Social survival: Humpback whales (Megaptera novaeangliae) use social structure to partition ecological niches within proposed critical habitat.

Animal culture and social bonds are relevant to wildlife conservation because they influence patterns of geography, behavior, and strategies of survival. Numerous examples of socially-driven habitat partitioning and ecological-niche specialization can be found among vertebrates, including toothed whales. But such social-ecological dynamics, described here as 'social niche partitioning', are not known among baleen whales, whose societies-particularly on foraging grounds-are largely perceived as unstructured and incidental to matters of habitat use and conservation. However, through 16 years of behavioral observations and photo-identifications of humpback whales (Megaptera novaeangliae) feeding within a fjord system in the Canadian Pacific (primarily within Gitga'at First Nation waters), we have documented long-term pair bonds (up to 12 years) as well as a complex societal structure, which corresponds closely to persistent patterns in feeding strategy, long-term site fidelity (extended occupancy and annual rate of return up to 75%), specific geographic preferences within the fjord system, and other forms of habitat use. Randomization tests of network congruency and clustering algorithms were used to test for overlap in patterns of social structure and habitat use, which confirmed the occurrence of social niche partitioning on the feeding grounds of this baleen whale species. In addition, we document the extensive practice of group bubble net feeding in Pacific Canada. This coordinated feeding behavior was found to strongly mediate the social structure and habitat use within this humpback whale society. Additionally, during our 2004-2019 study, we observed a shift in social network structure in 2010-2012, which corresponded with environmental and demographic shifts including a sudden decline in the population's calving rate. Our findings indicate that the social lives of humpback whales, and perhaps baleen whales generally, are more complex than previously supposed and should be a primary consideration in the assessment of potential impacts to important habitat.

Vessel noise levels drive behavioural responses of humpback whales with implications for whale-watching.

Disturbance from whale-watching can cause significant behavioural changes with fitness consequences for targeted whale populations. However, the sensory stimuli triggering these responses are unknown, preventing effective mitigation. Here, we test the hypothesis that vessel noise level is a driver of disturbance, using humpback whales (Megaptera novaeangliae) as a model species. We conducted controlled exposure experiments (n = 42) on resting mother-calf pairs on a resting ground off Australia, by simulating whale-watch scenarios with a research vessel (range 100 m, speed 1.5 knts) playing back vessel noise at control/low (124/148 dB), medium (160 dB) or high (172 dB) low frequency-weighted source levels (re 1 µPa RMS@1 m). Compared to control/low treatments, during high noise playbacks the mother's proportion of time resting decreased by 30%, respiration rate doubled and swim speed increased by 37%. We therefore conclude that vessel noise is an adequate driver of behavioural disturbance in whales and that regulations to mitigate the impact of whale-watching should include noise emission standards.

Whale-watching is a multi-billion-dollar industry that is growing around the world. Typically, tour operators use boats to transport tourists into coastal waters to see groups of whales, dolphins or porpoises. There is, however, accumulating evidence that boat-based whale-watching negatively affects the way these animals behave and so many countries have put guidelines in place to mitigate activities that may disturb the animals. These guidelines generally stipulate the boat's angle of approach, how close the boat can get and the speed at which it can pass by the animals. In general, these guidelines are based on the assumption that the animals are disturbed by the closeness of the whale-watching boats. However, whales, dolphins and porpoises have very sensitive hearing, and only have a short range of vision underwater. Therefore, it seems plausible that the animals hear whale-watching boats long before they see them and so the loudness of underwater noise from the boats may be enough to disturb these animals' behaviour. To test this hypothesis, Sprogis et al. performed experiments where they simulated a whale-watching vessel approaching humpback whale mothers and calves who were resting off the northwest coast of Australia. A small motorised research boat travelling at a low speed passed different mother-calf pairs at a target distance of 100 meters, which is a common whale-watching distance guideline in many countries. The boat had an underwater speaker that played recordings of the boat noise at different volumes, while a drone with a video camera flew overhead to record the whales' behaviours in detail and to identify individual animals. These "controlled exposure experiments" showed that the quiet boat noise did not appear to disturb the mothers and calves. However, compared to when the quiet boat passed the animals the louder boat noise decreased how long the mother whale rested on the surface by 30%, made her swim 37% faster, and doubled the number of breaths she took per minute. If there are many disturbances from humans, then it can negatively impact the energy the mother and calf have available for nursing, fending off males and predators, and migrating back to their feeding ground nearer the Earth's poles. Based on these findings, it is shown that the loudness of the underwater noise from boats can explain why whales may be disturbed during whale-watching activities. To help reduce this disturbance, Sprogis et al. recommend that noise emission standards should be added to the current whale-watching regulations such that boats should be as quiet as possible and ideally around the volume of the ambient background noise. This would allow operators to approach the animals in a responsible, sustainable manner and offer tourists a view of undisturbed wildlife.

