Laura N. Kloepper

Audiogram of a formerly stranded long-finned pilot whale (Globicephala melas) measured using auditory evoked potentials.

SUMMARY Long-finned pilot whales are highly social odontocetes found in temperate and subpolar regions. This species is particularly known for its interaction with fisheries as well as its mass strandings. Recent tagging work has provided some information about pilot whales in the wild but, even though they have been successfully kept in captivity, little is known about their sensory capabilities. This study investigates the hearing abilities of a rehabilitated 2 year old male long-finned pilot whale. A complete audiogram was collected using auditory evoked potential techniques that included measurements of nine frequencies from 4 to 100 kHz presented as sinusoidally amplitude-modulated tones. The results indicated that the region of best hearing was between 11.2 and 50 kHz and the subject had relatively poor high frequency hearing compared with other odontocete species. This study emphasizes the importance of collecting basic hearing measurements from new species, understanding diagnostic life histories as well as continuously increasing the sample size of audiometry measurements within and between odontocete species as animals become available.

Active echolocation beam focusing in the false killer whale, Pseudorca crassidens

SUMMARY The odontocete sound production system is highly complex and produces intense, directional signals that are thought to be focused by the melon and the air sacs. Because odontocete echolocation signals are variable and the emitted click frequency greatly affects the echolocation beam shape, investigations of beam focusing must account for frequency-related beam changes. In this study we tested whether the echolocation beam of a false killer whale changed depending on target difficulty and distance while also accounting for frequency-related changes in the echolocation beam. The data indicate that the false killer whale changes its beam size according to target distance and difficulty, which may be a strategy of maximizing the energy of the target echo. We propose that the animal is using a strategy of changing the focal region according to target distance and that this strategy is under active control.

Decreased echolocation performance following high-frequency hearing loss in the false killer whale (Pseudorca crassidens)

SUMMARY Toothed whales and dolphins possess a hypertrophied auditory system that allows for the production and hearing of ultrasonic signals. Although the fossil record provides information on the evolution of the auditory structures found in extant odontocetes, it cannot provide information on the evolutionary pressures leading to the hypertrophied auditory system. Investigating the effect of hearing loss may provide evidence for the reason for the development of high-frequency hearing in echolocating animals by demonstrating how high-frequency hearing assists in the functioning echolocation system. The discrimination abilities of a false killer whale (Pseudorca crassidens) were measured prior to and after documented high-frequency hearing loss. In 1992, the subject had good hearing and could hear at frequencies up to 100 kHz. In 2008, the subject had lost hearing at frequencies above 40 kHz. First in 1992, and then again in 2008, the subject performed an identical echolocation task, discriminating between machined hollow aluminum cylinder targets of differing wall thickness. Performances were recorded for individual target differences and compared between both experimental years. Performances on individual targets dropped between 1992 and 2008, with a maximum performance reduction of 36.1%. These data indicate that, with a loss in high-frequency hearing, there was a concomitant reduction in echolocation discrimination ability, and suggest that the development of a hypertrophied auditory system capable of hearing at ultrasonic frequencies evolved in response to pressures for fine-scale echolocation discrimination.

Change in echolocation signals with hearing loss in a false killer whale (Pseudorca crassidens)

The echolocation signals of a false killer whale (Pseudorca crassidens) were collected during a wall thickness discrimination task and compared to clicks recorded during an identical experiment in 1992. During the sixteen year time period, the subject demonstrated a loss of high frequency hearing of about 70 kHz. Clicks between the two experiments were compared to investigate the effect of hearing loss on echolocation signals. There was a significant reduction in the peak frequency, center frequency and source level of clicks between the two time periods. Additionally, the subject currently produces more signals with low frequency peaks and fewer signals with high frequency peaks than she did in 1992. These results indicate the subject changed its echolocation signals to match its range of best hearing.

A re-evaluation of auditory filter shape in delphinid odontocetes: Evidence of constant-bandwidth filters

The auditory filter shape of delphinid odontocetes was previously considered to be typically mammalian constant-quality in which filter bandwidths increase proportionally with frequency. Recent studies with porpoises demonstrate constant-bandwidth portions of the auditory filter. The critical ratios for a bottlenose dolphin were measured between 40 and 120  kHz by behaviorally determining the subjects ability to detect pure tones in the presence of white noise. Critical ratios as a function of frequency were constant, indicating the auditory filter acts as a constant-bandwidth system in this frequency range. Re-analysis of past studies supports these findings, and suggests the delphinid auditory system is best characterized as a constant-Q system below 40  kHz and a constant-bandwidth-like system between 40  kHz and 120  kHz before returning to a constant Q pattern at the highest frequencies.

