Accurate species classification of Arctic toothed whale echolocation clicks using one-third octave ratios.

Passive acoustic monitoring has been an effective tool to study cetaceans in remote regions of the Arctic. Here, we advance methods to acoustically identify the only two Arctic toothed whales, the beluga (Delphinapterus leucas) and narwhal (Monodon monoceros), using echolocation clicks. Long-term acoustic recordings collected from moorings in Northwest Greenland were analyzed. Beluga and narwhal echolocation signals were distinguishable using spectrograms where beluga clicks had most energy >30 kHz and narwhal clicks had a sharp lower frequency limit near 20 kHz. Changes in one-third octave levels (TOL) between two pairs of one-third octave bands were compared from over one million click spectra. Narwhal clicks had a steep increase between the 16 and 25 kHz TOL bands that was absent in beluga click spectra. Conversely, beluga clicks had a steep increase between the 25 and 40 kHz TOL bands that was absent in narwhal click spectra. Random Forest classification models built using the 16 to 25 kHz and 25 to 40 kHz TOL ratios accurately predicted the species identity of 100% of acoustic events. Our findings support the use of echolocation TOL ratios in future automated click classifiers for acoustic monitoring of Arctic toothed whales and potentially for other odontocete species.

First documented movement of a humpback whale between the Cape Verde Islands and West Greenland.

The endangered population of humpback whales (Megaptera novaeangliae) breeding and calving off the Cape Verde Islands (CVI) are known to migrate to feeding areas located along the eastern margin of the North Atlantic Ocean (Iceland, and Norway). Here, we report for the first time a confirmed migration of an individual humpback whale from CVI breeding ground to a western North Atlantic feeding ground of West Greenland. This individual humpback, which was photographed and identified off the coast of West Greenland in 2021, was previously documented in CVI 22 years before (1999). An annual subsistence hunt for humpbacks occurs in West Greenland and the resighting at this location with a humpback whale from CVI has strong implications for the conservation efforts of the small CVI population.

Closing the Organofluorine Mass Balance in Marine Mammals Using Suspect Screening and Machine Learning-Based Quantification.

High-resolution mass spectrometry (HRMS)-based suspect and nontarget screening has identified a growing number of novel per- and polyfluoroalkyl substances (PFASs) in the environment. However, without analytical standards, the fraction of overall PFAS exposure accounted for by these suspects remains ambiguous. Fortunately, recent developments in ionization efficiency (IE) prediction using machine learning offer the possibility to quantify suspects lacking analytical standards. In the present work, a gradient boosted tree-based model for predicting log IE in negative mode was trained and then validated using 33 PFAS standards. The root-mean-square errors were 0.79 (for the entire test set) and 0.29 (for the 7 PFASs in the test set) log IE units. Thereafter, the model was applied to samples of liver from pilot whales (n = 5; East Greenland) and white beaked dolphins (n = 5, West Greenland; n = 3, Sweden) which contained a significant fraction (up to 70%) of unidentified organofluorine and 35 unquantified suspect PFASs (confidence level 2-4). IE-based quantification reduced the fraction of unidentified extractable organofluorine to 0-27%, demonstrating the utility of the method for closing the fluorine mass balance in the absence of analytical standards.

Quantifying marine traffic intensity in Northwest Greenland and the potential disturbance of two seabird colonies.

Marine traffic poses a growing threat to wildlife in the marine environment, including Arctic seabirds, which are exposed to high vessel densities when breeding in coastal areas. However, little is known about the magnitude of the problem. Here, we utilized underwater acoustic monitoring to quantify marine traffic and above-water disturbances at two thick-billed murre colonies in Greenland in 2016. We detected a total of 307 vessels, and only 4 % was known from automatic monitoring systems. Based on proximity, noise emission, and boating behavior, we classified 11 vessels as disturbing and an additional 12 as potentially disturbing for the seabirds. One colony facing population decline was located closest to the main boating route and experienced 2-5 times more disturbances than the other (increasing) colony, suggesting a negative impact of marine traffic. Our study shows that underwater acoustics can be a useful method to quantify above-water disturbances of seabird colonies.

Alternative and legacy flame retardants in marine mammals from three northern ocean regions.

