Michael J. Noad

Songs of male humpback whales, Megaptera novaeangliae, are involved in intersexual interactions

Male humpback whales produce complex songs during the breeding season, yet the singing behaviour of males and whether songs function in male contests and/or through female choice are still poorly understood. We investigated song function by obtaining simultaneous observations of the positions and movements of singing and nonsinging whales in real time during their migration off the east coast of Australia. We collected movement data by acoustic tracking using a hydrophone array, land-based visual tracking and observations from a small boat. Of the 114 singers analysed, 66 (58%) associated with conspecifics. Singers were significantly more likely to join groups containing a mother–calf pair than other groups. Males started to sing after joining groups only if they consisted of a mother–calf pair not escorted by another male. Singers also associated longer and sang for a significantly greater proportion of time with mother–calf pairs than any other group type. Associating with mother–calf pairs has been shown to be a reproductively successful strategy for males. In contrast, whales that joined singers were usually lone males; these associations were brief and singers typically stopped singing in the presence of other males. This is the highest reported incidence in humpback whales of males singing when escorting females and supports an intersexual function of song in humpback whales. We suggest that males joining singers are prospecting for females rather than engaging in male social ordering and that singing may incur the cost of attracting competing males.

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.

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

SUMMARY 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.

Humpback whale song on the southern ocean feeding grounds : implications for cultural transmission

Male humpback whales produce a long, complex, and stereotyped song on low-latitude breeding grounds; they also sing while migrating to and from these locations, and occasionally in high-latitude summer feeding areas. All males in a population sing the current version of the constantly evolving display and, within an ocean basin, populations sing similar songs; however, this sharing can be complex. In the western and central South Pacific region there is repeated cultural transmission of song types from eastern Australia to other populations eastward. Song sharing is hypothesized to occur through several possible mechanisms. Here, we present the first example of feeding ground song from the Southern Ocean Antarctic Area V and compare it to song from the two closest breeding populations. The early 2010 song contained at least four distinct themes; these matched four themes from the eastern Australian 2009 song, and the same four themes from the New Caledonian 2010 song recorded later in the year. This provides evidence for at least one of the hypothesized mechanisms of song transmission between these two populations, singing while on shared summer feeding grounds. In addition, the feeding grounds may provide a point of acoustic contact to allow the rapid horizontal cultural transmission of song within the western and central South Pacific region and the wider Southern Ocean.

Vertical distribution of lipids, fatty acids and organochlorine contaminants in the blubber of southern hemisphere humpback whales (Megaptera novaeangliae)

Persistent organic pollutants (POPs), such as toxic lipophilic organochlorine (OC) compounds, accumulate in the blubber tissue of marine mammals. Toxicological sampling methods most frequently target only the superficial blubber layer. Vertical distribution of these contaminants through the blubber mantle may, however, not be homogenous and could reflect any dissemination of lipids and fatty acids (FAs). It is therefore critical to assess stratification patterns in a species of interest as a quality control measure for interpretation of toxicological data. Here, we analysed and compared the distribution of lipids, FAs, and OCs in the outermost and innermost blubber layer of southern hemisphere humpback whales. FA stratification was evident for short-chain (≤18) monounsaturated fatty acids (SC-MUFA), which were concentrated in the outer layer, consistent with the thermoregulatory role of this blubber layer. This stratification was, however, not reflected in OC distribution, which was similar in the inner and outer blubber layers of male humpback whales. By comparison, a noticeable gradient in total blubber lipid from the outer to the inner layer was observed in two lactating females, which coincided with higher lipid normalised contaminant levels in the inner layer. This study contains the most comprehensive assessment of humpback whale blubber stratification to date, however, further investigation of biological and ecological influencing factors is required.

Improved versions of the Levenshtein distance method for comparing sequence information in animals’ vocalisations: tests using humpback whale song

Animals can communicate using visual and acoustic displays to convey information to conspecifics. In some cases, such displays are produced in highly stereotyped and repetitive sequences. Here we use a quantitative analysis technique, the Levenshtein distance, to assess similarity in sequences of displays at both the population and individual levels. We review two existing variations of the method and present two new variations that complement and extend these existing techniques. Three of the methods include the use of a median string sequence and three use a normalisation of the original equation. Humpback whale song theme sequences from multiple populations, years and song types (different variations of the display) are used as examples to illustrate the application and success of each variation. A novel outcome of this technique is that it can produce a threshold measure of similarity to assess when behavioural sequences are so dissimilar that they must be considered different, with a measure of the probability of such clusters being distinct. The Levenshtein distance is applicable to all behavioural data produced in sequences and its use should not be limited to acoustical studies.

