Acoustic sequences have been described in a range of species and in varying complexity. Cetaceans are known to produce complex song displays but these are generally limited to mysticetes; little is ...known about call combinations in odontocetes. Here we investigate call combinations produced by killer whales (Orcinus orca), a highly social and vocal species. Using acoustic recordings from 22 multisensor tags, we use a first order Markov model to show that transitions between call types or subtypes were significantly different from random, with repetitions and specific call combinations occurring more often than expected by chance. The mixed call combinations were composed of two or three calls and were part of three call combination clusters. Call combinations were recorded over several years, from different individuals, and several social clusters. The most common call combination cluster consisted of six call (sub-)types. Although different combinations were generated, there were clear rules regarding which were the first and last call types produced, and combinations were highly stereotyped. Two of the three call combination clusters were produced outside of feeding contexts, but their function remains unclear and further research is required to determine possible functions and whether these combinations could be behaviour- or group-specific.
Monitoring the body condition of free-ranging marine mammals at different life-history stages is essential to understand their ecology as they must accumulate sufficient energy reserves for survival ...and reproduction. However, assessing body condition in free-ranging marine mammals is challenging. We cross-validated two independent approaches to estimate the body condition of humpback whales (
) at two feeding grounds in Canada and Norway: animal-borne tags (
= 59) and aerial photogrammetry (
= 55). Whales that had a large length-standardized projected area in overhead images (i.e. whales looked fatter) had lower estimated tissue body density (TBD) (greater lipid stores) from tag data. Linking both measurements in a Bayesian hierarchical model to estimate the true underlying (hidden) tissue body density (uTBD), we found uTBD was lower (-3.5 kg m
) in pregnant females compared to adult males and resting females, while in lactating females it was higher (+6.0 kg m
). Whales were more negatively buoyant (+5.0 kg m
) in Norway than Canada during the early feeding season, possibly owing to a longer migration from breeding areas. While uTBD decreased over the feeding season across life-history traits, whale tissues remained negatively buoyant (1035.3 ± 3.8 kg m
) in the late feeding season. This study adds confidence to the effectiveness of these independent methods to estimate the body condition of free-ranging whales.
Impact assessments for sonar operations typically use received sound levels to predict behavioural disturbance in marine mammals. However, there are indications that cetaceans may learn to associate ...exposures from distant sound sources with lower perceived risk. To investigate the roles of source distance and received level in an area without frequent sonar activity, we conducted multi-scale controlled exposure experiments ( n = 3) with 12 northern bottlenose whales near Jan Mayen, Norway. Animals were tagged with high-resolution archival tags ( n = 1 per experiment) or medium-resolution satellite tags ( n = 9 in total) and subsequently exposed to sonar. We also deployed bottom-moored recorders to acoustically monitor for whales in the exposed area. Tagged whales initiated avoidance of the sound source over a wide range of distances (0.8-28 km), with responses characteristic of beaked whales. Both onset and intensity of response were better predicted by received sound pressure level (SPL) than by source distance. Avoidance threshold SPLs estimated for each whale ranged from 117-126 dB re 1 µPa, comparable to those of other tagged beaked whales. In this pristine underwater acoustic environment, we found no indication that the source distances tested in our experiments modulated the behavioural effects of sonar, as has been suggested for locations where whales are frequently exposed to sonar.
Killer whales (KW) may be predators or competitors of other cetaceans. Since their foraging behavior and acoustics differ among populations (‘ecotypes’), we hypothesized that other cetaceans can ...eavesdrop on KW sounds and adjust their behavior according to the KW ecotype. We performed playback experiments on long-finned pilot whales (
Globicephala melas
) in Norway using familiar fish-eating KW sounds (fKW) simulating a sympatric population that might compete for foraging areas, unfamiliar mammal-eating KW sounds (mKW) simulating a potential predator threat, and two control sounds. We assessed behavioral responses using animal-borne multi-sensor tags and surface visual observations. Pilot whales barely changed behavior to a broadband noise (CTRL−), whereas they were attracted and exhibited spyhops to fKW, mKW, and to a repeated-tonal upsweep signal (CTRL+). Whales never stopped nor started feeding in response to fKW, whereas they reduced or stopped foraging to mKW and CTRL+. Moreover, pilot whales joined other subgroups in response to fKW and CTRL+, whereas they tightened individual spacing within group and reduced time at surface in response to mKW. Typical active intimidation behavior displayed to fKW might be an antipredator strategy to a known low-risk ecotype or alternatively a way of securing the habitat exploited by a heterospecific sympatric population. Cessation of feeding and more cohesive approach to mKW playbacks might reflect an antipredator behavior towards an unknown KW ecotype of potentially higher risk. We conclude that pilot whales are able to acoustically discriminate between familiar and unfamiliar KW ecotypes, enabling them to adjust their behavior according to the perceived disturbance type.
Naval sonar disrupts foraging in humpback whales Sivle, Lise Doksæter; Wensveen, Paul J.; Kvadsheim, Petter H. ...