Flame retardants are globally distributed contaminants that have been linked to negative health effects in humans and wildlife. As top predators, marine mammals bioaccumulate flame retardants and other contaminants in their tissues which is one of many human-imposed factors threatening population health. While some flame retardants, such as the polybrominated diphenyl ethers (PBDE), have been banned because of known toxicity and environmental persistence, limited data exist on the presence and distribution of current-use alternative flame retardants in marine mammals from many industrialized and remote regions of the world. Therefore, this study measured 44 legacy and alternative flame retardants in nine marine mammal species from three ocean regions: the Northwest Atlantic, the Arctic, and the Baltic allowing for regional, species, age, body condition, temporal, and tissue comparisons to help understand global patterns. PBDE concentrations were 100-1000 times higher than the alternative brominated flame retardants (altBFRs) and Dechloranes. 2,2',4,5,5'-pentabromobiphenyl (BB-101) and hexabromobenzene (HBBZ) were the predominant altBFRs, while Dechlorane-602 was the predominant Dechlorane. This manuscript also reports only the second detection of hexachlorocyclopentadienyl-dibromocyclooctane (HCDBCO) in marine mammals. The NW Atlantic had the highest PBDE concentrations followed by the Baltic and Arctic which reflects greater historical use of PBDEs in North America compared to Europe and greater industrialization of North America and Baltic countries compared to the Arctic. Regional patterns for other compounds were more complicated, and there were significant interactions among species, regions, body condition and age class. Lipid-normalized PBDE concentrations in harbor seal liver and blubber were similar, but HBBZ and many Dechloranes had higher concentrations in liver, indicating factors other than lipid dynamics affect the distribution of these compounds. The health implications of contamination by this mixture of compounds are of concern and require further research.

Cheap gulp foraging of a giga-predator enables efficient exploitation of sparse prey.

The giant rorqual whales are believed to have a massive food turnover driven by a high-intake lunge feeding style aptly described as the world's largest biomechanical action. This high-drag feeding behavior is thought to limit dive times and constrain rorquals to target only the densest prey patches, making them vulnerable to disturbance and habitat change. Using biologging tags to estimate energy expenditure as a function of feeding rates on 23 humpback whales, we show that lunge feeding is energetically cheap. Such inexpensive foraging means that rorquals are flexible in the quality of prey patches they exploit and therefore more resilient to environmental fluctuations and disturbance. As a consequence, the food turnover and hence the ecological role of these marine giants have likely been overestimated.

Scaling of maneuvering performance in baleen whales: larger whales outperform expectations.

Despite their enormous size, whales make their living as voracious predators. To catch their much smaller, more maneuverable prey, they have developed several unique locomotor strategies that require high energetic input, high mechanical power output and a surprising degree of agility. To better understand how body size affects maneuverability at the largest scale, we used bio-logging data, aerial photogrammetry and a high-throughput approach to quantify the maneuvering performance of seven species of free-swimming baleen whale. We found that as body size increases, absolute maneuvering performance decreases: larger whales use lower accelerations and perform slower pitch-changes, rolls and turns than smaller species. We also found that baleen whales exhibit positive allometry of maneuvering performance: relative to their body size, larger whales use higher accelerations, and perform faster pitch-changes, rolls and certain types of turns than smaller species. However, not all maneuvers were impacted by body size in the same way, and we found that larger whales behaviorally adjust for their decreased agility by using turns that they can perform more effectively. The positive allometry of maneuvering performance suggests that large whales have compensated for their increased body size by evolving more effective control surfaces and by preferentially selecting maneuvers that play to their strengths.

Passive acoustic monitoring reveals year-round marine mammal community composition off Tasiilaq, Southeast Greenland.

Climate-driven changes are affecting sea ice conditions off Tasiilaq, Southeast Greenland, with implications for marine mammal distributions. Knowledge about marine mammal presence, biodiversity, and community composition is key to effective conservation and management but is lacking, especially during winter months. Seasonal patterns of acoustic marine mammal presence were investigated relative to sea ice concentration at two recording sites between 2014 and 2018, with one (65.6°N, 37.4°W) or three years (65.5°N, 38.0°W) of passive acoustic recordings. Seven marine mammal species were recorded. Bearded seals were acoustically dominant during winter and spring, whereas sperm, humpback, and fin whales dominated during the sea ice-free summer and autumn. Narwhals, bowhead, and killer whales were recorded only rarely. Song-fragments of humpback whales and acoustic presence of fin whales in winter suggest mating-associated behavior taking place in the area. Ambient noise levels in 1/3-octave level bands (20, 63, 125, 500, 1000, and 4000 Hz), ranged between 75.6 to 105 dB re 1 µPa. This study provides multi-year insights into the coastal marine mammal community composition off Southeast Greenland and suggests that the Tasiilaq area provides suitable habitat for various marine mammal species year-round.

