Animal acoustic communication often takes the form of complex sequences, made up of multiple distinct acoustic units. Apart from the well-known example of birdsong, other animals such as insects, amphibians, and mammals (including bats, rodents, primates, and cetaceans) also generate complex acoustic sequences. Occasionally, such as with birdsong, the adaptive role of these sequences seems clear (e.g. mate attraction and territorial defence). More often however, researchers have only begun to characterise – let alone understand – the significance and meaning of acoustic sequences. Hypotheses abound, but there is little agreement as to how sequences should be defined and analysed. Our review aims to outline suitable methods for testing these hypotheses, and to describe the major limitations to our current and near-future knowledge on questions of acoustic sequences. This review and prospectus is the result of a collaborative effort between 43 scientists from the fields of animal behaviour, ecology and evolution, signal processing, machine learning, quantitative linguistics, and information theory, who gathered for a 2013 workshop entitled, “Analysing vocal sequences in animals”. Our goal is to present not just a review of the state of the art, but to propose a methodological framework that summarises what we suggest are the best practices for research in this field, across taxa and across disciplines. We also provide a tutorial-style introduction to some of the most promising algorithmic approaches for analysing sequences. We divide our review into three sections: identifying the distinct units of an acoustic sequence, describing the different ways that information can be contained within a sequence, and analysing the structure of that sequence. Each of these sections is further subdivided to address the key questions and approaches in that area. We propose a uniform, systematic, and comprehensive approach to studying sequences, with the goal of clarifying research terms used in different fields, and facilitating collaboration and comparative studies. Allowing greater interdisciplinary collaboration will facilitate the investigation of many important questions in the evolution of communication and sociality.

Animals often use acoustic signals to communicate in groups or social aggregations in which multiple individuals signal within a receiver's hearing range. Consequently, receivers face challenges related to acoustic interference and auditory masking that are not unlike the human "cocktail party problem," which refers to the problem of perceiving speech in noisy social settings. Understanding the sensory solutions to the cocktail party problem has been a goal of research on human hearing and speech communication for several decades. Despite a general interest in acoustic signaling in groups, animal behaviorists have devoted comparatively less attention toward understanding how animals solve problems equivalent to the human cocktail party problem. After illustrating how humans and nonhuman animals experience and overcome similar perceptual challenges in cocktail-party-like social environments, this article reviews previous psychophysical and physiological studies of humans and non-human animals to describe how the cocktail party problem can be solved. This review also outlines several basic and applied benefits that could result from studies of the cocktail party problem in the context of animal acoustic communication.In many animals, acoustic communication occurs in large groups or aggregations of signaling

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Auditory stream segregation refers to the perceptual grouping of sounds, to form coherent representations of objects in the acoustic scene, and is a fundamental aspect of hearing and speech perception. The perceptual segregation of simple interleaved tone sequences has been studied in humans and European starlings (Sturnus vulgaris) using sequences of 2 alternating tones differing in frequency (ABA-ABA-ABA-...). The segregation of A and B tones into separate auditory streams is believed to be promoted by preattentive auditory processes that increase the separation of excitation patterns along a tonotopic gradient. We tested the hypothesis that frequency selectivity and forward masking operate as 2 preattentive processes in sequential stream segregation by recording neural responses in the auditory forebrain of awake starlings to repeated ABA- sequences in which we varied the frequency separation (DeltaF) between the A and B tones and the tone repetition time (TRT). The A tones were presented at the neurons' characteristic frequency (CF), and B tones differed from the CF over a one-octave range. Larger DeltaF values and shorter TRTs promote the perceptual segregation of alternating tone sequences in humans and also resulted in larger differences in neural responses to alternating CF (A) and non-CF (B) tones. Our results are consistent with the hypothesis that preattentive auditory processes, such as frequency selectivity and forward masking, contribute to the perceptual segregation of sequential acoustic events having different frequencies into separate auditory streams, but also suggest that additional processes may be required to account for all known perceptual effects related to sequential auditory stream segregation.

2000: Individual variation in advertisement calls of territorial male green frogs, Rana clamitans: implications for individual discrimination. Ethology 107, 65Ð84. AbstractIndividuals of many territorial species discriminate between familiar territorial neighbors and unfamiliar strangers based on individual dierences in acoustic signals. Many anuran amphibians are territorial and use primarily acoustic communication during social interactions, but evidence for acoustically mediated individual discrimination is available only for one species. As a ®rst step in research designed to investigate individual discrimination in a second species of territorial frog, we examined patterns of within-male and among-male variability in 198 advertisement calls of 20 male green frogs, Rana clamitans. The aim was to determine which acoustic properties could be used by males to recognize their territorial neighbors and to estimate limits of reliability aorded by these properties for identifying dierent neighbors. All of the call properties that we examined exhibited signi®cant inter-individual variation. Discriminant function analyses assigned between 52% and 100% of calls to the correct individual, depending on sample size and the call properties included in the model. This suggests that there is sucient among-male variability to statistically identify individuals by their advertisement calls. The call properties of fundamental frequency and dominant frequency contributed the most towards discrimination among individuals. Based on their natural history and behavior and the results reported here, we suggest that male green frogs likely discriminate between strangers and adjacently territorial neighbors based on individual variation in advertisement calls.

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Anuran amphibians are superb animal models for investigating the mechanisms underlying acoustic signal perception amid high levels of background noise generated by large social aggregations of vocalizing individuals. Yet there are not well-established methods for quantifying a number of key measures of auditory perception in frogs, in part, because frogs are notoriously difficult subjects for traditional psychoacoustic experiments based on classical or operant conditioning. A common experimental approach for studying frog hearing and acoustic communication involves behavioral phonotaxis experiments, in which patterns of movement directed toward sound sources indicate the subjects' perceptual experiences. In this study, three different phonotaxis experiments were conducted using the same target signals and noise maskers to compare different experimental methods and analytical tools for deriving estimates of signal recognition thresholds in the presence or absence of "chorus-shaped noise" ͑i.e., artificial noise with a spectrum similar to that of real breeding choruses͒. Estimates of recognition thresholds based on measures of angular orientation, response probabilities, and response latencies were quite similar in both two-choice and no-choice phonotaxis tests. These results establish important baselines for comparing different methods of estimating signal recognition thresholds in frogs tested in various masking noise conditions.