This study measured the degree of behavioral responses in blue whales (Balaenoptera musculus) to controlled noise exposure off the southern California coast. High-resolution movement and passive acoustic data were obtained from non-invasive archival tags (n=42) whereas surface positions were obtained with visual focal follows. Controlled exposure experiments (CEEs) were used to obtain direct behavioral measurements before, during and after simulated and operational military mid-frequency active sonar (MFAS), pseudorandom noise (PRN) and controls (no noise exposure). For a subset of deep-feeding animals (n=21), active acoustic measurements of prey were obtained and used as contextual covariates in response analyses. To investigate potential behavioral changes within individuals as a function of controlled noise exposure conditions, two parallel analyses of timeseries data for selected behavioral parameters (e.g. diving, horizontal movement and feeding) were conducted. This included expert scoring of responses according to a specified behavioral severity rating paradigm and quantitative change-point analyses using Mahalanobis distance statistics. Both methods identified clear changes in some conditions. More than 50% of blue whales in deep-feeding states responded during CEEs, whereas no changes in behavior were identified in shallow-feeding blue whales. Overall, responses were generally brief, of low to moderate severity, and highly dependent on exposure context such as behavioral state, source-to-whale horizontal range and prey availability. Response probability did not follow a simple exposure-response model based on received exposure level. These results, in combination with additional analytical methods to investigate different aspects of potential responses within and among individuals, provide a comprehensive evaluation of how free-ranging blue whales responded to mid-frequency military sonar.

No abstract

The evolutionary origin of rhythm perception, a cognitive ability essential to musicality, remains unresolved [1-5]. The ability to perceive and memorize rhythmic sounds is widely shared among humans [6] but seems rare among other mammals [7, 8]. Although the perception of temporal metrical patterns has been found in a few species, this ability has only been demonstrated through behavioral training [9] (but see [10] for an example of spontaneous tempo coordination in a bonobo), and there is no experimental evidence to indicate its biological function. Furthermore, there is no example of a non-human mammal able to remember and recognize auditory rhythmic patterns among a wide range of tempi. In the northern elephant seal Mirounga angustirostris, the calls of mature males comprise a rhythmic series of pulses, with the call of each individual characterized by its tempo and timbre; these individual vocal signatures are stable over years and across contexts [11]. Here, we report that northern elephant seal males routinely memorize and recognize the unique tempo and timbre of their rivals' voices and use this rhythmic information to individually identify competitors, which facilitates navigation within the social network of the rookery. By performing playbacks with natural and modified vocalizations, we show that males are sensitive to call rhythm disruption independently of modification of spectral features and that they use both temporal and spectral cues to identify familiar rivals. While spectral features of calls typically encode individual identity in mammalian vocalizations [12], this is the first example of this phenomenon involving sound rhythm.

Specialized signals emitted by competing males often convey honest information about fighting ability. It is generally believed that receivers use these signals to directly assess their opponents. Here, we demonstrate an alternative communication strategy used by males in a breeding system where the costs of conflict are extreme. We evaluated the acoustic displays of breeding male northern elephant seals (Mirounga angustirostris), and found that social knowledge gained through prior experience with signallers was sufficient to maintain structured dominance relationships. Using sound analysis and playback experiments with both natural and modified signals, we determined that males do not rely on encoded information about size or dominance status, but rather learn to recognize individual acoustic signatures produced by their rivals. Further, we show that behavioural responses to competitors' calls are modulated by relative position in the hierarchy: the highest ranking (alpha) males defend their harems from all opponents, whereas mid-ranking (beta) males respond differentially to familiar challengers based on the outcome of previous competitive interactions. Our findings demonstrate that social knowledge of rivals alone can regulate dominance relationships among competing males within large, spatially dynamic social groups, and illustrate the importance of combining descriptive and experimental methods when deciphering the biological relevance of animal signals.