Strong and lasting impacts of past global warming on baleen whales and their prey.

Global warming is affecting the population dynamics and trophic interactions across a wide range of ecosystems and habitats. Translating these real-time effects into their long-term consequences remains a challenge. The rapid and extreme warming period that occurred after the Last Glacial Maximum (LGM) during the Pleistocene-Holocene transition (7-12 thousand years ago) provides an opportunity to gain insights into the long-term responses of natural populations to periods with global warming. The effects of this post-LGM warming period have been assessed in many terrestrial taxa, whereas insights into the impacts of rapid global warming on marine taxa remain limited, especially for megafauna. In order to understand how large-scale climate fluctuations during the post-LGM affected baleen whales and their prey, we conducted an extensive, large-scale analysis of the long-term effects of the post-LGM warming on abundance and inter-ocean connectivity in eight baleen whale and seven prey (fish and invertebrates) species across the Southern and the North Atlantic Ocean; two ocean basins that differ in key oceanographic features. The analysis was based upon 7032 mitochondrial DNA sequences as well as genome-wide DNA sequence variation in 100 individuals. The estimated temporal changes in genetic diversity during the last 30,000 years indicated that most baleen whale populations underwent post-LGM expansions in both ocean basins. The increase in baleen whale abundance during the Holocene was associated with simultaneous changes in their prey and climate. Highly correlated, synchronized and exponential increases in abundance in both baleen whales and their prey in the Southern Ocean were indicative of a dramatic increase in ocean productivity. In contrast, the demographic fluctuations observed in baleen whales and their prey in the North Atlantic Ocean were subtle, varying across taxa and time. Perhaps most important was the observation that the ocean-wide expansions and decreases in abundance that were initiated by the post-LGM global warming, continued for millennia after global temperatures stabilized, reflecting persistent, long-lasting impacts of global warming on marine fauna.

Soundscape and ambient noise levels of the Arctic waters around Greenland.

A longer Arctic open water season is expected to increase underwater noise levels due to anthropogenic activities such as shipping, seismic surveys, sonar, and construction. Many Arctic marine mammal species depend on sound for communication, navigation, and foraging, therefore quantifying underwater noise levels is critical for documenting change and providing input to management and legislation. Here we present long-term underwater sound recordings from 26 deployments around Greenland from 2011 to 2020. Ambient noise was analysed in third octave and decade bands and further investigated using generic detectors searching for tonal and transient sounds. Ambient noise levels partly overlap with previous Arctic observations, however we report much lower noise levels than previously documented, specifically for Melville Bay and the Greenland Sea. Consistent seasonal noise patterns occur in Melville Bay, Baffin Bay and Greenland Sea, with noise levels peaking in late summer and autumn correlating with open water periods and seismic surveys. These three regions also had similar tonal detection patterns that peaked in May/June, likely due to bearded seal vocalisations. Biological activity was more readily identified using detectors rather than band levels. We encourage additional research to quantify proportional noise contributions from geophysical, biological, and anthropogenic sources in Arctic waters.

Cryptic vocal behavior of foraging humpback whales on feeding grounds in West Greenland.

Male humpback whales (Megaptera novaeangliae) sing in mating aggregations in the form of song displays, but much less is known about how both sexes use sound on their feeding grounds. Here, we test different hypotheses about the function of vocalizations in 14 foraging humpback whales tagged with sound and movement recording Dtags in Greenland. We show that this population of foraging humpback whales have an overall low call rate of 11.9 calls h-1 (inter-quartile range = 12.1) with no support for the hypotheses that they employ sound in the localization or manipulation of prey nor in the coordination of lunge feeding. The calls had a mean received level of 135 ± 5dB re 1 µPa, which is some 30 dB lower than maximum levels of song recorded on similar deployed tags, suggesting a much smaller active space of these vocalizations. This reduced active space might, in concert with low call rates, serve to mitigate eavesdropping by predatory killer whales or conspecifics competing for the same prey resources. We conclude that feeding humpback whales in Greenland produce low level, infrequent calls suggesting that calling is not a prerequisite for successful feeding, but likely serves to mediate within group social interactions.