Marine ecology. Progress series (Halstenbek),
12/2016, Letnik:
562
Journal Article
Recenzirano
Odprti dostop
Modern long-range naval sonars are a potential disturbance for marine mammals and can cause disruption of feeding in cetaceans. We examined the lunge-feeding behaviour of humpback whales Megaptera ...novaeangliae before, during and after controlled exposure experiments with naval sonar by use of acoustic and motion sensor archival tags attached to each animal. Lunge-feeding by humpback whales entails a strong acceleration to increase speed before engulfing a large volume of prey-laden water, which can be identified by an acoustic signature characterized by a few seconds of high-level flow-noise followed by a rapid reduction, coinciding with a peak in animal acceleration. Over 2 successive seasons, 13 humpback whales were tagged. All were subject to a no-sonar control exposure, and 12 whales were exposed to 2 consecutive sonar exposure sessions, with 1 h between sessions. The first sonar session resulted in an average 68% reduction in lunge rate during exposure compared to pre-exposure, and this reduction was significantly greater than any changes observed during the no-sonar control. During the second sonar session, reduction in lunge rate was 66% during sonar exposure compared to the pre-exposure level, but was not significant compared to the no-sonar control, likely due to a larger inter-individual variability because some individuals appeared to have habituated whereas others had not. Our results indicate that naval sonars operating near humpback whale feeding grounds may lead to reduced foraging and negative impacts on energy balance.
The underwater hearing sensitivity of a young male harbor porpoise for tonal signals of various signal durations was quantified by using a behavioral psychophysical technique. The animal was trained ...to respond only when it detected an acoustic signal. Fifty percent detection thresholds were obtained for tonal signals (15 frequencies between 0.25-160 kHz, durations 0.5-5000 ms depending on the frequency; 134 frequency-duration combinations in total). Detection thresholds were quantified by varying signal amplitude by the 1-up 1-down staircase method. The hearing thresholds increased when the signal duration fell below the time constant of integration. The time constants, derived from an exponential model of integration Plomp and Bouman, J. Acoust. Soc. Am. 31, 749-758 (1959), varied from 629 ms at 2 kHz to 39 ms at 64 kHz. The integration times of the porpoises were similar to those of other mammals including humans, even though the porpoise is a marine mammal and a hearing specialist. The results enable more accurate estimations of the distances at which porpoises can detect short-duration environmental tonal signals. The audiogram thresholds presented by Kastelein et al. J. Acoust. Soc. Am. 112, 334-344 (2002), after correction for the frequency bandwidth of the FM signals, are similar to the results of the present study for signals of 1500 ms duration. Harbor porpoise hearing is more sensitive between 2 and 10 kHz, and less sensitive above 10 kHz, than formerly believed.
Eight experimentally controlled exposures to 1-2 kHz or 6-7 kHz sonar signals were conducted with four killer whale groups. The source level and proximity of the source were increased during each ...exposure in order to reveal response thresholds. Detailed inspection of movements during each exposure session revealed sustained changes in speed and travel direction judged to be avoidance responses during six of eight sessions. Following methods developed for Phase-I clinical trials in human medicine, response thresholds ranging from 94 to 164 dB re 1 μPa received sound pressure level (SPL) were fitted to Bayesian dose-response functions. Thresholds did not consistently differ by sonar frequency or whether a group had previously been exposed, with a mean SPL response threshold of 142 ± 15 dB (mean ± s.d.). High levels of between- and within-individual variability were identified, indicating that thresholds depended upon other undefined contextual variables. The dose-response functions indicate that some killer whales started to avoid sonar at received SPL below thresholds assumed by the U.S. Navy. The predicted extent of habitat over which avoidance reactions occur depends upon whether whales responded to proximity or received SPL of the sonar or both, but was large enough to raise concerns about biological consequences to the whales.
Abstract
Killer whales (
Orcinus orca
) have group‐specific call repertoires that can be used to track groups and populations using passive acoustic monitoring. To provide a detailed description of ...the Icelandic killer whale repertoire and its variation, we analyzed acoustic data collected in five locations between 1985 and 2016. Calls were classified manually, and CART and random forest analyses were employed to validate the manual classification. A total of 91 call categories (including call types and subtypes) were defined. Most call categories were recorded in more than one location, with the highest proportion shared between herring grounds in Vestmannaeyjar (South) and Breiðafjörður (West). However, both locations included call categories that were not recorded elsewhere in Iceland. Recordings from past herring wintering grounds in eastern Iceland included few call categories that matched other locations. Sample sizes from Reykjanes (Southwest) and Skjálfandi (North) were small and did not include unique call categories. The relative occurrence of call categories in Vestmannaeyjar changed little over a 14‐year period (2002–2016), although shorter‐term changes between years were observed that appeared to correlate to changes in individuals identified. This comparison of acoustic repertoires provides valuable information on the social structure and movement patterns of herring‐eating killer whales around Iceland.
Anti-predator strategies are often defined as ‘flight’ or ‘fight’, based upon prey anatomical adaptations for size, morphology and weapons, as well as observed behaviours in the presence of ...predators. The humpback whale Megaptera novaeangliae is considered a ‘fight’ specialist based upon anatomy and observations of grouping behaviour and active defence when attacked by killer whales. However, the early stage of humpback whale anti-predator strategy, when the prey detects the presence of a distant potential predator that may not have perceived it, has never been described. Our aim was to experimentally examine this initial stage of anti-predator responses. Humpbacks are likely to hear well at the frequencies of killer whale vocalisations, thus the perception of killer whale sounds could trigger anti-predator responses. To address this hypothesis, we played mammal-eating killer whale sounds to 8 solitary or paired humpback whales in North Atlantic feeding grounds and monitored their behavioural responses. We found that predator sound playbacks induced a cessation of feeding, a change in the diving pattern and a clear directional and rapid horizontal avoidance away from the speaker. Interestingly, in mother-calf pairs with young calves, the directional horizontal avoidance was atypically alternated by 90 degree turns, which may serve as a mechanism to better track the pre dator or a stealth tactic when more vulnerable animals are present. These results provide experimental evidence that humpback whales can exhibit a strong horizontal avoidance as an initial stage of anti-predator defence, indicating that anti-predator responses may be more graded and mixed than previously recognized.
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