Behavioural responses of humpback whales to food-related chemical stimuli.

Baleen whales face the challenge of finding patchily distributed food in the open ocean. Their relatively well-developed olfactory structures suggest that they could identify the specific odours given off by planktonic prey such as krill aggregations. Like other marine predators, they may also detect dimethyl sulfide (DMS), a chemical released in areas of high marine productivity. However, dedicated behavioural studies still have to be conducted in baleen whales in order to confirm the involvement of chemoreception in their feeding ecology. We implemented 56 behavioural response experiments in humpback whales using two food-related chemical stimuli, krill extract and DMS, as well as their respective controls (orange clay and vegetable oil) in their breeding (Madagascar) and feeding grounds (Iceland and Antarctic Peninsula). The whales approached the stimulus area and stayed longer in the trial zone during krill extract trials compared to control trials, suggesting that they were attracted to the chemical source and spent time exploring its surroundings, probably in search of prey. This response was observed in Iceland, and to a lesser extend in Madagascar, but not in Antarctica. Surface behaviours indicative of sensory exploration, such as diving under the stimulus area and stopping navigation, were also observed more often during krill extract trials than during control trials. Exposure to DMS did not elicit such exploration behaviours in any of the study areas. However, acoustic analyses suggest that DMS and krill extract both modified the whales' acoustic activity in Madagascar. Altogether, these results provide the first behavioural evidence that baleen whales actually perceive prey-derived chemical cues over distances of several hundred metres. Chemoreception, especially olfaction, could thus be used for locating prey aggregations and for navigation at sea, as it has been shown in other marine predators including seabirds.

The echolocation transmission beam of free-ranging Indo-Pacific humpback dolphins (Sousa chinensis).

While the transmission beam of odontocetes has been described in a number of studies, the majority of them that have measured the transmission beam in two dimensions were focused on captive animals. Within the current study, a dedicated cross hydrophone array with nine elements was used to investigate the echolocation transmission beam of free-ranging Indo-Pacific humpback dolphins. A total of 265 on-axis clicks were analyzed, from which the apparent peak to peak source levels ranged between 168 to 207 dB (mean 184.5 dB ± 6.6 dB). The 3-dB beam width along the horizontal and vertical plane was 9.6° and 7.4°, respectively. Measured separately, the directivity index of the horizontal and vertical plane was 12.6 and 13.5 dB, respectively, and the overall directivity index (both planes combined) was 29.5 dB. The beam shape was slightly asymmetrical along the horizontal and vertical axis. Compared to other species, the characteristics of the transmitting beam of Indo-Pacific humpback dolphins were relatively close to the bottlenose dolphin (Tursiops truncatus), likely due to the similarity in the peak frequency and waveform of echolocation clicks and comparable body sizes of the two species.

A blind source separation approach for humpback whale song separation.