Energetic and physical limitations on the breaching performance of large whales.

The considerable power needed for large whales to leap out of the water may represent the single most expensive burst maneuver found in nature. However, the mechanics and energetic costs associated with the breaching behaviors of large whales remain poorly understood. In this study we deployed whale-borne tags to measure the kinematics of breaching to test the hypothesis that these spectacular aerial displays are metabolically expensive. We found that breaching whales use variable underwater trajectories, and that high-emergence breaches are faster and require more energy than predatory lunges. The most expensive breaches approach the upper limits of vertebrate muscle performance, and the energetic cost of breaching is high enough that repeated breaching events may serve as honest signaling of body condition. Furthermore, the confluence of muscle contractile properties, hydrodynamics, and the high speeds required likely impose an upper limit to the body size and effectiveness of breaching whales.

Scaling of swimming performance in baleen whales.

The scale dependence of locomotor factors has long been studied in comparative biomechanics, but remains poorly understood for animals at the upper extremes of body size. Rorqual baleen whales include the largest animals, but we lack basic kinematic data about their movements and behavior below the ocean surface. Here, we combined morphometrics from aerial drone photogrammetry, whale-borne inertial sensing tag data and hydrodynamic modeling to study the locomotion of five rorqual species. We quantified changes in tail oscillatory frequency and cruising speed for individual whales spanning a threefold variation in body length, corresponding to an order of magnitude variation in estimated body mass. Our results showed that oscillatory frequency decreases with body length (∝length-0.53) while cruising speed remains roughly invariant (∝length0.08) at 2 m s-1 We compared these measured results for oscillatory frequency against simplified models of an oscillating cantilever beam (∝length-1) and an optimized oscillating Strouhal vortex generator (∝length-1). The difference between our length-scaling exponent and the simplified models suggests that animals are often swimming non-optimally in order to feed or perform other routine behaviors. Cruising speed aligned more closely with an estimate of the optimal speed required to minimize the energetic cost of swimming (∝length0.07). Our results are among the first to elucidate the relationships between both oscillatory frequency and cruising speed and body size for free-swimming animals at the largest scale.

Don't forget the porpoise: acoustic monitoring reveals fine scale temporal variation between bottlenose dolphin and harbour porpoise in Cardigan Bay SAC.

Populations of bottlenose dolphin and harbour porpoise inhabit Cardigan Bay, which was designated a Special Area of Conservation (SAC), with bottlenose dolphin listed as a primary feature for its conservation status. Understanding the abundance, distribution and habitat use of species is fundamental for conservation and the implementation of management. Bottlenose dolphin and harbour porpoise usage of feeding sites within Cardigan Bay SAC was examined using passive acoustic monitoring. Acoustic detections recorded with calibrated T-PODs (acoustic data loggers) indicated harbour porpoise to be present year round and in greater relative abundance than bottlenose dolphin. Fine-scale temporal partitioning between the species occurred at three levels: (1) seasonal differences, consistent between years, with porpoise detections peaking in winter months and dolphin detections in summer months; (2) diel variation, consistent across sites, seasons and years, with porpoise detections highest at night and dolphin detections highest shortly after sunrise; and (3) tidal variation was observed with peak dolphin detections occurring during ebb at the middle of the tidal cycle and before low tide, whereas harbour porpoise detections were highest at slack water, during and after high water with a secondary peak recorded during and after low water. General Additive Models (GAMs) were applied to better understand the effects of each covariate. The reported abundance and distribution of the two species, along with the temporal variation observed, have implications for the design and management of protected areas. Currently, in the UK, no SACs have been formally designated for harbour porpoise while three exist for bottlenose dolphins. Here, we demonstrate a need for increased protection and species-specific mitigation measures for harbour porpoise.

Eye lens radiocarbon reveals centuries of longevity in the Greenland shark (Somniosus microcephalus).

