Michael Poole

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.

Song hybridization events during revolutionary song change provide insights into cultural transmission in humpback whales

Cultural processes occur in a wide variety of animal taxa, from insects to cetaceans. The songs of humpback whales are one of the most striking examples of the transmission of a cultural trait and social learning in any nonhuman animal. To understand how songs are learned, we investigate rare cases of song hybridization, where parts of an existing song are spliced with a new one, likely before an individual totally adopts the new song. Song unit sequences were extracted from over 9,300 phrases recorded during two song revolutions across the South Pacific Ocean, allowing fine-scale analysis of composition and sequencing. In hybrid songs the current and new songs were spliced together in two specific ways: (i) singers placed a single hybrid phrase, in which content from both songs were combined, between the two song types when transitioning from one to the other, and/or (ii) singers spliced complete themes from the revolutionary song into the current song. Sequence analysis indicated that both processes were governed by structural similarity rules. Hybrid phrases or theme substitutions occurred at points in the songs where both songs contained “similar sounds arranged in a similar pattern.” Songs appear to be learned as segments (themes/phrase types), akin to birdsong and human language acquisition, and these can be combined in predictable ways if the underlying structural pattern is similar. These snapshots of song change provide insights into the mechanisms underlying song learning in humpback whales, and comparative perspectives on the evolution of human language and culture.

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

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.

Migratory interchange of humpback whales ( Megaptera novaeangliae ) among breeding grounds of Oceania and connections to Antarctic feeding areas based on genotype matching

Humpback whales (Megaptera novaeangliae) congregate to breed during the austral winter near tropical islands of the South Pacific (Oceania). It has long been assumed that humpback whales from Oceania migrate primarily to Antarctic feeding grounds directly south (International Whaling Commission Management Areas V and VI); however, there are few records of individual movement connecting these seasonal habitats. Based on genetic samples of living whales collected over nearly two decades, we demonstrate interchange between the breeding grounds of Oceania and Antarctic feeding Areas V, VI, and I (i.e., from 130°E to 60°W), as well as with the eastern Pacific (Colombia), and the migratory corridors of eastern Australia and New Zealand. We first compared genotype profiles (up to 16 microsatellite loci) of samples collected from Oceania breeding grounds to each other and to those from the eastern Pacific. The matching profiles documented 47 individuals that were present on more than one breeding ground, including the first record of movement between Oceania and Colombia. We then compared the 1179 genotypes from the breeding grounds to 777 from the migratory corridors of east Australia and New Zealand, confirming the connection of these corridors with New Caledonia. Finally, we compared genotypes from breeding grounds to 166 individuals from Antarctic feeding Areas I–VI. This comparison of genotypes revealed five matches: one between New Caledonia and Area V, one between Tonga and Area VI, two between Tonga and Area I (western edge), and one between Colombia and Area I (Antarctic Peninsula). Despite the relatively small number of samples from the Antarctic, our comparison has doubled the number of recorded connections with Oceania available from previous studies during the era of commercial whaling.

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

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 u...

Song learning in humpback whales: Lessons from song hybridization events during revolutionary song change

Humpback whale songs are one of the most startling examples of transmission of a cultural trait and social learning in any non-human animal. Here, we investigate extremely rare cases of song hybridization, where parts of an existing song are spliced with a novel, revolutionary song, to understand how songs are learnt. Song unit sequences were extracted from over 800 phrases recorded during a song revolution (French Polynesia 2005), to allow fine-scale analysis of composition and sequencing. Clustering of song sequences (i.e., phrases) using the Levenshtein distance indicated songs clustered into three song types; a single hybrid phrase was identified representing the transition of one singer between two of these song types. A predictive model was fitted to the data and tested against the only other known recordings of humpback song hybridization: the eastern Australia 1996-97 song revolution. Songs change during revolutions through combining multiple complete phrases and themes from one song type, before ...