Time-varying auditory gain control in response to double-pulse stimuli in harbour porpoises is not mediated by a stapedial reflex

Echolocating animals reduce their output level and hearing sensitivity with decreasing echo delays, presumably to stabilize the perceived echo intensity during target approaches. In bats, this variation in hearing sensitivity is formed by a call-induced stapedial reflex that tapers off over time after the call. Here, we test the hypothesis that a similar mechanism exists in toothed whales by subjecting a trained harbour porpoise to a series of double sound pulses varying in delay and frequency, while measuring the magnitudes of the evoked auditory brainstem responses (ABRs). We find that the recovery of the ABR to the second pulse is frequency dependent, and that a stapedial reflex therefore cannot account for the reduced hearing sensitivity at short pulse delays. We propose that toothed whale auditory time-varying gain control during echolocation is not enabled by the middle ear as in bats, but rather by frequency-dependent mechanisms such as forward masking and perhaps higher-order control of efferent feedback to the outer hair cells

Characteristics and propagation of airgun pulses in shallow water with implications for effects on small marine mammals

Airguns used in seismic surveys are among the most prevalent and powerful anthropogenic noise sources in marine habitats. They are designed to produce most energy below 100 Hz, but the pulses have also been reported to contain medium-to-high frequency components with the potential to affect small marine mammals, which have their best hearing sensitivity at higher frequencies. In shallow water environments, inhabited by many of such species, the impact of airgun noise may be particularly challenging to assess due to complex propagation conditions. To alleviate the current lack of knowledge on the characteristics and propagation of airgun pulses in shallow water with implications for effects on small marine mammals, we recorded pulses from a single airgun with three operating volumes (10 in3, 25 in3 and 40 in3) at six ranges (6, 120, 200, 400, 800 and 1300 m) in a uniform shallow water habitat using two calibrated Reson 4014 hydrophones and four DSG-Ocean acoustic data recorders. We show that airgun pulses in this shallow habitat propagated out to 1300 meters in a way that can be approximated by a 18log(r) geometric transmission loss model, but with a high pass filter effect from the shallow water depth. Source levels were back-calculated to 192 dB re µPa2s (sound exposure level) and 200 dB re 1 µPa dB Leq-fast (rms over 125 ms duration), and the pulses contained substantial energy up to 10 kHz, even at the furthest recording station at 1300 meters. We conclude that the risk of causing hearing damage when using single airguns in shallow waters is small for both pinnipeds and porpoises. However, there is substantial potential for significant behavioral responses out to several km from the airgun, well beyond the commonly used shut-down zone of 500 meters

Estimated communication range and energetic cost of bottlenose dolphin whistles in a tropical habitat

Author Posting. © Acoustical Society of America, 2012. This article is posted here by permission of Acoustical Society of America for personal use, not for redistribution. The definitive version was published in Journal of the Acoustical Society of America 131 (2012): 582-592, doi:10.1121/1.3662067.Bottlenose dolphins (Tursiops sp.) depend on frequency-modulated whistles for many aspects of their social behavior, including group cohesion and recognition of familiar individuals. Vocalization amplitude and frequency influences communication range and may be shaped by many ecological and physiological factors including energetic costs. Here, a calibrated GPS-synchronized hydrophone array was used to record the whistles of bottlenose dolphins in a tropical shallow-water environment with high ambient noise levels. Acoustic localization techniques were used to estimate the source levels and energy content of individual whistles. Bottlenose dolphins produced whistles with mean source levels of 146.7±6.2 dB re. 1 μPa(RMS). These were lower than source levels estimated for a population inhabiting the quieter Moray Firth, indicating that dolphins do not necessarily compensate for the high noise levels found in noisy tropical habitats by increasing their source level. Combined with measured transmission loss and noise levels, these source levels provided estimated median communication ranges of 750 m and maximum communication ranges up to 5740 m. Whistles contained less than 17 mJ of acoustic energy, showing that the energetic cost of whistling is small compared to the high metabolic rate of these aquatic mammals, and unlikely to limit the vocal activity of toothed whales.This study received support from the Danish Ph.D. School of Aquatic Sciences (SOAS), Aarhus University, DK, WWF Verdensnaturfonden and Aase & Ejnar Danielsens Foundation, the Siemens Foundation, the Faculty of Science at the University of Aarhus, DK, and the Danish Natural Science Foundation via a Steno scholarship and a logistics grant to PTM

