Effects of Airgun Sounds on Bowhead Whale Calling Rates: Evidence for Two Behavioral Thresholds

In proximity to seismic operations, bowhead whales (Balaena mysticetus) decrease their calling rates. Here, we investigate the transition from normal calling behavior to decreased calling and identify two threshold levels of received sound from airgun pulses at which calling behavior changes. Data were collected in August–October 2007–2010, during the westward autumn migration in the Alaskan Beaufort Sea. Up to 40 directional acoustic recorders (DASARs) were deployed at five sites offshore of the Alaskan North Slope. Using triangulation, whale calls localized within 2 km of each DASAR were identified and tallied every 10 minutes each season, so that the detected call rate could be interpreted as the actual call production rate. Moreover, airgun pulses were identified on each DASAR, analyzed, and a cumulative sound exposure level was computed for each 10-min period each season (CSEL_10-min). A Poisson regression model was used to examine the relationship between the received CSEL_10-min from airguns and the number of detected bowhead calls. Calling rates increased as soon as airgun pulses were detectable, compared to calling rates in the absence of airgun pulses. After the initial increase, calling rates leveled off at a received CSEL_10-min of ~94 dB re 1 μPa²-s (the lower threshold). In contrast, once CSEL_10-min exceeded ~127 dB re 1 μPa²-s (the upper threshold), whale calling rates began decreasing, and when CSEL_10-min values were above ~160 dB re 1 μPa²-s, the whales were virtually silent.     

SEAL RESPONSES TO AIRGUN SOUNDS DURING SUMMER SEISMIC SURVEYS IN THE ALASKAN BEAUFORT SEA

Numbers, sighting distances, and behavior of seals were studied during a nearshore seismic program off northern Alaska in July-September 1996. We observed from the seismic vessel for 885.6 h, including all periods (day and night) when airguns operated and many periods without airguns. Of 422 seals seen, 421 were seen in daylight; 91-8% were ringed seals, 7.3% were bearded seals, and 0.9% were spotted seals. About 79% were first seen within 250 m of the seismic boat, and sighting rate declined rapidly at lateral distances > 50 m. During daylight, seals were seen at nearly identical rates (0.60-0.63/ h) during periods with no airguns firing, one airgun, and a "full-array" of 8-11 120-in³ airguns. However, seals tended to be farther away ( P < 0.0001) during full-array seismic. There was partial avoidance of the zone <150 m from the boat during full-array seismic, but seals apparently did not move much beyond 250 m. "Swimming away" was more common during full-array than no-airgun periods, but relative frequencies of five behaviors did not differ significantly among distance categories. Airgun operations were interrupted 112 times when seals were sighted within safety radii (150–250 m). The National Marine Fisheries Service specified these radii in the Incidental Harassment Authorization issued for the project; they are based on a 190 dB re 1 μPa (rms) criterion for broadband received level. Methods for estimating numbers of seals potentially affected by the seismic program are described, and effectiveness of monitoring and mitigation is discussed. There is an urgent need for more data on effects of strong seismic pulses on seals.     

Effects of seismic surveys on New Zealand fur seals during daylight hours: Do fur seals respond to obstacles rather than airgun noise?

The responses of New Zealand fur seals (Arctocephalus forsteri) during a three‐dimensional marine seismic survey were recorded in the first quantified investigation of the effects of seismic exploration operations on otariid seals. The survey was over the continental slope off southern New Zealand, where fur seals dive to forage at night and rest on the surface during daylight hours. Data were restricted to daylight sightings from the source vessel while towed seismic gear was fully deployed, with comparisons made between fur seal responses when airguns were off and when airguns were operating at full power. Results were inconclusive. Comparisons were confounded because both sighting rate and distance first seen decreased with deteriorating sea state. The key finding of this study was that the source vessel and towed gear created physical obstacles that often generated responses from fur seals. A more thorough investigation is recommended in order to differentiate between responses to airgun noise and responses to physical obstacles, and improvements are suggested to rectify shortfalls in data collection that should generate conclusive outcomes.     

