Narwhal (Monodon monoceros) echolocation click rates to support cue counting passive acoustic density estimation.

Estimating animal abundance is fundamental for effective management and conservation. It is increasingly done by combining passive acoustics with knowledge about rates at which animals produce cues (cue rates). Narwhals (Monodon monoceros) are elusive marine mammals for which passive acoustic density estimation might be plausible, but for which cue rates are lacking. Clicking rates in narwhals were investigated using a dataset from sound and movement tag records collected in August 2013-2016 and 2019 in East Greenland. Clicking rates were quantified for ∼1200 one-second-long systematic random samples from 8 different whales. Generalized additive models were used to model (1) the probability of being in a clicking state versus depth and (2) the clicking rate while in a clicking state, versus time and depth. The probability of being in a clicking state increased with depth, reaching ∼1.0 at ∼500 m, while the number of clicks per second (while in a clicking state) increased with depth. The mean cue production rate, weighted by tag duration, was 1.28 clicks per second (se = 0.13, CV = 0.10). This first cue rate for narwhals may be used for cue counting density estimation, but care should be taken if applying it to other geographical areas or seasons, given sample size, geographical, and temporal limitations.

The active space of sperm whale codas: inter-click information for intra-unit communication.

Sperm whales (Physeter macrocephalus) are social mega-predators who form stable matrilineal units that often associate within a larger vocal clan. Clan membership is defined by sharing a repertoire of coda types consisting of specific temporal spacings of multi-pulsed clicks. It has been hypothesized that codas communicate membership across socially segregated sympatric clans, but others propose that codas are primarily used for behavioral coordination and social cohesion within a closely spaced social unit. Here, we test these hypotheses by combining measures of ambient noise levels and coda click source levels with models of sound propagation to estimate the active space of coda communication. Coda clicks were localized off the island of Dominica with a four- or five-element 80 m vertical hydrophone array, allowing us to calculate the median RMS source levels of 1598 clicks from 444 codas to be 161 dB re. 1 µPa (IQR 153-167), placing codas among the most powerful communication sounds in toothed whales. However, together with measured ambient noise levels, these source levels lead to a median active space of coda communication of ∼4 km, reflecting the maximum footprint of a single foraging sperm whale unit. We conclude that while sperm whale codas may contain information about clan affiliation, their moderate active space shows that codas are not used for long range acoustic communication between units and clans, but likely serve to mediate social cohesion and behavioral transitions in intra-unit communication.

Directional biosonar beams allow echolocating harbour porpoises to actively discriminate and intercept closely spaced targets.

Echolocating toothed whales face the problem that high sound speeds in water mean that echoes from closely spaced targets will arrive at time delays within their reported auditory integration time of some 264 µs. Here, we test the hypothesis that echolocating harbour porpoises cannot resolve and discriminate targets within a clutter interference zone given by their integration time. To do this, we trained two harbour porpoises (Phocoena phocoena) to actively approach and choose between two spherical targets at four varying inter-target distances (13.5, 27, 56 and 108 cm) in a two-alternative forced-choice task. The free-swimming, blindfolded porpoises were tagged with a sound and movement tag (DTAG4) to record their echoic scene and acoustic outputs. The known ranges between targets and the porpoise, combined with the sound levels received on target-mounted hydrophones revealed how the porpoises controlled their acoustic gaze. When targets were close together, the discrimination task was more difficult because of smaller echo time delays and lower echo level ratios between the targets. Under these conditions, buzzes were longer and started from farther away, source levels were reduced at short ranges, and the porpoises clicked faster, scanned across the targets more, and delayed making their discrimination decision until closer to the target. We conclude that harbour porpoises can resolve and discriminate closely spaced targets, suggesting a clutter rejection zone much shorter than their auditory integration time, and that such clutter rejection is greatly aided by spatial filtering with their directional biosonar beam.

Echolocation click parameters and biosonar behaviour of the dwarf sperm whale (Kogia sima).

