Characterising resting patterns of mother-calf humpback whale groups in a semi-enclosed embayment along the Australian east coast migration pathway.

On migration from low latitude breeding grounds to high latitude feeding grounds, humpback whale mothers and calves spend time resting in coastal embayments. Unlike other areas where resting has been documented, Jervis Bay, on Australia's east coast, is remote from both breeding and feeding grounds, and provides a unique opportunity to compare resting behaviour observed within a semi-enclosed embayment to observations offshore. Land-based, and UAV surveys were conducted in Jervis Bay in 2018, 2019, and 2021. We show that (i) a disproportionately high percentage of groups with a calf enter Jervis Bay during the southbound migration, (ii) travelling speeds are significantly slower in the Bay compared to offshore, indicating resting behaviour, and (iii) aerial observations highlight resting and nurturing behaviour. Subsequently, we conclude that Jervis Bay is an important area for resting mother-calf humpback whale groups. Comparison with reports of resting behaviour during migration in areas nearer the breeding grounds shows commonalities that characterise resting behaviour in mothers and calves. This characterisation will allow improved monitoring and management of humpback whales in nearshore embayments during a critical stage of calf development, particularly those with increased anthropogenic activities.

Source levels of humpback whales decrease with frequency suggesting an air-filled resonator is used in sound production.

Source level and frequency are important in determining how far an acoustic signal can travel. However, in some species these sound characteristics have been found to be biomechanically linked, and therefore cannot be modified independently to achieve optimal transmission. This study investigates the variability in source levels and their relationship with frequency in the songs of humpback whales (Megaptera novaeangliae). Songs were recorded off eastern Australia using a fixed hydrophone array deployed on the whales' migratory corridor. Singing whales were acoustically tracked. An empirical, frequency-dependent model was used to estimate transmission loss. Source levels and frequency were measured for 2408 song units from 19 singers. Source levels varied from 138 to 187 dB re 1 µPa at 1 m (root mean squared), while peak frequency ranged between 52 and 3877 Hz. Much of the variability in source levels was accounted for by differences between the unit types, with mean source levels for each unit type varying by up to 17 dB. Source levels were negatively correlated with peak frequency and decreased by 2.3 dB per octave. The negative correlation between source levels and frequency is consistent with the presence of an air-filled resonator in the whales' sound production system.

A behavioural dose-response model for migrating humpback whales and seismic air gun noise.

The behavioural responses of migrating humpback whales to an air gun, a small clustered seismic array and a commercial array were used to develop a dose-response model, accounting for the presence of the vessel, array towpath relative to the migration and social and environmental parameters. Whale groups were more likely to show an avoidance response (increasing their distance from the source) when the received sound exposure level was over 130 dB re 1 µPa2·s and they were within 4 km of the source. The 50% probability of response occurred where received levels were 150-155 dB re 1 µPa2·s and they were within 2.5 km of the source. A small number of whales moving rapidly close to the source vessel did not exhibit an avoidance response at the highest received levels (160-170 dB re 1 µPa2·s) meaning it was not possible to estimate the maximum response threshold.

The behavioural response of migrating humpback whales to a full seismic airgun array.

Despite concerns on the effects of noise from seismic survey airguns on marine organisms, there remains uncertainty as to the biological significance of any response. This study quantifies and interprets the response of migrating humpback whales (Megaptera novaeangliae) to a 3130 in3 (51.3l) commercial airgun array. We compare the behavioural responses to active trials (array operational; n = 34 whale groups), with responses to control trials (source vessel towing the array while silent; n = 33) and baseline studies of normal behaviour in the absence of the vessel (n = 85). No abnormal behaviours were recorded during the trials. However, in response to the active seismic array and the controls, the whales displayed changes in behaviour. Changes in respiration rate were of a similar magnitude to changes in baseline groups being joined by other animals suggesting any change group energetics was within their behavioural repertoire. However, the reduced progression southwards in response to the active treatments, for some cohorts, was below typical migratory speeds. This response was more likely to occur within 4 km from the array at received levels over 135 dB re 1 µPa2s.

Determining the behavioural dose-response relationship of marine mammals to air gun noise and source proximity.

The effect of various anthropogenic sources of noise (e.g. sonar, seismic surveys) on the behaviour of marine mammals is sometimes quantified as a dose-response relationship, where the probability of an animal behaviourally 'responding' (e.g. avoiding the source) increases with 'dose' (or received level of noise). To do this, however, requires a definition of a 'significant' response (avoidance), which can be difficult to quantify. There is also the potential that the animal 'avoids' not only the source of noise but also the vessel operating the source, complicating the relationship. The proximity of the source is an important variable to consider in the response, yet difficult to account for given that received level and proximity are highly correlated. This study used the behavioural response of humpback whales to noise from two different air gun arrays (20 and 140 cubic inch air gun array) to determine whether a dose-response relationship existed. To do this, a measure of avoidance of the source was developed, and the magnitude (rather than probability) of this response was tested against dose. The proximity to the source, and the vessel itself, was included within the one-analysis model. Humpback whales were more likely to avoid the air gun arrays (but not the controls) within 3 km of the source at levels over 140 re. 1 µPa2 s-1, meaning that both the proximity and the received level were important factors and the relationship between dose (received level) and response is not a simple one.

