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Flexible Information in the Social Sounds of Humpback Whales

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Flexible Information in the Social Sounds of Humpback Whales Flexible Information in the Social Sounds of Humpback Whales Dana Anne Cusano Bachelor of Arts (cum laude), Master of Research 0000-0002-4186-4206 A thesis submitted for the degree of Doctor of Philosophy at The University of Queensland in 2020 School of Veterinary Science i Abstract Animals living in a highly social environment typically have frequent and diverse interactions. To facilitate these relationships, social animals often have complex communication systems consisting of both between- and within-call variation. Such variability may manifest in a diverse number of call types (between-call variation) as well as the potential for conveying information on the signaller’s internal motivational state or arousal (within-call variation). These aspects may be particularly important for social species or during complex social interactions, a concept known as the ‘social complexity hypothesis for communicative complexity’. However, not all species appear to conform to these trends. The humpback whale (Megaptera novaeangliae), like other baleen whales, has a supposedly simple social system characterised by small, temporary, and unstable groups. Despite this, humpback whales have one of the most complex communication systems of any non-human animal, especially during breeding-related social interactions. Although these interactions are undoubtedly mediated using acoustic signals, how potential information is conveyed (e.g. through call types and/or through changes in the structure of the calls) is poorly understood. This thesis examines the potential information in the social calls of humpback whales, with a particular emphasis on within-call structural variation (e.g. changes in frequency, duration, or bandwidth). As humpback whales are thought to have a simple social system, this thesis also aims to determine the potential link between the complex communication system of this species and social interactions during behaviours associated with breeding. To do this, data were collected during interactions between males and females on the breeding grounds in the Great Barrier Reef and during their annual southward migration back towards the feeding grounds in Antarctica. The variety of these breeding interactions ranges from simple social interactions between a single female and her calf, to moderately social groups consisting of a female-calf pair escorted by a single male, and ultimately to highly social competitive groups comprised of multiple males aggressively competing for access to a female. Using the breeding ground data, competitive groups were further split into three ‘intensity’ levels, defined by a progressive increase in their levels of aggression and arousal. A novel quantitative method was used to classify the acoustic repertoire of social calls in humpback whales and determine which call types were structurally stereotyped (‘discrete’) or structurally variable (‘graded’). The repertoire was found to consist of 15 call types, made up of six discrete and ii nine graded calls. However, even the discrete calls showed significant variability in their acoustic structure, especially during social interactions. As acoustic variation is correlated with relatively high informative value, this indicates that all humpback whale calls have the potential to convey a large amount of information. Some calls also likely contain ‘multiple messages’ and may convey information on more than one attribute, such as temporary dominance status and individual identity. In female-calf only groups, humpback whales produced calls at lower rates and used fewer call types. As the number and diversity of social interactions increased (defined as the addition of one or more male escorts), individual call rates increased, as did the use of long bouts of calls. Further, new discrete call types were introduced into the repertoire which were never heard in the more simple female-calf dyads. This meant that some calls demonstrated a degree of context-specificity, and were only heard in groups with one or more escorts. The use of certain calls in limited contexts is another characteristic linked with increased communicative capacity. This may be especially true if they also display acoustic features that are indicative of certain motivational states or arousal levels. Graded call types were used more frequently than discrete call types across all levels of social interaction. However, during competitive behaviour, there was a significant increase in the overall proportion of graded calls used compared with discrete calls as the levels of aggression and group arousal increased. There was also a significant increase in the rate of individual call production with increased intensity. These results demonstrate a striking similarity between the displays and ritualised fighting in terrestrial animals, a concept not described in other cetaceans. This thesis represents a dedicated and comprehensive assessment of the information content of humpback whale social calls, and one of the first to investigate the relationship between social interactions and communicative complexity in a marine mammal. The results of this research have provided insight into the repertoire of the humpback whale, and which call types are likely to contain information relevant during social interactions related to breeding. Overall, the results presented here support high communicative complexity in this species, both in the repertoire and within call types. Not only does this represent a critical step towards identifying the function of calls, it has broad implications for animal behaviour and communication in general. Using the classification methods for future analyses of other whale species will allow for comparisons which can inform researchers about the possible drivers of communicative complexity in this group of animals. iii Declaration by author This thesis is composed of my original work, and contains no material previously published or written by another person except where due reference has been made in the text. I have clearly stated the contribution by others to jointly-authored works that I have included in my thesis. I have clearly stated the contribution of others to my thesis as a whole, including statistical assistance, survey design, data analysis, significant technical procedures, professional editorial advice, financial support and any other original research work used or reported in my thesis. The content of my thesis is the result of work I have carried out since the commencement of my higher degree by research candidature and does not include a substantial part of work that has been submitted to qualify for the award of any other degree or diploma in any university or other tertiary institution. I have clearly stated which parts of my thesis, if any, have been submitted to qualify for another award. I acknowledge that an electronic copy of my thesis must be lodged with the University Library and, subject to the policy and procedures of The University of Queensland, the thesis be made available for research and study in accordance with the Copyright Act 1968 unless a period of embargo has been approved by the Dean of the Graduate School. I acknowledge that copyright of all material contained in my thesis resides with the copyright holder(s) of that material. Where appropriate I have obtained copyright permission from the copyright holder to reproduce material in this thesis and have sought permission from co-authors for any jointly authored works included in the thesis. iv Publications included in this thesis No publications included. Submitted manuscripts included in this thesis The manuscript appearing as Chapter 2 ‘Humpback whale social call production reflects both motivational state and arousal’ has been accepted to the peer-reviewed journal Bioacoustics. This manuscript was a collaborative effort, with specific author contributions listed on page 17. Other publications during candidature Peer-reviewed papers 1. Parks SE, Cusano DA, Van Parijs SM, Nowacek DP. 2019. Evidence for acoustic crypsis in communication by right whale mother-calf pairs on the calving grounds. Biology Letters 15:1- 6. 2. Parks SE, Cusano DA, Van Parijs SM, Nowacek DP. 2019. North Atlantic right whale (Eubalaena glacialis) acoustic behavior on the calving grounds. Journal of the Acoustical Society of America 146: EL15-21. 3. Cusano DA, Conger LA, Van Parijs SM, Parks SE. 2018. Implementing conservation measures for the North Atlantic right whale: considering the behavioral ontogeny of mother- calf pairs. Animal Conservation 22: 228-237. 4. Root-Gutteridge H, Cusano DA, Shiu Y, Nowacek DP, Van Parijs SM, Parks SE. 2018. A lifetime of changing calls: North Atlantic right whales, Eubalaena glacialis, refine call production as they age. Animal Behaviour 137: 21-34. 5. Petraccione J, Root-Gutteridge H, Cusano DA, Parks SE. 2017. Exploring the early social affiliations and behaviour of a captive Asian elephant (Elephas maximus) calf. Journal of Zoo and Aquarium Research 5: 131-136. 6. Hunsinger E, Root-Gutteridge H, Cusano DA, Parks SE. 2017. A description of defensive hiss types in the flat horned hissing cockroach (Aeluropoda insignis). Bioacoustics 27: 261-271. 7. Parks SE, Cusano DA, Bocconcelli A, Friedlaender AS, Wiley DN. 2016. Noise impacts on social sound production by foraging humpback whales. Proceedings of Meetings on Acoustics 27: 01009. v 8. Cusano DA, Matthews LP, Grapsten E, Parks SE. 2016. Effects of Increasing Temperature on Acoustic Advertisement in the Tettigoniidae.
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