High Source Levels and Small Active Space of High-Pitched Song in Bowhead Whales (Balaena mysticetus)

The low-frequency, powerful vocalizations of blue and fin whales may potentially be detected by conspecifics across entire ocean basins. In contrast, humpback and bowhead whales produce equally powerful, but more complex broadband vocalizations composed of higher frequencies that suffer from higher attenuation. Here we evaluate the active space of high frequency song notes of bowhead whales (Balaena mysticetus) in Western Greenland using measurements of song source levels and ambient noise. Four independent, GPS-synchronized hydrophones were deployed through holes in the ice to localize vocalizing bowhead whales, estimate source levels and measure ambient noise. The song had a mean apparent source level of 185±2 dB rms re 1 µPa @ 1 m and a high mean centroid frequency of 444±48 Hz. Using measured ambient noise levels in the area and Arctic sound spreading models, the estimated active space of these song notes is between 40 and 130 km, an order of magnitude smaller than the estimated active space of low frequency blue and fin whale songs produced at similar source levels and for similar noise conditions. We propose that bowhead whales spatially compensate for their smaller communication range through mating aggregations that co-evolved with broadband song to form a complex and dynamic acoustically mediated sexual display.     

Non-song acoustic communication in migrating humpback whales (Megaptera novaeangliae)

Humpback whales are renowned for the complex structured songs produced by males. A second, relatively understudied area of humpback acoustic communication concerns un-patterned sounds known as "social sounds," produced by both males and females. These include vocalizations as well as sounds produced at the surface of the water as a result of surface behaviors ( e.g. , breaching, pectoral slapping). This study describes a portion of the non-song social sound repertoire of southward migrating humpbacks in Australian waters, and explores the social relevance of these sounds. On migration, humpback whales travel in social groups of varying compositions. These social groups are not stable in that humpback whales continually change group composition by splitting from, or joining with, other groups. The results of this study suggest that "breaching" and "slapping" have a communicative function. Other sounds such as "underwater blows" and "cries" were heard mainly in competitive groups while other low-frequency sounds such as "grumbles,""snorts,""thwops," and "wops" may function in intra- or inter-group communication. Particular sounds ("grunts,""groans," and "barks") were almost exclusive to joining pods suggesting a role in social integration. Social sounds in humpbacks may have specific social and behavioral functions relating to social group composition, and the mediation of interactions between these social groups.     

Your attention please: increasing ambient noise levels elicits a change in communication behaviour in humpback whales (Megaptera novaeangliae)

High background noise is an important obstacle in successful signal detection and perception of an intended acoustic signal. To overcome this problem, many animals modify their acoustic signal by increasing the repetition rate, duration, amplitude or frequency range of the signal. An alternative method to ensure successful signal reception, yet to be tested in animals, involves the use of two different types of signal, where one signal type may enhance the other in periods of high background noise. Humpback whale communication signals comprise two different types: vocal signals, and surface-generated signals such as ‘breaching’ or ‘pectoral slapping’. We found that humpback whales gradually switched from primarily vocal to primarily surface-generated communication in increasing wind speeds and background noise levels, though kept both signal types in their repertoire. Vocal signals have the advantage of having higher information content but may have the disadvantage of loosing this information in a noisy environment. Surface-generated sounds have energy distributed over a greater frequency range and may be less likely to become confused in periods of high wind-generated noise but have less information content when compared with vocal sounds. Therefore, surface-generated sounds may improve detection or enhance the perception of vocal signals in a noisy environment.     

Recurring patterns in the songs of humpback whales (Megaptera novaeangliae)

Humpback whales, unlike most mammalian species, learn new songs as adults. Populations of singers progressively and collectively change the sounds and patterns within their songs throughout their lives and across generations. In this study, humpback whale songs recorded in Hawaii from 1985 to 1995 were analyzed using self-organizing maps (SOMs) to classify the sounds within songs, and to identify sound patterns that were present across multiple years. These analyses supported the hypothesis that recurring, persistent patterns exist within whale songs, and that these patterns are defined at least in part by acoustic relationships between adjacent sounds within songs. Sound classification based on acoustic differences between adjacent sounds yielded patterns within songs that were more consistent from year to year than classifications based on the properties of single sounds. Maintenance of fixed ratios of acoustic modulation across sounds, despite large variations in individual sounds, suggests intrinsic constraints on how sounds change within songs. Such acoustically invariant cues may enable whales to recognize and assess variations in songs despite propagation-related distortion of individual sounds and yearly changes in songs.     