Short-term disturbance by a commercial two-dimensional seismic survey does not lead to long-term displacement of harbour porpoises

Assessments of the impact of offshore energy developments are constrained because it is not known whether fine-scale behavioural responses to noise lead to broader-scale displacement of protected small cetaceans. We used passive acoustic monitoring and digital aerial surveys to study changes in the occurrence of harbour porpoises across a 2000 km² study area during a commercial two-dimensional seismic survey in the North Sea. Acoustic and visual data provided evidence of group responses to airgun noise from the 470 cu inch array over ranges of 5–10 km, at received peak-to-peak sound pressure levels of 165–172 dB re 1 µPa and sound exposure levels (SELs) of 145–151 dB re 1 µPa² s⁻¹. However, animals were typically detected again at affected sites within a few hours, and the level of response declined through the 10 day survey. Overall, acoustic detections decreased significantly during the survey period in the impact area compared with a control area, but this effect was small in relation to natural variation. These results demonstrate that prolonged seismic survey noise did not lead to broader-scale displacement into suboptimal or higher-risk habitats, and suggest that impact assessments should focus on sublethal effects resulting from changes in foraging performance of animals within affected sites.     

Comparing marine mammal acoustic habitats in Atlantic and Pacific sectors of the High Arctic: year-long records from Fram Strait and the Chukchi Plateau

During the International Polar Year (IPY), acoustic recorders were deployed on oceanographic moorings in Fram Strait and on the Chukchi Plateau, representing the first coordinated year-round sampling of underwater acoustic habitats at two sites in the High Arctic. Examination of species-specific marine mammal calls recorded from autumn 2008–2009 revealed distinctly different acoustic habitats at each site. Overall, the Fram Strait site was acoustically complex compared with the Chukchi Plateau site. In Fram Strait, calls from bowhead whales (Balaena mysticetus) and a variety of toothed whales (odontocetes) were recorded year-round, as were airgun pulses from seismic surveys. In addition, calls from blue whales (Balaenoptera musculus) and fin whales (B. physalus) were recorded from June to October and August to March, respectively. Conversely, at the Chukchi Plateau site, beluga (Delphinapterus leucas) and bowhead whale calls were recorded primarily from May to August, with airgun signals detected only in September–October. Ribbon seal (Phoca fasciata) calls were detected in October–November, with no marine mammals calls at all recorded from December to February. Of note, ice-adapted bearded seals (Erignathus barbatus) were recorded at both sites, primarily in spring and summer, corresponding with the mating season for that species. Differences in acoustic habitats between the two sites were related to contrasts in sea ice cover, temperature, patterns of ocean circulation and contributions from anthropogenic noise sources. These data provide a provisional baseline for the comparison of underwater acoustic habitats between Pacific and Atlantic sectors of the High Arctic.     

Postural instability and akinesia in a pantropical spotted dolphin,Stenella attenuata,in proximity to operating airguns of a geophysical seismic vessel

Aberrant behaviour including erratic locomotion was observed in a pantropical spotted dolphin 600 m ahead of an airgun array during 3D seismic explorations off Liberia in March 2009. The dolphin, presumably in acoustic distress, lifted its head and cervical region above the surface in an oblique, strikingly rigid posture during 5 min. Turbulent white-water evidenced a major propulsory thrust. Incremental postural instability and apparent exhaustion progressed to a catatonic-like state of akinesia as the dolphin rolled over onto one side, then its back before sinking virtually motionless close to the airgun array. Unless it recovered full locomotory control, asphyxiation was inevitable. Potential internal injury is discussed, both acoustic-mediated and from extreme exertion (exertional myopathy, rhabdomyolysis and myoglobinuric nephrosis). As behaviour was spatially and temporally closely associated with firing seismic airguns, we suggest a cause–effect relationship. Differential diagnoses of pre-existing morbidity, senescence, or intoxication are considered possible but unlikely.     

Through-ice communication by Weddell seals may not be practicable

Possible communication between territorial male Weddell seals (Leptonychotes weddellii) under the ice with females on the ice was investigated. In-air and underwater recordings of underwater calls were made at three locations near Davis, Antarctica. Most underwater calls were not detectable in air, often because of wind noise. In-air call amplitudes of detectable calls ranged from 32–74 dB re. 20 Pa at 86 Hz down to 4–38 dB re. 20 Pa at 3.6 kHz. Most of these would be audible to humans. Only 26 of 582 amplitude measurements (from 230 calls) ranged from 5 dB to a maximum of 15 dB above the minimum harbour-seal (Phoca vitulina) in-air detection threshold. Seals on the ice could likely hear a few very loud underwater calls but only if the caller was nearby and there were no wind noises. The low detectability of underwater calls in air likely precludes effective communication between underwater seals and those on the ice.     