Evening choruses in the Perth Canyon and their potential link with Myctophidae fishes.

An evening chorus centered at near 2.2 kHz was detected across the years 2000 to 2014 from seabed receivers in 430-490 m depth overlooking the Perth Canyon, Western Australia. The chorus reached a maximum level typically 2.1 h post-sunset and normally ran for 2.1 h (between 3 dB down points). It was present at lower levels across most of the hours of darkness. Maximum chorus spectrum levels were 74-76 dB re 1 µPa2/Hz in the 2 kHz 1/3 octave band, averaging 6-12 dB and up to 30 dB greater than pre-sunset levels. The chorus displayed highest levels over April to August each year with up to 10 dB differences between seasons. The spatial extent of the chorus was not determined but exceeded the sampling range of 13-15 km offshore from the 300 m depth contour and 33 km along the 300 m depth contour. The chorus comprised short damped pulses. The most likely chorus source is considered to be fishes of the family Myctophidae foraging in the water column. The large chorus spatial extent and its apparent correlation with regions of high productivity suggest it may act as an acoustic beacon to marine fauna indicating regions of high biomass.

Response of humpback whales (Megaptera novaeangliae) to ramp-up of a small experimental air gun array.

'Ramp-up', or 'soft start', is a mitigation measure used in seismic surveys and involves increasing the radiated sound level over 20-40 min. This study compared the behavioural response in migrating humpback whales to the first stages of ramp-up with the response to a 'constant' source, 'controls' (in which the array was towed but not operated) with groups in the absence of the source vessel used as the 'baseline'. Although the behavioural response, in most groups, resulted in an increase in distance from the source (potential avoidance), there was no evidence that either 'ramp-up' or the constant source at a higher level was superior for triggering whales to move away from the source vessel. 'Control' groups also responded suggesting the presence of the source vessel had some effect. However, the majority of groups appeared to avoid the source vessel at distances greater than the radius of most mitigation zones.

Addressing Challenges in Studies of Behavioral Responses of Whales to Noise.

Studying the behavioral response of whales to noise presents numerous challenges. In addition to the characteristics of the noise exposure, many factors may affect the response and these must be measured and accounted for in the analysis. An adequate sample size that includes matching controls is crucial if meaningful results are to be obtained. Field work is thus complicated, logistically difficult, and expensive. This paper discusses some of the challenges and how they are being met in a large-scale multiplatform project in which humpback whales are exposed to the noise of seismic air guns.

Regional Variations and Trends in Ambient Noise: Examples from Australian Waters.

Studies of ambient noise south of Australia show higher levels at low frequencies in the deep water off the continental shelf compared with locations on the shelf. The difference arises because of differences in transmission loss. Marine animals would experience significantly different noise levels and directionality in the two regions and while crossing the boundary, provide positional information. Opportunities for long-range, low-frequency communication by animals would be significantly limited by the higher background noise in the open ocean. Measures of long-term sea noise trends highlight the influence of biological sources and the importance of local sound transmission regimens.

Characterization of the Sounds Produced by Temperate and Tropical Sea Urchins During Feeding (Diadematidae and Echinometridae).

Despite the abundance and ecological importance of sea urchins in eastern Australia, it is not known how they may contribute to ambient noise. The sounds of feeding of two temperate and two tropical species were recorded in captivity. Most sound was produced between 2.3 and 9.2 kHz, but there were differences between species and substrate types.

Evidence of a Lombard response in migrating humpback whales (Megaptera novaeangliae).

The Lombard reflex is an increase in the subject's vocal levels in response to increased noise levels. This functions to maintain an adequate signal-to-noise ratio at the position of the receiver when noise levels vary. While it has been demonstrated in a small number of mammals and birds including some whales, it has not yet been shown to occur in one of the most vocal species of baleen whale, the humpback whale (Megaptera novaeangliae). Humpback whales were simultaneously visually and acoustically tracked (using an array of calibrated hydrophone buoys) as they migrated southward. Source levels of social vocalizations were estimated from measured received levels and a site-specific empirical sound propagation model developed. In total, 226 social vocalizations from 16 passing groups of whales were selected for final analysis. Noise levels were predominantly wind-dependent (from sea surface motion) and ranged from 81 to 108 dB re 1 µPa in the 36 Hz-2.8 kHz band. Vocalization source levels increased by 0.9 dB for every 1 dB increase in wind-dependent background noise levels, with source levels (at 1 m) being maintained ∼60 dB above the noise level.

Source levels of social sounds in migrating humpback whales (Megaptera novaeangliae).

