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 mu Pa in the 36Hz-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 similar to 60 dB above the noise level. (C) 2014 Acoustical Society of America

Temporal stability and change in the social call repertoire of migrating humpback whales

Quantifying the stability of a species vocal repertoire is fundamental for further investigations into repertoire function and geographic variation. Changes to the repertoire of sounds used in the song displays of male humpback whales have been well studied. In contrast, little is known about the stability of this species' non-song vocal calls. The stability of the social call repertoire of east Australian humpback whales was investigated from 1997, 2003-2004, and 2008. Out of 46 qualitatively defined call types, 19 were classified as "song-unit calls" that tended to change with the song, and 15 were "inconsistent" and only found in one or two years. Twelve call types were "stable" and present in all years and were commonly produced (64.2% of calls). Stable calls tended to vary in some of the measured call parameters but there was no clear trend between years. This result could indicate that minor changes to calls are not permanent, but reflect individual differences in call production or the graded nature of calls within different social environments. This research has clearly identified stable calls in the call repertoire of humpback whales and while their function is not well understood, their stability suggests an important role in social interactions

Network analysis reveals underlying syntactic features in a vocally learnt mammalian display, humpback whale song

J.A.A. was funded by an Australian Government Research Training Program Scholarship and the Australian American Association University of Queensland Fellowship. E.C.G. was funded by a Royal Society Newton International Fellowship and a Royal Society University Research Fellowship. HARC was funded by the US Office of Naval Research, the Australian Defence Science and Technology Organisation, and the Australian Marine Mammal Centre. BRAHSS was funded by the E&P Sound and Marine Life Joint Industry Programme and the US Bureau of Ocean Energy Management.Vocal communication systems have a set of rules that govern the arrangement of acoustic signals, broadly defined as ‘syntax’. However, there is a limited understanding of potentially shared or analogous rules across vocal displays in different taxa. Recent work on songbirds has investigated syntax using network-based modelling. This technique quantifies features such as connectivity (adjacent signals in a sequence) and recurring patterns. Here, we apply network-based modelling to the complex, hierarchically structured songs of humpback whales (Megaptera novaeangliae) from east Australia. Given the song's annual evolving pattern and the cultural conformity of males within a population, network modelling captured the patterns of multiple song types over 13 consecutive years. Song arrangements in each year displayed clear ‘small-world’ network structure, characterized by clusters of highly connected sounds. Transitions between these connected sounds further suggested a combination of both structural stability and variability. Small-world network structure within humpback songs may facilitate the characteristic and persistent vocal learning observed. Similar small-world structures and transition patterns are found in several birdsong displays, indicating common syntactic patterns among vocal learning in multiple taxa. Understanding the syntactic rules governing vocal displays in multiple, independently evolving lineages may indicate what rules or structural features are important to the evolution of complex communication, including human language.PostprintPostprintPeer reviewe

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 mu Pa @ 1 m with a median of 158 dB re 1 mu Pa @ 1 m. Source levels of surface-generated social sounds ("breaches" and "slaps") were narrower in range (133 to 171 dB re 1 mu Pa @ 1 m) but slightly higher in level (median of 162 dB re 1 mu Pa @ 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. VC 2013 Acoustical Society of America

Quantifying humpback whale song sequences to understand the dynamics of song exchange at the ocean basin scale

Humpback whales have a continually evolving vocal sexual display, or "song," that appears to undergo both evolutionary and "revolutionary" change. All males within a population adhere to the current content and arrangement of the song. Populations within an ocean basin share similarities in their songs; this sharing is complex as multiple variations of the song (song types) may be present within a region at any one time. To quantitatively investigate the similarity of song types, songs were compared at both the individual singer and population level using the Levenshtein distance technique and cluster analysis. The highly stereotyped sequences of themes from the songs of 211 individuals from populations within the western and central South Pacific region from 1998 through 2008 were grouped together based on the percentage of song similarity, and compared to qualitatively assigned song types. The analysis produced clusters of highly similar songs that agreed with previous qualitative assignments. Each cluster contained songs from multiple populations and years, confirming the eastward spread of song types and their progressive evolution through the study region. Quantifying song similarity and exchange will assist in understanding broader song dynamics and contribute to the use of vocal displays as population identifiers