Are Tropical Highland Frog Calls Cold–adapted? The Case of the Andean Frog Hyla labialis1

Across a wide range of temperatures established in the laboratory, we tape–recorded the advertisement calls of 76 freshly caught Hyla labialis males from three elevationally separated populations in the Eastern Andes of Colombia. Each male was tested once at a single temperature and returned to his capture site after measurement of his snout–vent length. We measured and averaged three characteristics of five to ten successive calls for each individual: number of pulses per call, pulse repetition rate, and call duration. We found that calling activity occurred within temperature ranges that overlapped among frogs from different elevations, but widened and shifted downward with increasing altitude of origin. Males from all sites called at temperatures higher, but not lower, than those naturally occurring during their nightly activity period. No decline in vocal performance was apparent when frogs extended their calling activity into the range of high temperatures selected for basking. Both snout–vent length and temperature affected pulse repetition rate and call duration, while the number of pulses per call was temperature–independent. Compared to the smaller males from lower elevations, the larger, high–mountain males had calls with significandy more pulses, a lower pulse repetition rate, and longer duration. Within each population, rising temperatures caused pulse repetition rate to increase and call duration to decrease significantly, whereas the number of pulses per call remained unchanged. Pulse repetition rate of highland males was the factor least affected by temperature, and it was less sensitive to night temperatures than to day temperatures. This, together with their capacity to call at low temperatures, suggests that highland frog calls are cold adapted.     

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.     

How long do females really listen? Assessment time for female mate choice in the grey treefrog, Hyla versicolor

A satisfactory understanding of the process of mate choice in many species of anurans requires that we know how sensitive females are to the variation in male calls under natural conditions and what is the timescale or ‘window’ over which females compare potential mates. In natural choruses, grey treefrog females may sit near calling males for many minutes before approaching a particular individual to mate, whereas in laboratory-based phonotaxis tests, they may approach a speaker within 30 s of exposure to broadcast calls. Females prefer long versus short calls. To estimate ‘assessment time’ of females in nature, we broadcast calls from four pairs of 360-degree speakers suspended within screen cages at four locations at the edge of a pond. One speaker per pair presented calls of constant duration while the other speaker shifted between broadcasts of calls that were longer or shorter than the constant duration call. The time period over which this change in call duration occurred differed between the four venues. Laboratory-based choice tests indicated that females preferred call sources with variable numbers of pulses to those with constant numbers of pulses when the former had more total pulses per time window. Accordingly, we assigned the probabilities of field captures at the different speakers based on the summed pulses from the constant and cycling speakers within the possible assessment windows. These probabilities, together with the numbers of females captured at the speaker array over the breeding season, indicated that the most likely assessment time is close to 2 min.     

Correspondence between evoked vocal responses and auditory thresholds in Pleurodema thaul (Amphibia; Leptodactylidae)

Thresholds for evoked vocal responses and thresholds of multiunit midbrain auditory responses to pure tones and synthetic calls were investigated in males of Pleurodema thaul, as behavioral thresholds well above auditory sensitivity have been reported for other anurans. Thresholds for evoked vocal responses to synthetic advertisement calls played back at increasing intensity averaged 43 dB RMS SPL (range 31–52 dB RMS SPL), measured at the subjects’ position. Number of pulses increased with stimulus intensities, reaching a plateau at about 18–39 dB above threshold and decreased at higher intensities. Latency to call followed inverse trends relative to number of pulses. Neural audiograms yielded an average best threshold in the high frequency range of 46.6 dB RMS SPL (range 41–51 dB RMS SPL) and a center frequency of 1.9 kHz (range 1.7–2.6 kHz). Auditory thresholds for a synthetic call having a carrier frequency of 2.1 kHz averaged 44 dB RMS SPL (range 39–47 dB RMS SPL). The similarity between thresholds for advertisement calling and auditory thresholds for the advertisement call indicates that male P. thaul use the full extent of their auditory sensitivity in acoustic interactions, likely an evolutionary adaptation allowing chorusing activity in low-density aggregations.     