The source level of an animal sound is important in communication, since it affects the distance over which the sound is audible. Several measurements of source levels of whale sounds have been reported, but the accuracy of many is limited because the distance to the source and the acoustic transmission loss were estimated rather than measured. This paper presents measurements of source levels of social sounds (surface-generated and vocal sounds) of humpback whales from a sample of 998 sounds recorded from 49 migrating humpback whale groups. Sources were localized using a wide baseline five hydrophone array and transmission loss was measured for the site. Social vocalization source levels were found to range from 123 to 183 dB re 1 µPa @ 1 m with a median of 158 dB re 1 µPa @ 1 m. Source levels of surface-generated social sounds ("breaches" and "slaps") were narrower in range (133 to 171 dB re 1 µPa @ 1 m) but slightly higher in level (median of 162 dB re 1 µPa @ 1 m) compared to vocalizations. The data suggest that group composition has an effect on group vocalization source levels in that singletons and mother-calf-singing escort groups tend to vocalize at higher levels compared to other group compositions.

Multivariate analysis of behavioural response experiments in humpback whales (Megaptera novaeangliae).

The behavioural response study (BRS) is an experimental design used by field biologists to determine the function and/or behavioural effects of conspecific, heterospecific or anthropogenic stimuli. When carrying out these studies in marine mammals it is difficult to make basic observations and achieve sufficient samples sizes because of the high cost and logistical difficulties. Rarely are other factors such as social context or the physical environment considered in the analysis because of these difficulties. This paper presents results of a BRS carried out in humpback whales to test the response of groups to one recording of conspecific social sounds and an artificially generated tone stimulus. Experiments were carried out in September/October 2004 and 2008 during the humpback whale southward migration along the east coast of Australia. In total, 13 'tone' experiments, 15 'social sound' experiments (using one recording of social sounds) and three silent controls were carried out over two field seasons. The results (using a mixed model statistical analysis) suggested that humpback whales responded differently to the two stimuli, measured by changes in course travelled and dive behaviour. Although the response to 'tones' was consistent, in that groups moved offshore and surfaced more often (suggesting an aversion to the stimulus), the response to 'social sounds' was highly variable and dependent upon the composition of the social group. The change in course and dive behaviour in response to 'tones' was found to be related to proximity to the source, the received signal level and signal-to-noise ratio (SNR). This study demonstrates that the behavioural responses of marine mammals to acoustic stimuli are complex. In order to tease out such multifaceted interactions, the number of replicates and factors measured must be sufficient for multivariate analysis.

Physical biologists and biological physicists: combining biology and physics in research on the effects of noise on aquatic life.

Research in aquatic bioacoustics and the effects of noise is interdisciplinary and to be effective requires a collaboration of experts from all the fields involved. The full range of expertise is needed for adequate understanding of the processes involved, adequate experimental design, analysis and interpretation, and adequate knowledge of the research already published. The biologists need to understand how physicists work and make allowance, and vice versa. Both need to understand that the other will not be familiar with their practices and approach and that there will be a certain amount of negotiation and education on both sides.However, the best reason to develop collaborations with other experts in interdisciplinary research is that it is such a rewarding experience from the insights it provides into other disciplines and from the opportunity to do really effective and very significant research, well beyond what the individuals might have achieved on their own.

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.

Information theory analysis of Australian humpback whale song.

Songs produced by migrating whales were recorded off the coast of Queensland, Australia, over six consecutive weeks in 2003. Forty-eight independent song sessions were analyzed using information theory techniques. The average length of the songs estimated by correlation analysis was approximately 100 units, with song sessions lasting from 300 to over 3100 units. Song entropy, a measure of structural constraints, was estimated using three different methodologies: (1) the independently identically distributed model, (2) a first-order Markov model, and (3) the nonparametric sliding window match length (SWML) method, as described by Suzuki et al. [(2006). "Information entropy of humpback whale song," J. Acoust. Soc. Am. 119, 1849-1866]. The analysis finds that the song sequences of migrating Australian whales are consistent with the hierarchical structure proposed by Payne and McVay [(1971). "Songs of humpback whales," Science 173, 587-597], and recently supported mathematically by Suzuki et al. (2006) for singers on the Hawaiian breeding grounds. Both the SWML entropy estimates and the song lengths for the Australian singers in 2003 were lower than that reported by Suzuki et al. (2006) for Hawaiian whales in 1976-1978; however, song redundancy did not differ between these two populations separated spatially and temporally. The average total information in the sequence of units in Australian song was approximately 35 bits/song. Aberrant songs (8%) yielded entropies similar to the typical songs.

The social vocalization repertoire of east Australian migrating humpback whales (Megaptera novaeangliae).

Although the songs of humpback whales have been extensively studied, other vocalizations and percussive sounds, referred to as "social sounds," have received little attention. This study presents the social vocalization repertoire of migrating east Australian humpback whales from a sample of 660 sounds recorded from 61 groups of varying composition, over three years. The social vocalization repertoire of humpback whales was much larger than previously described with a total of 34 separate call types classified aurally and by spectrographic analysis as well as statistically. Of these, 21 call types were the same as units of the song current at the time of recording but used individually instead of as part of the song sequence, while the other 13 calls were stable over the three years of the study and were not part of the song. This study provides a catalog of sounds that can be used as a basis for future studies. It is an essential first step in determining the function, contextual use and cultural transmission of humpback social vocalizations.