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.
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, the authors 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. This 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.
2 Short title: Vocally derived population structure Word count: 5,851 (first word of Abstract through last word of Literature Cited) Abstract 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/behaviour 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 using the Levenshtein distance technique, to group previously defined populations into (vocally based) clusters using 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 two 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 4 vocal patterns be incorporated by the International Whaling Commission in conjunction with traditional methods in the assessment of structure. 5
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.
Male humpback whales (Megaptera novaeangliae) produce a continually evolving vocal sexual display, or “song,” which is shared by all males within a population. The rapid cultural transmission of this display between distinct but interconnected populations within the western and central South Pacific region presents a unique opportunity to investigate population connectivity based on a vocal display. Quantitative analyses were conducted on eleven years of data to investigate vocal groupings based on song types shared between populations, to produce an acoustically derived population structure for the region. Four distinct vocal groupings resulted; the western group contained a single population, off eastern Australia, the central group was comprised of whales around New Caledonia, Tonga and American Samoa, and finally the whales of the eastern region were split into two groups, one around the Cook Islands and the other in the waters of French Polynesia. These groupings broadly agree with results obtained using genetic and photo-identification methods, and confirm that humpback whales are likely to form separate breeding populations rather than panmictic subpopulations. This study demonstrates the utility of using culturally transmitted vocal patterns as a way of defining populations, at least in this species.
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