To do so, we collected morphology data on 245 adult male narwhals over the course of 35 years. Based on the disproportional growth and large variation in tusk length we found, we provide the best evidence to date that narwhal tusks are indeed sexually selected. By combining our results on tusk scaling with known material properties of the tusk, we suggest that the narwhal tusk is a sexually selected signal that is used during male-male contests.

Narwhal Genome Reveals Long-Term Low Genetic Diversity despite Current Large Abundance Size.

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.