SEASONAL AND DIURNAL TRENDS OF CHORUSING HUMPBACK WHALES WINTERING IN WATERS OFF WESTERN MAUI

A portable data logger controlled by a Tattletale 7 microcontroller was used to record humpback whale choruses during the 1998 humpback whale winter season in Hawaii. The data logger sampled the sounds for four minutes every half hour using a digitizing rate of 2 kHz, and the data were stored on a hard disk. The results between January and April showed a peak in the sound pressure level between mid-February and mid-March. This peak of approximately 120 dB re 1 μPa coincided with the peak in the number of whales sighted by aerial survey on 7 March 1998. The choruses had spectral peaks at 315 Hz and 630 Hz. Some of the sounds at 630 Hz were second harmonics of the 315 Hz peak and others were not. The data also indicated a diurnal pattern in the sound pressure level, with levels at night significantly louder than the daytime levels. The sound levels began to increase during sunset and remained relatively high until sunrise, when they progressively decreased to a minimum. The nighttime peak occurred within an hour before and after midnight, and the daytime minimum occurred between 1100 and 1500. That more humpback whales appear to sing at night may reflect a switch to sexual advertisement as the primary male mating strategy at this time. It may also indicate that daylight and vision play key roles in the formation of competitive groups. It is suggested that the relative number of humpback whales in a given locale may be estimated by monitoring changes in sound pressure levels.     

Blue whales respond to anthropogenic noise

Anthropogenic noise may significantly impact exposed marine mammals. This work studied the vocalization response of endangered blue whales to anthropogenic noise sources in the mid-frequency range using passive acoustic monitoring in the Southern California Bight. Blue whales were less likely to produce calls when mid-frequency active sonar was present. This reduction was more pronounced when the sonar source was closer to the animal, at higher sound levels. The animals were equally likely to stop calling at any time of day, showing no diel pattern in their sensitivity to sonar. Conversely, the likelihood of whales emitting calls increased when ship sounds were nearby. Whales did not show a differential response to ship noise as a function of the time of the day either. These results demonstrate that anthropogenic noise, even at frequencies well above the blue whales' sound production range, has a strong probability of eliciting changes in vocal behavior. The long-term implications of disruption in call production to blue whale foraging and other behaviors are currently not well understood.     

Diversity in sound pressure levels and estimated active space of resident killer whale vocalizations

Signal source intensity and detection range, which integrates source intensity with propagation loss, background noise and receiver hearing abilities, are important characteristics of communication signals. Apparent source levels were calculated for 819 pulsed calls and 24 whistles produced by free-ranging resident killer whales by triangulating the angles-of-arrival of sounds on two beamforming arrays towed in series. Levels in the 1–20 kHz band ranged from 131 to 168 dB re 1 μPa at 1 m, with differences in the means of different sound classes (whistles: 140.2±4.1 dB; variable calls: 146.6±6.6 dB; stereotyped calls: 152.6±5.9 dB), and among stereotyped call types. Repertoire diversity carried through to estimates of active space, with “long-range” stereotyped calls all containing overlapping, independently-modulated high-frequency components (mean estimated active space of 10–16 km in sea state zero) and “short-range” sounds (5–9 km) included all stereotyped calls without a high-frequency component, whistles, and variable calls. Short-range sounds are reported to be more common during social and resting behaviors, while long-range stereotyped calls predominate in dispersed travel and foraging behaviors. These results suggest that variability in sound pressure levels may reflect diverse social and ecological functions of the acoustic repertoire of killer whales.     

Beaked whales respond to simulated and actual navy sonar

Beaked whales have mass stranded during some naval sonar exercises, but the cause is unknown. They are difficult to sight but can reliably be detected by listening for echolocation clicks produced during deep foraging dives. Listening for these clicks, we documented Blainville's beaked whales, Mesoplodon densirostris, in a naval underwater range where sonars are in regular use near Andros Island, Bahamas. An array of bottom-mounted hydrophones can detect beaked whales when they click anywhere within the range. We used two complementary methods to investigate behavioral responses of beaked whales to sonar: an opportunistic approach that monitored whale responses to multi-day naval exercises involving tactical mid-frequency sonars, and an experimental approach using playbacks of simulated sonar and control sounds to whales tagged with a device that records sound, movement, and orientation. Here we show that in both exposure conditions beaked whales stopped echolocating during deep foraging dives and moved away. During actual sonar exercises, beaked whales were primarily detected near the periphery of the range, on average 16 km away from the sonar transmissions. Once the exercise stopped, beaked whales gradually filled in the center of the range over 2-3 days. A satellite tagged whale moved outside the range during an exercise, returning over 2-3 days post-exercise. The experimental approach used tags to measure acoustic exposure and behavioral reactions of beaked whales to one controlled exposure each of simulated military sonar, killer whale calls, and band-limited noise. The beaked whales reacted to these three sound playbacks at sound pressure levels below 142 dB re 1 μPa by stopping echolocation followed by unusually long and slow ascents from their foraging dives. The combined results indicate similar disruption of foraging behavior and avoidance by beaked whales in the two different contexts, at exposures well below those used by regulators to define disturbance.     