Many marine mammal species are highly social and are frequently encountered in groups or aggregations. When conducting passive acoustic monitoring in such circumstances, recordings commonly contain vocalizations of multiple individuals which overlap in time and frequency. This paper considers the use of blind source separation as a method for processing these recordings to separate the calls of individuals. The example problem considered here is that of the songs of humpback whales. The high levels of noise and long impulse responses can make source separation in underwater contexts a challenging proposition. The approach present here is based on time-frequency masking, allied to a noise reduction process. The technique is assessed using simulated and measured data sets, and the results demonstrate the effectiveness of the method for separating humpback whale songs.

Potential motivational information encoded within humpback whale non-song vocal sounds.

Acoustic signals in terrestrial animals follow motivational-structural rules to inform receivers of the signaler's motivational state, valence and level of arousal. Low-frequency "harsh" signals are produced in aggressive contexts, whereas high-frequency tonal sounds are produced in fearful/appeasement contexts. Using the non-song social call catalogue of humpback whales (Megaptera novaeangliae), this study tested for potential motivational-structural rules within the call catalogue of a baleen whale species. A total of 32 groups within different social contexts (ranging from stable, low arousal groups, such as a female with her calf, to affiliating, higher arousal, groups containing multiple males competing for access to the central female) were visually and acoustically tracked as they migrated southwards along the eastern coast of Australia. Social calls separated into four main cluster types, with signal structures in two categories consistent with "aggressive" signals and, "fearful/appeasement" signals in terrestrial animals. The group's use of signals within these clusters matched their context in that presumed low arousal non-affiliating groups almost exclusively used "low-arousal" signals (a cluster of low frequency unmodulated or upsweep sounds). Affiliating groups used a higher proportion of an intermediate cluster of signal types deemed "higher arousal" signals and groups containing three or more adults used a higher proportion of "aggressive" signal types.

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.

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 µPa in the 36 Hz-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 ∼60 dB above the noise level.

Understanding the intentional acoustic behavior of humpback whales: a production-based approach.

Following a production-based approach, this paper deals with the acoustic behavior of humpback whales. This approach investigates various physical factors, which are either internal (e.g., physiological mechanisms) or external (e.g., environmental constraints) to the respiratory tractus of the whale, for their implications in sound production. This paper aims to describe a functional scenario of this tractus for the generation of vocal sounds. To do so, a division of this tractus into three different configurations is proposed, based on the air recirculation process which determines air sources and laryngeal valves. Then, assuming a vocal function (in sound generation or modification) for several specific anatomical components, an acoustic characterization of each of these configurations is proposed to link different spectral features, namely, fundamental frequencies and formant structures, to specific vocal production mechanisms. A discussion around the question of whether the whale is able to fully exploit the acoustic potential of its respiratory tractus is eventually provided.

Site specific probability of passive acoustic detection of humpback whale calls from single fixed hydrophones.

Passive acoustic monitoring of marine mammal calls is an increasingly important method for assessing population numbers, distribution, and behavior. A common mistake in the analysis of marine mammal acoustic data is formulating conclusions about these animals without first understanding how environmental properties such as bathymetry, sediment properties, water column sound speed, and ocean acoustic noise influence the detection and character of vocalizations in the acoustic data. The approach in this paper is to use Monte Carlo simulations with a full wave field acoustic propagation model to characterize the site specific probability of detection of six types of humpback whale calls at three passive acoustic monitoring locations off the California coast. Results show that the probability of detection can vary by factors greater than ten when comparing detections across locations, or comparing detections at the same location over time, due to environmental effects. Effects of uncertainties in the inputs to the propagation model are also quantified, and the model accuracy is assessed by comparing calling statistics amassed from 24,690 humpback units recorded in the month of October 2008. Under certain conditions, the probability of detection can be estimated with uncertainties sufficiently small to allow for accurate density estimates.

Network-based diffusion analysis reveals cultural transmission of lobtail feeding in humpback whales.