Estimating colony sizes of emerging bats using acoustic recordings

The decline of bats demands more widespread monitoring of populations for conservation and management. Current censusing methods are either prone to bias or require costly equipment. Here, we report a new method using passive acoustics to determine bat count census from overall acoustic amplitude of the emerging bat stream. We recorded the video and audio of an emerging colony of Mexican free-tailed bats from two cave locations across multiple nights. Instantaneous bat counts were calculated from the video frames, and the bat stream’s acoustic amplitude corresponding to each video frame was determined using three different methods for calculating acoustic intensity. We found a significant link between all three acoustic parameters and bat count, with the highest R2 of 0.742 linking RMS pressure and bat count. Additionally, the relationship between acoustics and population size at one cave location could accurately predict the population size at another cave location. The data were gathered with low-cost, easy-to-operate equipment, and the data analysis can be easily accomplished using automated scripts or with open-source acoustic software. These results are a potential first step towards creating an acoustic model to estimate bat population at large cave colonies worldwide.

Single-lobed frequency-dependent beam shape in an echolocating false killer whale (Pseudorca crassidens).

Recent studies indicate some odontocetes may produce echolocation beams with a dual-lobed vertical structure. The shape of the odontocete echolocation beam was further investigated in a false killer whale performing an echolocation discrimination task. Clicks were recorded with an array of 16 hydrophones and frequency-dependent amplitude plots were constructed to assess beam shape. The majority of the echolocation clicks were single-lobed in structure with most energy located between 20 and 80 kHz. These data indicate the false killer whale does not produce a dual-lobed structure, as has been shown in bottlenose dolphins, which may be a function of lowered frequencies in the emitted signal due to hearing loss.

Transmission beam characteristics of a Risso's dolphin (Grampus griseus)

The echolocation system of the Rissos dolphin (Grampus griseus) remains poorly studied compared to other odontocete species. In this study, echolocation signals were recorded from a stationary Rissos dolphin with an array of 16 hydrophones and the two-dimensional beam shape was explored using frequency-dependent amplitude plots. Click source parameters were similar to those already described for this species. Centroid frequency of click signals increased with increasing sound pressure level, while the beamwidth decreased with increasing center frequency. Analysis revealed primarily single-lobed, and occasionally vertically dual-lobed, beam shapes. Overall beam directivity was found to be greater than that of the harbor porpoise, bottlenose dolphin, and a false killer whale. The relationship between frequency content, beam directivity, and head size for this Rissos dolphin deviated from the trend described for other species. These are the first reported measurements of echolocation beam shape and directivity in G. griseus.

Spatial orientation of different frequencies within the echolocation beam of a Tursiops truncatus and Pseudorca crassidens

A two-dimensional array of 16 hydrophones was created to map the spatial distribution of different frequencies within the echolocation beam of a Tursiops truncatus and a Pseudorca crassidens. It was previously shown that both the Tursiops and Pseudorca only paid attention to frequencies between 29 and 42 kHz while echolocating. Both individuals tightly focused the 30 kHz frequency and the spatial location of the focus was consistently pointed toward the target. At 50 kHz the beam was less focused and less precisely pointed at the target. At 100 kHz the focus was often completely lost and was not pointed at the target. This indicates that these individuals actively focused the beam toward the target only in the frequency range they paid attention to. Frequencies outside this range were left unfocused and undirected. This focusing was probably achieved through sensorimotor control of the melon morphology and nasal air sacs. This indicates that both morphologically different species can control the spatial distribution of different frequency ranges within the echolocation beam to create consistent ensonation of desired targets.

Cognitive Adaptation of Sonar Gain Control in the Bottlenose Dolphin

Echolocating animals adjust the transmit intensity and receive sensitivity of their sonar in order to regulate the sensation level of their echoes; this process is often termed automatic gain control. Gain control is considered not to be under the animals cognitive control, but previous investigations studied animals ensonifying targets or hydrophone arrays at predictable distances. To test whether animals maintain gain control at a fixed level in uncertain conditions, we measured changes in signal intensity for a bottlenose dolphin (Tursiops truncatus) detecting a target at three target distances (2.5, 4 and 7 m) in two types of sessions: predictable and unpredictable. Predictable sessions presented the target at a constant distance; unpredictable sessions moved the target randomly between the three target positions. In the predictable sessions the dolphin demonstrated intensity distance compensation, increasing the emitted click intensity as the target distance increased. Additionally, as trials within sessions progressed, the animal adjusted its click intensity even from the first click in a click train, which is consistent with the animal expecting a target at a certain range. In the unpredictable sessions there was no significant difference of intensity with target distance until after the 7th click in a click train. Together, these results demonstrate that the bottlenose dolphin uses learning and expectation for sonar gain control.