Effect of harmonicity on the detection of a signal in a complex masker and on spatial release from masking

The amount of masking of sounds from one source (signals) by sounds from a competing source (maskers) heavily depends on the sound characteristics of the masker and the signal and on their relative spatial location. Numerous studies investigated the ability to detect a signal in a speech or a noise masker or the effect of spatial separation of signal and masker on the amount of masking, but there is a lack of studies investigating the combined effects of many cues on the masking as is typical for natural listening situations. The current study using free-field listening systematically evaluates the combined effects of harmonicity and inharmonicity cues in multi-tone maskers and cues resulting from spatial separation of target signal and masker on the detection of a pure tone in a multi-tone or a noise masker. A linear binaural processing model was implemented to predict the masked thresholds in order to estimate whether the observed thresholds can be accounted for by energetic masking in the auditory periphery or whether other effects are involved. Thresholds were determined for combinations of two target frequencies (1 and 8 kHz), two spatial configurations (masker and target either co-located or spatially separated by 90 degrees azimuth), and five different masker types (four complex multi-tone stimuli, one noise masker). A spatial separation of target and masker resulted in a release from masking for all masker types. The amount of masking significantly depended on the masker type and frequency range. The various harmonic and inharmonic relations between target and masker or between components of the masker resulted in a complex pattern of increased or decreased masked thresholds in comparison to the predicted energetic masking. The results indicate that harmonicity cues affect the detectability of a tonal target in a complex masker.     

200 kHz Commercial Sonar Systems Generate Lower Frequency Side Lobes Audible to Some Marine Mammals

The spectral properties of pulses transmitted by three commercially available 200 kHz echo sounders were measured to assess the possibility that marine mammals might hear sound energy below the center (carrier) frequency that may be generated by transmitting short rectangular pulses. All three sounders were found to generate sound at frequencies below the center frequency and within the hearing range of some marine mammals, e.g. killer whales, false killer whales, beluga whales, Atlantic bottlenose dolphins, harbor porpoises, and others. The frequencies of these sub-harmonic sounds ranged from 90 to 130 kHz. These sounds were likely detectable by the animals over distances up to several hundred meters but were well below potentially harmful levels. The sounds generated by the sounders could potentially affect the behavior of marine mammals within fairly close proximity to the sources and therefore the exclusion of echo sounders from environmental impact analysis based solely on the center frequency output in relation to the range of marine mammal hearing should be reconsidered.     

A Dominance Hierarchy of Auditory Spatial Cues in Barn Owls

Background

Barn owls integrate spatial information across frequency channels to localize sounds in space.

Methodology/Principal Findings

We presented barn owls with synchronous sounds that contained different bands of frequencies (3–5 kHz and 7–9 kHz) from different locations in space. When the owls were confronted with the conflicting localization cues from two synchronous sounds of equal level, their orienting responses were dominated by one of the sounds: they oriented toward the location of the low frequency sound when the sources were separated in azimuth; in contrast, they oriented toward the location of the high frequency sound when the sources were separated in elevation. We identified neural correlates of this behavioral effect in the optic tectum (OT, superior colliculus in mammals), which contains a map of auditory space and is involved in generating orienting movements to sounds. We found that low frequency cues dominate the representation of sound azimuth in the OT space map, whereas high frequency cues dominate the representation of sound elevation.

Conclusions/Significance

We argue that the dominance hierarchy of localization cues reflects several factors: 1) the relative amplitude of the sound providing the cue, 2) the resolution with which the auditory system measures the value of a cue, and 3) the spatial ambiguity in interpreting the cue. These same factors may contribute to the relative weighting of sound localization cues in other species, including humans.     