The Greenland shark (Somniosus microcephalus), an iconic species of the Arctic Seas, grows slowly and reaches >500 centimeters (cm) in total length, suggesting a life span well beyond those of other vertebrates. Radiocarbon dating of eye lens nuclei from 28 female Greenland sharks (81 to 502 cm in total length) revealed a life span of at least 272 years. Only the smallest sharks (220 cm or less) showed signs of the radiocarbon bomb pulse, a time marker of the early 1960s. The age ranges of prebomb sharks (reported as midpoint and extent of the 95.4% probability range) revealed the age at sexual maturity to be at least 156 ± 22 years, and the largest animal (502 cm) to be 392 ± 120 years old. Our results show that the Greenland shark is the longest-lived vertebrate known, and they raise concerns about species conservation.

Keeping momentum with a mouthful of water: behavior and kinematics of humpback whale lunge feeding.

Rorqual baleen whales lunge feed by engulfment of tons of prey-laden water in a large and expandable buccal pouch. According to prior interpretations, feeding rorquals are brought to a near-halt at the end of each lunge by drag forces primarily generated by the open mouth. Accelerating the body from a standstill is energetically costly and is purported to be the key factor determining oxygen consumption in lunge-feeding rorquals, explaining the shorter dive times than expected given their sizes. Here, we use multi-sensor archival tags (DTAGs) sampling at high rates in a fine-scale kinematic study of lunge feeding to examine the sequence of events within lunges and how energy may be expended and conserved in the process of prey capture. Analysis of 479 lunges from five humpback whales reveals that the whales accelerate as they acquire prey, opening their gape in synchrony with strong fluke strokes. The high forward speed (mean depth rate: 2.0±0.32 m s(-1)) during engulfment serves both to corral active prey and to expand the ventral margin of the buccal pouch and so maximize the engulfed water volume. Deceleration begins after mouth opening when the pouch nears full expansion and momentum starts to be transferred to the engulfed water. Lunge-feeding humpback whales time fluke strokes throughout the lunge to impart momentum to the engulfed water mass and so avoid a near or complete stop, but instead continue to glide at ~1-1.5 m s(-1) after the lunge has ended. Subsequent filtration and prey handling appear to take an average of 46 s and are performed in parallel with re-positioning for the next lunge.

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.

Source parameters of echolocation clicks from wild bottlenose dolphins (Tursiops aduncus and Tursiops truncatus).

The Indian Ocean and Atlantic bottlenose dolphins (Tursiops aduncus and Tursiops truncatus) are among the best studied echolocating toothed whales. However, almost all echolocation studies on bottlenose dolphins have been made with captive animals, and the echolocation signals of free-ranging animals have not been quantified. Here, biosonar source parameters from wild T. aduncus and T. truncatus were measured with linear three- and four-hydrophone arrays in four geographic locations. The two species had similar source parameters, with source levels of 177-228 dB re 1 µPa peak to peak, click durations of 8-72 µs, centroid frequencies of 33-109 kHz and rms bandwidths between 23 and 54 kHz. T. aduncus clicks had a higher frequency emphasis than T. truncatus. The transmission directionality index was up to 3 dB higher for T. aduncus (29 dB) as compared to T. truncatus (26 dB). The high directionality of T. aduncus does not appear to be only a physical consequence of a higher frequency emphasis in clicks, but may also be caused by differences in the internal properties of the sound production system.

Comparison of echolocation clicks from geographically sympatric killer whales and long-finned pilot whales (L).

The source characteristics of biosonar signals from sympatric killer whales and long-finned pilot whales in a Norwegian fjord were compared. A total of 137 pilot whale and more than 2000 killer whale echolocation clicks were recorded using a linear four-hydrophone array. Of these, 20 pilot whale clicks and 28 killer whale clicks were categorized as being recorded on-axis. The clicks of pilot whales had a mean apparent source level of 196 dB re 1 µPa pp and those of killer whales 203 dB re 1 µPa pp. The duration of pilot whale clicks was significantly shorter (23 µs, S.E.=1.3) and the centroid frequency significantly higher (55 kHz, S.E.=2.1) than killer whale clicks (duration: 41 µs, S.E.=2.6; centroid frequency: 32 kHz, S.E.=1.5). The rate of increase in the accumulated energy as a function of time also differed between clicks from the two species. The differences in duration, frequency, and energy distribution may have a potential to allow for the distinction between pilot and killer whale clicks when using automated detection routines for acoustic monitoring.