Marla M. Holt

Speaking up: Killer whales (Orcinus orca) increase their call amplitude in response to vessel noise

This study investigated the effects of anthropogenic sound exposure on the vocal behavior of free-ranging killer whales. Endangered Southern Resident killer whales inhabit areas including the urban coastal waters of Puget Sound near Seattle, WA, where anthropogenic sounds are ubiquitous, particularly those from motorized vessels. A calibrated recording system was used to measure killer whale call source levels and background noise levels (1-40 kHz). Results show that whales increased their call amplitude by 1 dB for every 1 dB increase in background noise levels. Furthermore, nearby vessel counts were positively correlated with these observed background noise levels.

Comparative assessment of amphibious hearing in pinnipeds

Auditory sensitivity in pinnipeds is influenced by the need to balance efficient sound detection in two vastly different physical environments. Previous comparisons between aerial and underwater hearing capabilities have considered media-dependent differences relative to auditory anatomy, acoustic communication, ecology, and amphibious life history. New data for several species, including recently published audiograms and previously unreported measurements obtained in quiet conditions, necessitate a re-evaluation of amphibious hearing in pinnipeds. Several findings related to underwater hearing are consistent with earlier assessments, including an expanded frequency range of best hearing in true seals that spans at least six octaves. The most notable new results indicate markedly better aerial sensitivity in two seals (Phoca vitulina and Mirounga angustirostris) and one sea lion (Zalophus californianus), likely attributable to improved ambient noise control in test enclosures. An updated comparative analysis alters conventional views and demonstrates that these amphibious pinnipeds have not necessarily sacrificed aerial hearing capabilities in favor of enhanced underwater sound reception. Despite possessing underwater hearing that is nearly as sensitive as fully aquatic cetaceans and sirenians, many seals and sea lions have retained acute aerial hearing capabilities rivaling those of terrestrial carnivores.

Effects of noise levels and call types on the source levels of killer whale calls

Accurate parameter estimates relevant to the vocal behavior of marine mammals are needed to assess potential effects of anthropogenic sound exposure including how masking noise reduces the active space of sounds used for communication. Information about how these animals modify their vocal behavior in response to noise exposure is also needed for such assessment. Prior studies have reported variations in the source levels of killer whale sounds, and a more recent study reported that killer whales compensate for vessel masking noise by increasing their call amplitude. The objectives of the current study were to investigate the source levels of a variety of call types in southern resident killer whales while also considering background noise level as a likely factor related to call source level variability. The source levels of 763 discrete calls along with corresponding background noise were measured over three summer field seasons in the waters surrounding the San Juan Islands, WA. Both noise level and call type were significant factors on call source levels (1-40 kHz band, range of 135.0-175.7 dB(rms) re 1 [micro sign]Pa at 1 m). These factors should be considered in models that predict how anthropogenic masking noise reduces vocal communication space in marine mammals.

A vessel noise budget for Admiralty Inlet, Puget Sound, Washington (USA)

One calendar year of Automatic Identification System (AIS) ship-traffic data was paired with hydrophone recordings to assess ambient noise in northern Admiralty Inlet, Puget Sound, WA (USA) and to quantify the contribution of vessel traffic. The study region included inland waters of the Salish Sea within a 20 km radius of the hydrophone deployment site. Spectra and hourly, daily, and monthly ambient noise statistics for unweighted broadband (0.02-30 kHz) and marine mammal, or M-weighted, sound pressure levels showed variability driven largely by vessel traffic. Over the calendar year, 1363 unique AIS transmitting vessels were recorded, with at least one AIS transmitting vessel present in the study area 90% of the time. A vessel noise budget was calculated for all vessels equipped with AIS transponders. Cargo ships were the largest contributor to the vessel noise budget, followed by tugs and passenger vessels. A simple model to predict received levels at the site based on an incoherent summation of noise from different vessels resulted in a cumulative probability density function of broadband sound pressure levels that shows good agreement with 85% of the temporal data.