Dwarf sperm whales (Kogia sima) are small toothed whales that produce narrow-band high-frequency (NBHF) echolocation clicks. Such NBHF clicks, subject to high levels of acoustic absorption, are usually produced by small, shallow-diving odontocetes, such as porpoises, in keeping with their short-range echolocation and fast click rates. Here, we sought to address the problem of how the little-studied and deep-diving Kogia can hunt with NBHF clicks in the deep sea. Specifically, we tested the hypotheses that Kogia produce NBHF clicks with longer inter-click intervals (ICIs), higher directionality and higher source levels (SLs) compared with other NBHF species. We did this by deploying an autonomous deep-water vertical hydrophone array in the Bahamas, where no other NBHF species are present, and by taking opportunistic recordings of a close-range Kogia sima in a South African harbour. Parameters from on-axis clicks (n=46) in the deep revealed very narrow-band clicks (root mean squared bandwidth, BWRMS, of 3±1 kHz), with SLs of up to 197 dB re. 1 µPa peak-to-peak (µPapp) at 1 m, and a half-power beamwidth of 8.8 deg. Their ICIs (mode of 245 ms) were much longer than those of porpoises (<100 ms), suggesting an inspection range that is longer than detection ranges of single prey, perhaps to facilitate auditory streaming of a complex echo scene. On-axis clicks in the shallow harbour (n=870) had ICIs and SLs in keeping with source parameters of other NBHF cetaceans. Thus, in the deep, dwarf sperm whales use a directional, but short-range echolocation system with moderate SLs, suggesting a reliable mesopelagic prey habitat.

Do echolocating toothed whales direct their acoustic gaze on- or off-target in a static detection task?

Echolocating mammals produce directional sound beams with high source levels to improve echo-to-noise ratios and reduce clutter. Recent studies have suggested that the differential spectral gradients of such narrow beams are exploited to facilitate target localization by pointing the beam slightly off targets to maximize the precision of angular position estimates [maximizing bearing Fisher information (FI)]. Here, we test the hypothesis that echolocating toothed whales focus their acoustic gaze askew during target detection to maximize spectral cues by investigating the acoustic gaze direction of two trained delphinids (Tursiops truncatus and Pseudorca crassidens) echolocating to detect an aluminum cylinder behind a hydrophone array in a go/no-go paradigm. The animals rarely placed their beam axis directly on the target, nor within the narrow range around the off-axis angle that maximizes FI. However, the target was, for each trial, ensonified within the swath of the half-power beam width, and hence we conclude that the animals solved the detection task using a strategy that seeks to render high echo-to-noise ratios rather than maximizing bearing FI. We posit that biosonar beam adjustment and acoustic gaze strategies are likely task-dependent and that maximizing bearing FI by pointing off-axis does not improve target detection performance.

High resolution three-dimensional beam radiation pattern of harbour porpoise clicks with implications for passive acoustic monitoring.

The source properties and radiation patterns of animal vocalisations define, along with propagation and noise conditions, the active space in which these vocalisations can be detected by conspecifics, predators, prey, and by passive acoustic monitoring (PAM). This study reports the 4π (360° horizontal and vertical) beam profile of a free-swimming, trained harbour porpoise measured using a 27-element hydrophone array. The forward echolocation beam is highly directional, as predicted by a piston model, and is consistent with previous measurements. However, at off-axis angles greater than ±30°, the beam attenuates more rapidly than the piston model and no side lobes are present. A diffuse back beam is also present with levels about -30 dB relative to the source level. In PAM, up to 50% of detections can be from portions of the beam profile with distorted click spectra, although this drops substantially for higher detection thresholds. Simulations of the probability of acoustically detecting a harbour porpoise show that a traditional piston model can underestimate the probability of detection compared to the actual three-dimensional radiation pattern documented here. This highlights the importance of empirical 4π measurements of beam profiles of toothed whales, both to improve understanding of toothed whale biology and to inform PAM.

Passive acoustic methods for fine-scale tracking of harbour porpoises in tidal rapids.

The growing interest in generating electrical power from tidal currents using tidal turbine generators raises a number of environmental concerns, including the risk that marine mammals might be injured or killed through collision with rotating turbine blades. To understand this risk, information on how marine mammals use tidal rapid habitats and in particular, their underwater movements and dive behaviour is required. Porpoises, which are the most abundant small cetacean at most European tidal sites, are difficult animals to tag, and the limited size of tidal habitats means that any telemetered animal would be likely to spend only a small proportion of time within them. Here, an alternative approach is explored, whereby passive acoustic monitoring (PAM) is used to obtain fine scale geo-referenced tracks of harbour porpoises in tidal rapid areas. Large aperture hydrophone arrays are required to obtain accurate locations of animals from PAM data and automated algorithms are necessary to process the large quantities of acoustic data collected on such systems during a typical survey. Methods to automate localisation, including a method to match porpoise detections on different hydrophones and separate different vocalising animals, and an assessment of the localisation accuracy of the large aperture hydrophone array are presented.