WEDDELL SEAL UNDERWATER CALLING RATES DURING THE WINTER AND SPRING NEAR MAWSON STATION, ANTARCTICA

Underwater vocalization monitoring and surveys, both on ice and underwater, were used to determine if Weddell seals ( Leptonychotes weddellii ) near Mawson Station, Antarctica, remain under the fast ice during winter within close range of breeding sites. Daytime and nighttime underwater calling rates were examined at seven breeding sites during austral winter and spring to identify seasonal and diel patterns. Seals rarely hauled out at any of the sites during winter, although all cohorts (adult males, females, and juveniles) were observed underwater and surfacing at breathing holes throughout winter (June–September) and spring (October–December). Seal vocalizations were recorded during each sampling session throughout the study ( n = 102 daytime at seven sites collectively, and n = 5 24-h samples at each of two sites). Mean daytime calling rate was low in mid-winter (July) (mean = 18.9 ± 7.1 calls/min) but increased monthly, reaching a peak during the breeding season (November) (mean = 62.6 ± 15.7 calls/min). Mean nighttime calling rate was high throughout the winter and early spring (July–October) with mean nocturnal calling rate in July (mean = 61.8 ± 35.1 calls/min) nearly equal to mean daytime calling rate in November (during 24-h daylight). Reduced vocal behavior during winter daylight periods may result from animals utilizing the limited daylight hours for nonvocal activities, possibly feeding.     

North Pacific right whale (Eubalaena japonica) seasonal and diel calling patterns from long‐term acoustic recordings in the southeastern Bering Sea, 2000–2006

We assessed North Pacific right whale (Eubalaena japonica) seasonal and daily calling patterns in the southeastern Bering Sea (SEBS) using long‐term hydrophone recordings from October 2000 through January 2006. We detected right whale calls on the SEBS middle shelf (<100 m depth) as early as May, intermittently throughout summer and fall, and as late as December. Calls also were detected on one day in June 2005 on the SEBS slope (>1,000 m), but were not detected near Kodiak Island from April to August 2003. In months with calls, detections occurred on more days in July–October (≥6 d/mo), than from May to June or November to December (≤3 d/mo). Calls were clustered in time and were usually detected on 1–3 consecutive days with a median interval of 6.5 d for calls >1 d apart. Hourly calling rates were significantly higher at night than during the day. These data indicate that right whales occur in the SEBS later in the year than previously known, intermittently pass through the middle‐shelf study region, and usually remain there no longer than a few days. Right whale habitat use in the SEBS may intensify in mid‐summer through early fall based on higher monthly and daily call detection rates.     

Patterns of Vocal Interactions in a Bullfrog (Rana catesbeiana) Chorus: Preferential Responding to Far Neighbors

In chorusing species, males seem to be spaced non-randomly, and their vocal interactions may be governed by particular behavioral rules. We monitored patterns of vocal interactions in a natural bullfrog ( Rana catesbeiana ) chorus to determine the probability with which calls of individual frogs would follow each other's in dyadic sequences. Expected probabilities of responses in a dyad were calculated based upon the joint probabilities of calling (relative calling rates) of the individual frogs; observed probabilities of response reflected the actual number of following responses in each dyad. Results of statistical tests comparing observed and expected probabilities of responding revealed that, when dyads were closely spaced, observed probabilities of a following response were significantly less than the expected probabilities. Conversely, when dyads were composed of more distant males, observed probabilities of responding were significantly greater than expected. Observed probabilities of response were correlated with inter-male distances; males called more frequently than expected following calls of far neighbors, and less frequently than expected following calls of near neighbors. These data suggest that males attend to both nearby and distant callers, and adjust the onset of their own vocalizations appropriately. Males may be actively inhibited by calls of their near neighbors, and their calling may be actively elicited by the calls of their far neighbors.     