Onset, growth, and recovery of in-air temporary threshold shift in a California sea lion (Zalophus californianus)

A California sea lion (Zalophus californianus) was tested in a behavioral procedure to assess noise-induced temporary threshold shift (TTS) in air. Octave band fatiguing noise was varied in both duration (1.5-50 min) and level (94-133 dB re 20 muPa) to generate a variety of equal sound exposure level conditions. Hearing thresholds were measured at the center frequency of the noise (2500 Hz) before, immediately after, and 24 h following exposure. Threshold shifts generated from 192 exposures ranged up to 30 dB. Estimates of TTS onset [159 dB re (20 muPa)(2) s] and growth (2.5 dB of TTS per dB of noise increase) were determined using an exponential function. Recovery for threshold shifts greater than 20 dB followed an 8.8 dB per log(min) linear function. Repeated testing indicated possible permanent threshold shift at the test frequency, but a later audiogram revealed no shift at this frequency or higher. Sea lions appear to be equally susceptible to noise in air and in water, provided that the noise exposure levels are referenced to absolute sound detection thresholds in both media. These data provide a framework within which to consider effects arising from more intense and/or sustained exposures.

Vocal performance affects metabolic rate in dolphins: implications for animals communicating in noisy environments

ABSTRACT Many animals produce louder, longer or more repetitious vocalizations to compensate for increases in environmental noise. Biological costs of increased vocal effort in response to noise, including energetic costs, remain empirically undefined in many taxa, particularly in marine mammals that rely on sound for fundamental biological functions in increasingly noisy habitats. For this investigation, we tested the hypothesis that an increase in vocal effort would result in an energetic cost to the signaler by experimentally measuring oxygen consumption during rest and a 2 min vocal period in dolphins that were trained to vary vocal loudness across trials. Vocal effort was quantified as the total acoustic energy of sounds produced. Metabolic rates during the vocal period were, on average, 1.2 and 1.5 times resting metabolic rate (RMR) in dolphin A and B, respectively. As vocal effort increased, we found that there was a significant increase in metabolic rate over RMR during the 2 min following sound production in both dolphins, and in total oxygen consumption (metabolic cost of sound production plus recovery costs) in the dolphin that showed a wider range of vocal effort across trials. Increases in vocal effort, as a consequence of increases in vocal amplitude, repetition rate and/or duration, are consistent with behavioral responses to noise in free-ranging animals. Here, we empirically demonstrate for the first time in a marine mammal, that these vocal modifications can have an energetic impact at the individual level and, importantly, these data provide a mechanistic foundation for evaluating biological consequences of vocal modification in noise-polluted habitats. Summary: Increased vocal effort by bottlenose dolphins has metabolic costs at the individual level, indicating energetic consequences for wild populations of marine mammals communicating in noise-polluted environments.

Spatial release from masking of aerial tones in pinnipeds

In most masking experiments, target signals and sound intended to mask are located in the same position. Spatial release from masking (SRM) occurs when signals and maskers are spatially separated, resulting in detection improvement relative to when they are spatially co-located. In this study, SRM was investigated in a harbor seal, who naturally lacks pinnae, and California sea lion, who possesses reduced pinnae. Subjects had to detect aerial tones at 1, 8, and 16 kHz in the presence of octave bands of white noise centered at the tone frequency. While the masker occurred in front of the subject (0 degree), the tone occurred at 0, 45, or 90 degrees in the horizontal plane. Unmasked thresholds were also measured at these angles to determine sensitivity differences based on source azimuth. Compared to when signal and masker where co-located, masked thresholds were lower by as much as 19 and 12 dB in the harbor seal and sea lion, respectively, when signal and masker were separated. Masked threshold differences of the harbor seal were larger than those previously measured under water. Performance was consistent with some measurements collected on terrestrial animals but differences between subjects at the highest frequency likely reflect variations in pinna anatomy.