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

Amazon river dolphins ( Inia geoffrensis ) modify biosonar output level and directivity during prey interception in the wild

Toothed whales have evolved to live in extremely different habitats and yet they all rely strongly on echolocation for finding and catching prey. Such biosonar-based foraging involves distinct phases of searching for, approaching and capturing prey, where echolocating animals gradually adjust sonar output to actively shape the flow of sensory information. Measuring those outputs in absolute levels requires hydrophone arrays centred on the biosonar beam axis, but this has never been done for wild toothed whales approaching and capturing prey. Rather, field studies make the assumption that toothed whales will adjust their biosonar in the same manner to arrays as they will when approaching prey. To test this assumption, we recorded wild botos (Inia geoffrensis) as they approached and captured dead fish tethered to a hydrophone in front of a star-shaped seven-hydrophone array. We demonstrate that botos gradually decrease interclick intervals and output levels during prey approaches, using stronger adjustment magnitudes than predicted from previous boto array data. Prey interceptions are characterised by high click rates, but although botos buzz during prey capture, they do so at lower click rates than marine toothed whales, resulting in a much more gradual transition from approach phase to buzzing. We also demonstrate for the first time that wild toothed whales broaden biosonar beamwidth when closing in on prey, as is also seen in captive toothed whales and bats, thus resulting in a larger ensonified volume around the prey, probably aiding prey tracking by decreasing the risk of prey evading ensonification

Behavioural impact assessment of unmanned aerial vehicles on Weddell seals (Leptonychotes weddellii)

The rapid increase in the use of unmanned aerial vehicles (UAVs) in wildlife research has raised concerns about its potential negative impact on animals. The paucity of studies and the variability of responses of pinnipeds to UAVs prompts the need for species-specific impact assessments. Here we assessed the potential behavioural impact of low altitude UAVs on Weddell seals (Leptonychotes weddellii). This is a preliminary step to envisage the feasibility of replacing and/or complementing traditional ground-based behavioural and morphometric measurements by potentially less invasive UAV aerial images. We flew a small UAV (DJI Mavic 2 zoom fitted with a phocid seal audiogram weighted source level of 84 dB re 20μPa (rms)) over 37 Weddell seals (3 adult males, 12 adult females and 22 mother-pup pairs) during the breeding season at Dumont D'Urville, East Antarctica. For each individual, we assessed the level of reaction during UAV overflights at three altitudes (25, 20 and 15 m) while factoring in pup presence and wind speed. For all altitudes and observations pooled together, Weddell seals predominantly (88%) showed little (vigilant) or no (resting) reactions towards the UAV. Moreover, only 27% of all individuals changed their initial activity during the sampling periods, and mothers rarely ended their nursing bouts (3%). While reactions were low overall, the probability of a stronger reaction occurring increased at lower altitudes, and varied among individuals. Neither the presence of pups nor a change in wind speed appear to influence individuals' response to the UAV significantly. However, on simpler histogram representations of the dataset, we observed the strongest reactions for females (n = 5) with a pup at wind speeds below 5 m.s−1 when ambient noise levels were lowest. While Weddell seals are likely to hear the UAV at 25 to 15 m altitude in low wind speeds, the low-level responses we observed are unlikely to negatively impact their energetic budget and/or reproductive success. Our results suggest a low impact of small UAV overflights of Weddell seals during the breeding season when flying ≥25 m. This allows for collection of high resolution images for behavioural and morphometric studies that can potentially replace more invasive data collection when capturing and handling the animals

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