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

A study of the behavioural response of whales to the noise of seismic air guns: Design, methods and progress

The concern about the effects of the noise of human activities on marine mammals, particularly whales, has led to a substantial amount of research but there is still much that is not understood, particularly in terms of the behavioural responses to noise and the longer term biological consequences of these responses. There are many challenges in conducting experiments that adequately assess behavioural reactions of whales to noise. These include the need to obtain an adequate sample size with the necessary controls and to measure the range of variables likely to affect the observed response. Analysis is also complex. Well designed experiments are complex and logistically difficult, and thus expensive. This paper discusses the challenges involved and how these are being met in a major series of experiments in Australian waters on the response of humpback whales to the noise of seismic airgun arrays. The project is known as BRAHSS (Behavioural Response of Australian Humpback whales to Seismic Surveys) and aims to provide the information that will allow seismic surveys to be conducted efficiently with minimal impact on whales. It also includes a study of the response to ramp-up in sound level which is widely used at the start of operations, but for which there is little information to show that it is effective. BRAHSS also aims to infer the longer term biological significance of the responses from the results and the knowledge of normal behaviour. The results are expected to have relevance to other sources and species

Migratory convergence facilitates cultural transmission of humpback whale song

E.C.G. was supported by a Royal Society Newton International Fellowship and a Royal Society University Research Fellowship. L.R. was supported by the MASTS pooling initiative (The Marine Alliance for Science and Technology for Scotland) and their support is gratefully acknowledged. MASTS is funded by the Scottish Funding Council (grant reference HR09011) and contributing institutions.Cultural transmission of behaviour is important in a wide variety of vertebrate taxa from birds to humans. Vocal traditions and vocal learning provide a strong foundation for studying culture and its transmission in both humans and cetaceans. Male humpback whales (Megaptera novaeangliae) perform complex, culturally transmitted song displays that can change both evolutionarily (through accumulations of small changes) or revolutionarily (where a population rapidly adopts a novel song). The degree of coordination and conformity underlying song revolutions makes their study of particular interest. Acoustic contact on migratory routes may provide a mechanism for cultural revolutions of song, yet these areas of contact remain uncertain. Here, we compared songs recorded from the Kermadec Islands, a recently discovered migratory stopover, to multiple South Pacific wintering grounds. Similarities in song themes from the Kermadec Islands and multiple wintering locations (from New Caledonia across to the Cook Islands) suggest a location allowing cultural transmission of song eastward across the South Pacific, active song learning (hybrid songs) and the potential for cultural convergence after acoustic isolation at the wintering grounds. As with the correlations in humans between genes, communication and migration, the migration patterns of humpback whales are written into their songs.Publisher PDFPeer reviewe

The devil is in the detail : quantifying vocal variation in a complex, multi-levelled, and rapidly evolving display

E.C.G. was funded by a Royal Society Newton International Fellowship. L.R. was supported by the MASTS pooling initiative (The Marine Alliance for Science and Technology for Scotland) and their support is gratefully acknowledged. MASTS is funded by the Scottish Funding Council (Grant Reference No. HR09011) and contributing institutions. Some funding and logistical support was provided to M.M.P. by the National Oceanic Society (USA), Dolphin & Whale Watching Expeditions (French Polynesia), Vista Press (USA), and the International Fund for Animal Welfare (via the South Pacific Whale Research Consortium).Identifying and quantifying variation in vocalizations is fundamental to advancing our understanding of processes such as speciation, sexual selection, and cultural evolution. The song of the humpback whale (Megaptera novaeangliae) presents an extreme example of complexity and cultural evolution. It is a long, hierarchically structured vocal display that undergoes constant evolutionary change. Obtaining robust metrics to quantify song variation at multiple scales (from a sound through to population variation across the seascape) is a substantial challenge. Here, we present a method to quantify song similarity at multiple levels within the hierarchy. To incorporate the complexity of these multiple levels, the calculation of similarity is weighted by measurements of sound units (lower levels within the display) to bridge the gap in information between upper and lower levels. Results demonstrate that the inclusion of weighting provides a more realistic and robust representation of song similarity at multiple levels within the display. Our method permits robust quantification of cultural patterns and processes that will also contribute to the conservation management of endangered humpback whale populations, and is applicable to any hierarchically structured signal sequence.PostprintPeer reviewe