Variability in blue whale acoustic behavior off southern California

To evaluate the acoustic behavior of blue whales (Balaenoptera musculus) located inshore and offshore of southern California, singular A and B calls, D calls, and AB phrases were analyzed from 12 mo of passive acoustic data collected at four locations within the Southern California Bight. The relative proportions of singular calls and phrases were used to evaluate spatial and temporal patterns in sound and song type usage, and singular call and phrase production rates were calculated to investigate spatial and temporal variability in call abundance. Blue whale sounds were recorded from spring through early winter, with the majority of all detections occurring between September and December. The proportions and production rates of singular calls and phrases varied between the inshore and offshore sites. In addition, the percentage of A units within repetitive song phrases was greater inshore than offshore, resulting from a higher proportion of AB song type inshore, in which A and B phrase units were alternating. The ABB song type, in which a single A unit was followed by multiple B units, was more common offshore. The observed differences in calling and singing behaviors may identify distinct and variable acoustic behavioral settings for blue whales off southern California.     

Habitat-dependent advertisement call variation in the monkey frog Phyllomedusa nordestina

The acoustic adaptation hypothesis (AAH) predicts that acoustic signals are selected to propagate more efficiently in the habitat where they are normally transmitted. Several studies corroborated the AAH for primates and birds, but evidence for frogs is contentious: While most studies failed to support the AAH, recent studies have shown that within-species variation conforms to the predictions of the AAH. Herein, we test the AAH by comparing advertisement calls of Phyllomedusa nordestina from two contrasting habitats (Atlantic Rainforest and Caatinga) and by testing the influence of the amount of vegetation around calling sites on acoustic parameters of frog calls. The interval between pulses was significantly different between individuals from the Atlantic Rainforest and from the Caatinga, and the number of pulses was correlated with the amount of vegetation around calling sites. Hence, our results indicate that multiple evolutionary forces may act simultaneously on the advertisement calls of frogs.     

The neuropeptide arginine vasotocin alters male call characteristics involved in social interactions in the grey treefrog, Hyla versicolor

We investigated the effects of different doses (0, 2.5, 25 and 250 μg) of the neuropeptide arginine vasotocin (AVT) on the calling characteristics of the grey treefrog in a chorus in its natural habitat. AVT changed some call characteristics known to influence social behaviour in grey treefrogs. It increased call duration and number of pulses in a call, but not dominant frequency, call rate or pulse effort. Saline injections and handling did not produce significant changes in any of the call characteristics. In addition, individual animals injected with AVT only rarely produced call characteristics that were outside of the range found in the preinjection measurements, suggesting that AVT does not cause abnormal calling behaviour. Other researchers have demonstrated that longer calls with more pulses are produced by males when chorus densities increase, and females display a strong preference for longer calls with more pulses. This suggests that the changes induced by AVT injections may have functional consequences in social interactions. We previously demonstrated that AVT-injected males (25 μg AVT) displaced resident males from calling sites through changes in calling behaviour under natural field conditions. Our results indicate that changes in call duration and pulse number could contribute to the unmanipulated resident male's behaviour towards the AVT-injected intruder, perhaps because the calls are more attractive to females or because the calls are perceived as more aggressive. Copyright 2000 The Association for the Study of Animal Behaviour.     

Flying south: Foraging locations of the Hutton's shearwater (Puffinus huttoni) revealed by Time‐Depth Recorders and GPS tracking

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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.     

UNDERWATER NOISE OF WHALE-WATCHING BOATS AND POTENTIAL EFFECTS ON KILLER WHALES (ORCINUS ORCA), BASED ON AN ACOUSTIC IMPACT MODEL

Underwater noise of whale-watching boats was recorded in the popular killer whale-watching region of southern British Columbia and northwestern Washington State. A software sound propagation and impact assessment model was applied to estimate zones around whale-watching boats where boat noise was audible to killer whales, where it interfered with their communication, where it caused behavioral avoidance, and where it possibly caused hearing loss. Boat source levels ranged from 145 to 169 dB re 1 μPa @ 1 m, increasing with speed. The noise of fast boats was modeled to be audible to killer whales over 16 km, to mask killer whale calls over 14 km, to elicit a behavioral response over 200 m, and to cause a temporary threshold shift (TTS) in hearing of 5 dB after 30–50 min within 450 m. For boats cruising at slow speeds, the predicted ranges were 1 km for audibility and masking, 50 m for behavioral responses, and 20 m for TTS. Superposed noise levels of a number of boats circulating around or following the whales were close to the critical level assumed to cause a permanent hearing loss over prolonged exposure. These data should be useful in developing whale-watching regulations. This study also gave lower estimates of killer whale call source levels of 105–124 dB re 1 μPa.