The metabolic cost of communicative sound production in bottlenose dolphins (Tursiops truncatus)

SUMMARY Bottlenose dolphins (Tursiops truncatus) produce various communicative sounds that are important for social behavior, maintaining group cohesion and coordinating foraging. For example, whistle production increases during disturbances, such as separations of mother–calf pairs and vessel approaches. It is clear that acoustic communication is important to the survival of these marine mammals, yet the metabolic cost of producing whistles and other socials sounds and the energetic consequences of modifying these sounds in response to both natural and anthropogenic disturbance are unknown. We used flow-through respirometry to determine whether the metabolic cost of sound production could be quantified in two captive dolphins producing social sounds (whistles and squawks). On average, we found that metabolic rates measured during 2 min periods of sound production were 1.2 times resting values. Up to 7 min were required for metabolism to return to resting values following vocal periods. The total metabolic cost (over resting values) of the 2 min vocal period plus the required recovery period (163.3 to 2995.9 ml O2 or 3279.6 to 60,166.7 J) varied by individual as well as by mean duration of sounds produced within the vocal period. Observed variation in received cumulative sound energy levels of vocalizations was not related to total metabolic costs. Furthermore, our empirical findings did not agree with previous theoretical estimates of the metabolic cost of whistles. This study provides the first empirical data on the metabolic cost of sound production in dolphins, which can be used to estimate metabolic costs of vocal responses to environmental perturbations in wild dolphins.

The Relationship between Vessel Traffic and Noise Levels Received by Killer Whales (Orcinus orca)

Whale watching has become increasingly popular as an ecotourism activity around the globe and is beneficial for environmental education and local economies. Southern Resident killer whales (Orcinus orca) comprise an endangered population that is frequently observed by a large whale watching fleet in the inland waters of Washington state and British Columbia. One of the factors identified as a risk to recovery for the population is the effect of vessels and associated noise. An examination of the effects of vessels and associated noise on whale behavior utilized novel equipment to address limitations of previous studies. Digital acoustic recording tags (DTAGs) measured the noise levels the tagged whales received while laser positioning systems allowed collection of geo-referenced data for tagged whales and all vessels within 1000 m of the tagged whale. The objective of the current study was to compare vessel data and DTAG recordings to relate vessel traffic to the ambient noise received by tagged whales. Two analyses were conducted, one including all recording intervals, and one that excluded intervals when only the research vessel was present. For all data, significant predictors of noise levels were length (inverse relationship), number of propellers, and vessel speed, but only 15% of the variation in noise was explained by this model. When research-vessel-only intervals were excluded, vessel speed was the only significant predictor of noise levels, and explained 42% of the variation. Simple linear regressions (ignoring covariates) found that average vessel speed and number of propellers were the only significant correlates with noise levels. We conclude that vessel speed is the most important predictor of noise levels received by whales in this study. Thus, measures that reduce vessel speed in the vicinity of killer whales would reduce noise exposure in this population.

Localization of aerial pure tones by pinnipeds

In this study, minimum audible angles (MAAs) of aerial pure tones were measured in and compared between a northern elephant seal (Mirounga angustirostris), a harbor seal (Phoca vitulina), and a California sea lion (Zalophus californianus). Testing was conducted between 0.8 and 16 kHz in the elephant seal and 0.8 and 20 kHz in the harbor seal and sea lion in a hemi-anechoic chamber using a left/right psychophysical procedure. Performance for the same frequencies was also quantified for discrete speaker separation of 5 degrees from the mid-line. For all subjects, MAAs ranged from approximately 3 degrees to 15 degrees and were generally equal to or larger than those previously measured in the same subjects with a broadband signal. Performance at 5 degrees ranged from chance to 97% correct, depending on frequency and subject. Poorest performance in the sea lion and harbor seal occurred at intermediate frequencies, which is consistent with the duplex theory of sound localization. In contrast, the elephant seals poorest performance occurred at higher frequencies. The elephant seals result suggests an inferior ability to utilize interaural level differences and is perhaps related to best hearing sensitivity shifted toward lower frequencies in this species relative to other pinnipeds.