Latencies of click-evoked auditory responses in a harbor porpoise exceed the time interval between subsequent echolocation clicks.

Most auditory evoked potential (AEP) studies in echolocating toothed whales measure neural responses to outgoing clicks and returning echoes using short-latency auditory brainstem responses (ABRs) arising a few ms after acoustic stimuli. However, little is known about longer-latency cortical AEPs despite their relevance for understanding echo processing and auditory stream segregation. Here, we used a non-invasive AEP setup with low click repetition rates on a trained harbor porpoise to test the long-standing hypothesis that echo information from distant targets is completely processed before the next click is emitted. We reject this hypothesis by finding reliable click-related AEP peaks with latencies of 90 and 160 ms, which are longer than 99% of click intervals used by echolocating porpoises, demonstrating that some higher-order echo processing continues well after the next click emission even during slow clicking. We propose that some of the echo information, such as range to evasive prey, is used to guide vocal-motor responses within 50-100 ms, but that information used for discrimination and auditory scene analysis is processed more slowly, integrating information over many click-echo pairs. We conclude by showing theoretically that the identified long-latency AEPs may enable hearing sensitivity measurements at frequencies ten times lower than current ABR methods.

Basin-wide contributions to the underwater soundscape by multiple seismic surveys with implications for marine mammals in Baffin Bay, Greenland.

Seismic surveys increasingly operate in deeper Arctic waters with propagation conditions and marine mammal fauna different from the better-studied temperate, or shallow-water, regions. Using 31 calibrated sound recorders, we quantified noise contributions from four concurrent seismic surveys in Baffin Bay, Greenland, to estimate their potential impacts on marine mammals. The impact was cumulative as the noise level rose in response to the onset of each survey: on a minute-by-minute scale the sound-exposure-levels varied by up to 70 dB (20 dB on average), depending on range to the seismic vessel, local bathymetry effects and interference patterns, representing a significant change in the auditory scene for marine mammals. Airgun pulse energy did not decrease to ambient before arrival of the next pulse leaving very little low-frequency masking-free time. Overall, the measured values matched well with pre-season-modeling, emphasizing the importance of noise-modeling in impact assessments, if responses of focal marine mammals are known.

Single source sound production and dynamic beam formation in echolocating harbour porpoises (Phocoena phocoena).

Echolocating toothed whales produce high-powered clicks by pneumatic actuation of phonic lips in their nasal complexes. All non-physeteroid toothed whales have two pairs of phonic lips allowing many of these species to produce both whistles and clicks at the same time. That has led to the hypothesis that toothed whales can increase the power outputs and bandwidths of clicks, and enable fast clicking and beam steering by acutely timed actuation of both phonic lip pairs simultaneously. Here we test that hypothesis by applying suction cup hydrophones on the sound-producing nasal complexes of three echolocating porpoises (Phocoena phocoena) with symmetrical pairs of phonic lips. Using time of arrival differences on three hydrophones, we show that all recorded clicks from these three porpoises are produced by the right pair of phonic lips with no evidence of simultaneous or independent actuation of the left pair. It is demonstrated that porpoises, despite actuation of only one sound source, can change their output and sound beam probably through conformation changes in the sound-producing soft tissues and nasal sacs, and that the coupling of the phonic lips and the melon acts as a waveguide for sound energy between 100 and 160 kHz to generate a forward-directed sound beam for echolocation.

Echolocation in sympatric Peale's dolphins (Lagenorhynchus australis) and Commerson's dolphins (Cephalorhynchus commersonii) producing narrow-band high-frequency clicks.

