Malene Simon

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

SUMMARY 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.

Behaviour and kinematics of continuous ram filtration in bowhead whales (Balaena mysticetus)

Balaenid whales perform long breath-hold foraging dives despite a high drag from their ram filtration of zooplankton. To maximize the volume of prey acquired in a dive with limited oxygen supplies, balaenids must either filter feed only occasionally when prey density is particularly high, or they must swim at slow speeds while filtering to reduce drag and oxygen consumption. Using digital tags with three-axis accelerometers, we studied bowhead whales feeding off West Greenland and present here, to our knowledge, the first detailed data on the kinematics and swimming behaviour of a balaenid whale filter feeding at depth. Bowhead whales employ a continuous fluking gait throughout the bottom phase of foraging dives, moving at very slow speeds (less than 1 m s−1), allowing them to filter feed continuously at depth. Despite the slow speeds, the large mouth aperture provides a water filtration rate of approximately 3 m3 s−1, amounting to some 2000 tonnes of water and prey filtered per dive. We conclude that a food niche of dense, slow-moving zooplankton prey has led balaenids to evolve locomotor and filtering systems adapted to work against a high drag at swimming speeds of less than 0.07 body length s−1 using a continuous fluking gait very different from that of nekton-feeding, aquatic predators.

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.

The relationship between the acoustic behaviour and surface activity of killer whales (Orcinus orca) that feed on herring (Clupea harengus)

We describe the acoustic behaviour of piscivorous killer whales in Norwegian and Icelandic waters. Whales were assigned to one of three activities (feeding, travelling or other), and sound recordings were made in their proximity with a single hydrophone and a digital audiotape (DAT) recorder. A quantitative analysis of the production of pulsed calls, whistles and echolocation clicks in the three activities revealed that there was a significant effect of activity on the production of these sound types. Both killer whales in Icelandic and Norwegian waters produced high rates of clicks and calls during feeding and low rates of click, calls and whistles during travelling. The differences can be used as acoustical markers and provides new possibilities for acoustic monitoring of killer whales in these areas. Based on the similarity between their prey choice, hunting strategies, phenotype and acoustic behaviour, we suggest that the killer whales in Icelandic and Norwegian waters belong to the same ecotype: Scandinavian herring-eating killer whales.

Singing behavior of fin whales in the Davis Strait with implications for mating, migration and foraging

Most baleen whales undertake migrations between low-latitude breeding grounds and high-latitude feeding grounds. Though little is known about the timing of their migration from the Arctic, fin whales are assumed to undertake a similar migratory pattern. To address questions about habitat use and migrations, the acoustic activity of fin whales in Davis Strait, between Greenland and Canada, was monitored continuously for two years using three bottom-moored acoustic recorders. The acoustic power in the fin whale call frequencies peaked in November-December, showing that fin whales are present in Davis Strait much later in the year than previously expected. The closely timed peaks in song activity and conception time imply that not all fin whales migrate south to mate, but rather start mating at high latitudes rather than or before migrating. Singing activity was strongly linked to daylight hours, suggesting that fin whales might feed during the few daylight hours of the late fall and early Arctic winter. A negative correlation between the advancing sea ice front and power in fin whale frequencies indicates that future changes in sea ice conditions from global warming might change the distribution and migratory patterns of fin whales near the poles.

Passive acoustic monitoring of bottlenose dolphin and harbour porpoise, in Cardigan Bay, Wales, with implications for habitat use and partitioning

Knowledge about harbour porpoise and bottlenose dolphin occurrence in Cardigan Bay Special Area of Conservation (SAC), Wales, is limited to daylight hours during summer, when conditions are suitable for traditional visual surveys. T-PODs are autonomous instruments programmed to log time-cues of species-specific echolocation signals for long periods of time. Here we investigated bottlenose dolphin and harbour porpoise habitat use and partitioning by deploying ten calibrated T-PODs in Cardigan Bay SAC for one year. The T-PODs detected both species all year round with a peak of detections in April-October for dolphins and in October-March for porpoise, revealing a previously unknown importance of the place to harbour porpoise during winter. Though the two species are sympatric, simultaneous detections of both species were rare and indication of temporal habitat partitioning between the two species in some parts of the SAC was observed. The one location where simultaneous detections were not as rare was close to the stretch of shoreline where stranding of porpoises killed by dolphins are most common, suggesting that the observed spatiotemporal overlap leads to inter-specific interactions, in some cases fatal for the porpoise.

Acoustic characteristics of underwater tail slaps used by Norwegian and Icelandic killer whales (Orcinus orca) to debilitate herring (Clupea harengus)

SUMMARY Norwegian killer whales debilitate prey by slapping their tails into herring schools. These underwater tail slaps produce a thud-like sound. It is unclear whether this sound is caused by cavitation and/or physical contact between herring and whale tail. Also the forces causing debilitation of the fish are not understood. Here we present an acoustic analysis of underwater tail slaps using a multi-channel wide (150 kHz) band recording system. Underwater tail slaps produced by Norwegian killer whales generated sounds consisting of multiple pulses with source levels of 186±5.4 dB (pp) re.1 μPa at 1 m (±1 s.d., N=4). The –3 dB and 97% energy bandwidths were 36.8±22.5 kHz and 130.5±17.5 kHz (±1 s.d., N=13), respectively, with a centre frequency of 46.1±22.3 kHz. The similarities between the acoustic properties of underwater tail slaps recorded from killer whales in Norway, and thud-like sounds recorded from killer whales in Iceland suggest that Norwegian and Icelandic killer whales use similar hunting techniques. The acoustic characteristics of sounds produced by underwater tail slaps were similar to the ones from other cavitation sound sources described in the literature, both in term of temporal and frequency features as well as in source level. We suggest that multiple factors generated by the tail slaps like particle fluctuations, turbulence, pressure changes and physical impact cause debilitation of herring.

Echolocation clicks from killer whales (Orcinus orca) feeding on herring (Clupea harengus)

Echolocation clicks from Norwegian killer whales feeding on herring schools were recorded using a four-hydrophone array. The clicks had broadband bimodal frequency spectra with low and high frequency peaks at 24 and 108 kHz, respectively. The -10 dB bandwidth was 35 kHz. The average source level varied from 173 to 202 dB re 1 microPa (peak-to-peak) at 1 m. This is considerably lower than source levels described for Canadian killer whales foraging on salmon. It is suggested that biosonar clicks of Norwegian killer whales are adapted for localization of prey with high target strength and acute hearing abilities.

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.

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.