We used network-based diffusion analysis to reveal the cultural spread of a naturally occurring foraging innovation, lobtail feeding, through a population of humpback whales (Megaptera novaeangliae) over a period of 27 years. Support for models with a social transmission component was 6 to 23 orders of magnitude greater than for models without. The spatial and temporal distribution of sand lance, a prey species, was also important in predicting the rate of acquisition. Our results, coupled with existing knowledge about song traditions, show that this species can maintain multiple independently evolving traditions in its populations. These insights strengthen the case that cetaceans represent a peak in the evolution of nonhuman culture, independent of the primate lineage.

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.

Turismo de observação de cetáceos como ferramenta no estudo do comportamento de baleias Jubarte (Megaptera novaeangliae) / Whale-watching as a tool to study humpback whales (Megaptera novaeangliae) behavior

O comportamento das baleias jubarte foi observado para detectar diferenças nas probabilidades de ocorrência de determinadas categorias comportamentais observadas a partir de cruzeiros de pesquisa e de turismo. Foram analisados dados de 256 grupos observados durante os cruzeiros de pesquisa e 122 grupos em cruzeiros de turismo, nas temporadas de 2001 a 2003, no litoral norte da Bahia. A análise de rede bayesiana forneceu maiores probabilidades de ocorrência para as categorias do tipo deslocamento, socialização e agressivo, enquanto a análise comparativa não forneceu diferenças expressivas entre as duas categorias de cruzeiro. De acordo com as probabilidades de ocorrência da conduta repouso para grupos com filhote e devido a uma possível interrupção do ato de amamentação causada pelas embarcações, sugere-se uma distância mais conservativa e um menor tempo de permanência com esses grupos(AU)

Humpback whales behavior was observed to determine the existence of differences between observations occurring from research vessels vs. whale-watching vessels. A total of 256 groups were observed from research vessels and 122 groups from whale-watching vessels during the breeding seasons from 2001 through 2003 in the northern coast of the state of Bahia, Brazil. A Bayesian network analysis yielded greatest probabilities of occurrence of the behavioral conducts traveling, socialization and aggressive. Significant behavioral differences between cruise types were not obtained through comparative analysis. Given the probability of occurrence of resting conducts in calf pods and the possibility of the interruption of suckling caused by vessels, it is suggested that a more conservative distance and shorter visits be adopted by whale-watching groups(AU)

Turismo de observação de cetáceos como ferramenta no estudo do comportamento de baleias Jubarte (Megaptera novaeangliae) / Whale-watching as a tool to study humpback whales (Megaptera novaeangliae) behavior

O comportamento das baleias jubarte foi observado para detectar diferenças nas probabilidades de ocorrência de determinadas categorias comportamentais observadas a partir de cruzeiros de pesquisa e de turismo. Foram analisados dados de 256 grupos observados durante os cruzeiros de pesquisa e 122 grupos em cruzeiros de turismo, nas temporadas de 2001 a 2003, no litoral norte da Bahia. A análise de rede bayesiana forneceu maiores probabilidades de ocorrência para as categorias do tipo deslocamento, socialização e agressivo, enquanto a análise comparativa não forneceu diferenças expressivas entre as duas categorias de cruzeiro. De acordo com as probabilidades de ocorrência da conduta repouso para grupos com filhote e devido a uma possível interrupção do ato de amamentação causada pelas embarcações, sugere-se uma distância mais conservativa e um menor tempo de permanência com esses grupos.

Humpback whales behavior was observed to determine the existence of differences between observations occurring from research vessels vs. whale-watching vessels. A total of 256 groups were observed from research vessels and 122 groups from whale-watching vessels during the breeding seasons from 2001 through 2003 in the northern coast of the state of Bahia, Brazil. A Bayesian network analysis yielded greatest probabilities of occurrence of the behavioral conducts traveling, socialization and aggressive. Significant behavioral differences between cruise types were not obtained through comparative analysis. Given the probability of occurrence of resting conducts in calf pods and the possibility of the interruption of suckling caused by vessels, it is suggested that a more conservative distance and shorter visits be adopted by whale-watching groups.