Imitating the neighbours: vocal dialect matching in a mimic-model system

Vocal mimicry provides a unique system for investigating song learning and cultural evolution in birds. Male lyrebirds produce complex vocal displays that include extensive and accurate mimicry of many other bird species. We recorded and analysed the songs of the Alberts lyrebird (Menura alberti) and its most commonly imitated model species, the satin bowerbird (Ptilonorhynchus violaceus), at six sites in southeast Queensland, Australia. We show that each population of lyrebirds faithfully reproduces the song of the local population of bowerbirds. Within a population, lyrebirds show less variation in song structure than the available variation in the songs of the models. These results provide the first quantitative evidence for dialect matching in the songs of two species that have no direct ecological relationship.

The Behavioural Response of Humpback Whales (Megaptera novaeangliae) to a 20 Cubic Inch Air Gun

Seismic surveys are widely used for exploration for oil and gas deposits below the sea floor. Despite concern they may have an impact on whale behaviour, our knowledge of marine mammal responses is limited. In the first of a series of experiments (the last one involving a full seismic array), this study tested the response of migrating humpback whale (Megaptera novaeangliae) groups to a 20 cubic inch air gun. Experiments were carried out during the southward migration of humpback whales along the east coast of Australia. Groups of whales were focally followed from land stations and/or small boats with observations before, during, and after exposure to a vessel towing the air gun. The source vessel moved either eastwards across the, migratory flow or northwards into the migratory flow In total, there were 18 control trials (where the source vessel ran the compressor and towed the air gun without it firing; n = 35 whale groups) and 16 active trials (where the air gun was firing every 11 s; a = 32 whale groups). The air gun source level was 199 dB re 1 mu Pa(2.)s (Sound Exposure Level [SEC]) at 1 m, and SELs received by the whales varied from 105 to 156 dB re 1 mu Pa(2.)s (modal value 128 dB re mu Pa(2.)s) for SELs at least 10 dB above the background noise (measured as dB re 1 mu Pa). Other baseline groups were focal followed when there was no source vessel in the area (n = 25). Results suggested that humpback whale, groups responded by decreasing both dive time and speed of southwards movement though the response magnitude was not found to be related to the proximity of the source vessel, the received level of the air gun, the tow path direction, or the exposure time, within the during phase. There was no evidence of orientation of the groups towards, or away from, the source vessel in the during phase. Interestingly, this behavioural response was found in the control trials as well as the active trials suggesting a response to the source vessel.

Non-song social call bouts of migrating humpback whales

The use of stereotyped calls within structured bouts has been described for a number of species and may increase the information potential of call repertoires. Humpback whales produce a repertoire of social calls, although little is known about the complexity or function of these calls. In this study, digital acoustic tag recordings were used to investigate social call use within bouts, the use of bouts across different social contexts, and whether particular call type combinations were favored. Call order within bouts was investigated using call transition frequencies and information theory techniques. Call bouts were defined through analysis of inter-call intervals, as any calls within 3.9 s of each other. Bouts were produced significantly more when new whales joined a group compared to groups that did not change membership, and in groups containing multiple adults escorting a female and calf compared to adult only groups. Although social calls tended to be produced in bouts, there were few repeated bout types. However, the order in which most call types were produced within bouts was non-random and dependent on the preceding call type. These bouts appear to be at least partially governed by rules for how individual components are combined.

Population structure of humpback whales in the western and central South Pacific Ocean as determined by vocal exchange among populations

For cetaceans, population structure is traditionally determined by molecular genetics or photographically identified individuals. Acoustic data, however, has provided information on movement and population structure with less effort and cost than traditional methods in an array of taxa. Male humpback whales (Megaptera novaeangliae) produce a continually evolving vocal sexual display, or song, that is similar among all males in a population. The rapid cultural transmission (the transfer of information or behavior between conspecifics through social learning) of different versions of this display between distinct but interconnected populations in the western and central South Pacific region presents a unique way to investigate population structure based on the movement dynamics of a song (acoustic) display. Using 11 years of data, we investigated an acoustically based population structure for the region by comparing stereotyped song sequences among populations and years. We used the Levenshtein distance technique to group previously defined populations into (vocally based) clusters based on the overall similarity of their song display in space and time. We identified the following distinct vocal clusters: western cluster, 1 population off eastern Australia; central cluster, populations around New Caledonia, Tonga, and American Samoa; and eastern region, either a single cluster or 2 clusters, one around the Cook Islands and the other off French Polynesia. These results are consistent with the hypothesis that each breeding aggregation represents a distinct population (each occupied a single, terminal node) in a metapopulation, similar to the current understanding of population structure based on genetic and photo-identification studies. However, the central vocal cluster had higher levels of song-sharing among populations than the other clusters, indicating that levels of vocal connectivity varied within the region. Our results demonstrate the utility and value of using culturally transmitted vocal patterns as a way of defining connectivity to infer population structure. We suggest vocal patterns be incorporated by the International Whaling Commission in conjunction with traditional methods in the assessment of structure.