North Atlantic right whales (Eubalaena glacialis) ignore ships but respond to alerting stimuli

North Atlantic right whales were extensively hunted during the whaling era and have not recovered. One of the primary factors inhibiting their recovery is anthropogenic mortality caused by ship strikes. To assess risk factors involved in ship strikes, we used a multi-sensor acoustic recording tag to measure the responses of whales to passing ships and experimentally tested their responses to controlled sound exposures, which included recordings of ship noise, the social sounds of conspecifics and a signal designed to alert the whales. The whales reacted strongly to the alert signal, they reacted mildly to the social sounds of conspecifics, but they showed no such responses to the sounds of approaching vessels as well as actual vessels. Whales responded to the alert by swimming strongly to the surface, a response likely to increase rather than decrease the risk of collision.     

Selective Attention Modulates Human Auditory Brainstem Responses: Relative Contributions of Frequency and Spatial Cues

Selective attention is the mechanism that allows focusing one’s attention on a particular stimulus while filtering out a range of other stimuli, for instance, on a single conversation in a noisy room. Attending to one sound source rather than another changes activity in the human auditory cortex, but it is unclear whether attention to different acoustic features, such as voice pitch and speaker location, modulates subcortical activity. Studies using a dichotic listening paradigm indicated that auditory brainstem processing may be modulated by the direction of attention. We investigated whether endogenous selective attention to one of two speech signals affects amplitude and phase locking in auditory brainstem responses when the signals were either discriminable by frequency content alone, or by frequency content and spatial location. Frequency-following responses to the speech sounds were significantly modulated in both conditions. The modulation was specific to the task-relevant frequency band. The effect was stronger when both frequency and spatial information were available. Patterns of response were variable between participants, and were correlated with psychophysical discriminability of the stimuli, suggesting that the modulation was biologically relevant. Our results demonstrate that auditory brainstem responses are susceptible to efferent modulation related to behavioral goals. Furthermore they suggest that mechanisms of selective attention actively shape activity at early subcortical processing stages according to task relevance and based on frequency and spatial cues.     

REACTION OF FIN WHALES BALAENOPTERA PHYSALUS TO AN EARTHQUAKE

ABSTRACT

Whales living within seismically active regions are subject to intense disturbances from strong sounds produced by earthquakes that can kill or injure individuals. Nishimura & Clark (1993) relate the possible effects of underwater earthquake noise levels in marine mammals, adducing that T-phase source signal level (10- to 30- Hz range) can exceed 200 dB re: 1 μPa at 1 m, for a magnitude 4–5 earthquake, sounds audible to fin whales which produce low frequency sounds of 16–20/25–44 Hz over 0.5–1s, typically of 183 dB re: 1 μPa at 1 m. Here we present the response of a fin whale to a 5.5 Richter scale earthquake that took place on 22 February 2005, in the Gulf of California. The whale covered 13 km in 26 min (mean speed = 30.2 km/h). We deduce that the sound heard by this whale might have triggered the costly energy expenditure of high speed swimming as a seismic-escape response. These observations support the hypothesis of Richardson et al. (1995) that cetaceans may flee from loud sounds before they are injured, when exposed to noise in excess of 140 dB re: 1 μPa 1 m.     

ATOC AND OTHER FACTORS AFFECTING THE DISTRIBUTION AND ABUNDANCE OF HUMPBACK WHALES (MEGAPTERA NOVAEANGLIAE) OFF THE NORTH SHORE OF KAUAI

Humpback whale diving behavior changes subtly when exposed to signals transmitted from the Acoustic Thermometry of Ocean Climate (ATOC) sound projector located 14 km offshore the island of Kauai. This study considered whether such responses would lead to changes in distribution and abundance. A land-based shore station measured humpback whale locations (scan samples) for both nearshore (<5 km) and offshore (5–10 km) areas. Control observations were made in 1994 and 1998. In 1998 multipleday blocks with ATOC transmissions were interspersed with multiple-day control blocks without transmissions. Sighting rates were higher in 1998 (ATOC) than in 1994 (control year), probably due to better sighting conditions, but may reflect increased population size. Sighting rates did not differ between control and ATOC conditions in 1998. A seasonal sighting peak was observed in both years. No vessel effect on sighting rate was detected in 1998.
There was no effect of ATOC on the distance from the shore station to whales, or the depth of water where pods were located. However, the distribution of whales shifted slightly eastward during the ATOC blocks and the mean distance between the ATOC source and pods was greater during transmissions. Nonetheless, more whales were found close to the source when it was on, suggesting a more variable response rather than simple avoidance, with whales found both closer to, and farther away from, the source during transmissions.     