An increasing number of smaller odontocetes have recently been shown to produce stereotyped narrow-band high-frequency (NBHF) echolocation clicks. Click source parameters of NBHF clicks are very similar, and it is unclear whether the sonars of individual NBHF species are adapted to specific habitats or the presence of other NBHF species. Here, we test whether sympatric NBHF species sharing the same habitat show similar adaptations in their echolocation clicks and whether their clicks display signs of character displacement. Wide-band sound recordings were obtained with a six-element hydrophone array from wild Peale's (Lagenorhynchus australis) and Commerson's (Cephalorhynchus commersonii) dolphins off the Falkland Islands. The centroid frequency was different between Commerson's (133+/-2 kHz) and Peale's (129+/-3 kHz) dolphins. The r.m.s. bandwidth was 12+/-3 kHz for both species. The source level was higher for Peale's dolphin (185+/-6 dB re 1 muPa p.-p.) than for Commerson's (177+/-5 dB re 1 muPa p.-p.). The mean directivity indexes were 25 dB for both species. The relatively low source levels in combination with the high directivity index may be an adaptation to reduce clutter when foraging in a coastal environment. We conclude that the small species-specific shifts in distribution of centroid frequencies around 130 kHz may reflect character displacement in otherwise-stereotyped NBHF clicks.

Feeding at a high pitch: source parameters of narrow band, high-frequency clicks from echolocating off-shore hourglass dolphins and coastal Hector's dolphins.

Toothed whales depend on echolocation for orientation and prey localization, and source parameters of echolocation clicks from free-ranging animals therefore convey valuable information about the acoustic physiology and behavioral ecology of the recorded species. Recordings of wild hourglass (Lagenorhynchus cruciger) and Hector's dolphins (Cephalorhynchus hectori) were made in the Drake Passage (between Tierra del Fuego and the Antarctic Peninsular) and Banks Peninsular (Akaroa Harbour, New Zealand) with a four element hydrophone array. Analysis of source parameters shows that both species produce narrow band high-frequency (NBHF) echolocation clicks. Coastal Hector's dolphins produce clicks with a mean peak frequency of 129 kHz, 3 dB bandwidth of 20 kHz, 57 micros, 10 dB duration, and mean apparent source level (ASL) of 177 dB re 1 microPa(p.-p.). The oceanic hourglass dolphins produce clicks with mean peak frequency of 126 kHz, 3 dB bandwidth of 8 kHz, 116 micros, 10 dB duration, and a mean estimated ASL of 197 dB re 1 microPa(p.-p.). Thus, hourglass dolphins apparently produce clicks of higher source level, which should allow them to detect prey at more than twice the distance compared to Hector's dolphins. The observed source parameter differences within these two NBHF species may be an adaptation to a coastal cluttered environment versus a deep water, pelagic habitat.

Source levels and harmonic content of whistles in white-beaked dolphins (Lagenorhynchus albirostris).

Recordings of white-beaked dolphin whistles were made in Faxafl6i Bay (Iceland) using a three-hydrophone towed linear array. Signals from the hydrophones were routed through an amplifier to a lunch box computer on board the boat and digitized using a sample rate of 125 kHz per channel. Using this method more than 5000 whistles were recorded. All recordings were made in sea states 0-1 (Beaufort scale). Dolphins were located in a 2D horizontal plane by using the difference of arrival time to the three hydrophones, and source levels were estimated from these positions using two different methods (I and II). Forty-three whistles gave a reliable location for the vocalizing dolphin when using method II and of these 12 when using method I. Source level estimates on the center hydrophone were higher using method I [average source level 148 (rms) +/- 12 dB, n = 36] than for method II [average source level 139 (rms) +/- 12 dB, n = 36]. Using these rms values the maximum possible communication range for whistling dolphins given the local ambient noise conditions was then estimated. The maximum range was 10.5 km for a dolphin whistle with the highest source level (167 dB) and about 140 m for a whistle with the lowest source level (118 dB). Only two of the 43 whistles contained an unequal number of harmonics recorded at the three hydrophones judging from the spectrograms. Such signals could be used to calculate the directionality of whistles, but more recordings are necessary to describe the directionality of white-beaked dolphin whistles.

Bat sonar: an alternative interpretation of the 10-ns jitter result.

In 1990 Simmons et al. reported evidence of a time resolution hitherto unknown in any animal, namely a 10-ns jitter detection threshold in echolocating bats. This result is discussed. The calibration data from the original papers are examined. Observations indicating other cues than delay being presented to the bats are given. We offer an alternative explanation for the psychometric jitter function, based on the assumption of a subtle distortion due to impedance mismatch in the delay-producing apparatus. We also report that effects of impedance mismatch are detectable by a human subject in a model experiment.