Low-Frequency Sounds Repel Male Mosquitoes <Emphasis Type="Italic">Aedes diantaeus</Emphasis> N.D.K. (Diptera,Culicidae)

We experimentally demonstrated that tonal acoustic signals with a carrier frequency of 140–200 Hz had a repellent effect on male mosquitoes (Culicidae). Swarming males of Aedes diantaeus were concentrated in a small space near the auxiliary attracting sound source which simulated the flight sound of conspecific females (carrier frequency 280–320 Hz). Then, the resulting cluster of attracted mosquitoes was stimulated with test signals of variable amplitude and carrier frequency from a second loudspeaker. The direction of mosquito flight from the source of test sounds and a decrease in their number above the attracting sound source were used as the criteria of behavioral response. Pronounced avoidance responses (negative phonotaxis) of swarming mosquitoes were observed in the range of 140–200 Hz. Most of the mosquitoes left the area above the attracting sound source within one second after the onset of the test signal. Mosquitoes mostly flew up, sideways, and backwards in relation to the test acoustic vector. We presume that mosquitoes develop defensive behavior against attacking predatory insects based on analysis of auditory information. The range of negative phonotaxis is limited at higher frequencies by the spectrum of the flight sounds of conspecific females, and in the low frequency range, by the increasing level of atmospheric noise.     

FISHING GEAR INVOLVED IN ENTANGLEMENTS OF RIGHT AND HUMPBACK WHALES

Interactions between marine mammals and fishing gear are an issue of global concern. Entanglements in the western North Atlantic are a major source of injury and mortality for endangered large whales. In this study, entanglements of 31 right whales ( Eubalaena glacialis ) and 30 humpback whales ( Megaptera novaeangliae ) were analyzed to determine the types and parts of gear involved. When gear was identified, 89% ( n = 32) of the entanglements were attributed to pot and gill net gear; however, a wide range of specific gear types were implicated. Despite gear recovery, gear type was not identified in 20% ( n = 9) of the cases. Although pot gear was recovered from both species equally, gill net gear was less frequently retrieved from right whales ( n = 2) than humpback whales ( n = 11). When gear part was identified, 81% ( n = 21) involved entanglements in buoy line and/or groundline. For right whales, the most common point of gear attachment was the mouth (77.4%); for humpback whales, the tail (53%) and the mouth (43%) were common attachment sites. Four right and three humpback whales in this sample were known to have died subsequent to entanglement. However, when identified, the gear types and parts involved in lethal cases were not substantially different from entanglements with non-lethal outcomes. Large whales can become entangled in a wide variety of fishing gear types and parts, and additional insight will depend on continued efforts to document entanglements and recover associated gear.     

The hearing of the Atlantic Salmon, Salmo salar

The hearing of the salmon, Salmo salar L., was studied by means of a cardiac conditioning technique. Fish were trained to show a slowing of the heart, on hearing a sound, in anticipation of a mild electric shock applied later. The minimum sound level to which the fish would respond was determined for a range of pure tones, both in the sea, and in the laboratory. The fish responded only to low frequency tones (below 380 Hz), and particle motion, rather than sound pressure, proved to be the relevant stimulus. The sensitivity of the fish to sound was not affected by the level of sea noise under natural conditions but hearing is likely to be masked by ambient noise in a turbulent river. Sound measurements made in the River Dee, near Aberdeen, lead to the conclusion that salmon are unlikely to detect sounds originating in air, but that they are sensitive to substrate borne sounds. Compared with the carp and cod the hearing of the salmon is poor, and more like that of the perch and plaice.     

Acoustic communication in two freshwater gobies: the relationship between ambient noise,hearing thresholds and sound spectrum

Two freshwater gobies Padogobius martensii and Gobius nigricans live in shallow (5-70 cm) stony streams, and males of both species produce courtship sounds. A previous study demonstrated high noise levels near waterfalls, a quiet window in the noise around 100 Hz at noisy locations, and extremely short-range propagation of noise and goby signals. To investigate the relationship of this acoustic environment to communication, we determined audiograms for both species and measured parameters of courtship sounds produced in the streams. We also deflated the swimbladder in P. martensii to determine its effect on frequency utilization in sound production and hearing. Both species are maximally sensitive at 100 Hz and produce low-frequency sounds with main energy from 70 to 100-150 Hz. Swimbladder deflation does not affect auditory threshold or dominant frequency of courtship sounds and has no or minor effects on sound amplitude. Therefore, both species utilize frequencies for hearing and sound production that fall within the low-frequency quiet region, and the equivalent relationship between auditory sensitivity and maximum ambient noise levels in both species further suggests that ambient noise shapes hearing sensitivity.