Comparison of methods for rhythm analysis of complex animals’ acoustic signals

Analyzing the rhythm of animals’ acoustic signals is of interest to a growing number of researchers: evolutionary biologists want to disentangle how these structures evolved and what patterns can be found, and ecologists and conservation biologists aim to discriminate cryptic species on the basis of parameters of acoustic signals such as temporal structures. Temporal structures are also relevant for research on vocal production learning, a part of which is for the animal to learn a temporal structure. These structures, in other words, these rhythms, are the topic of this paper. How can they be investigated in a meaningful, comparable and universal way? Several approaches exist. Here we used five methods to compare their suitability and interpretability for different questions and datasets and test how they support the reproducibility of results and bypass biases. Three very different datasets with regards to recording situation, length and context were analyzed: two social vocalizations of Neotropical bats (multisyllabic, medium long isolation calls of Saccopteryx bilineata, and monosyllabic, very short isolation calls of Carollia perspicillata) and click trains of sperm whales, Physeter macrocephalus. Techniques to be compared included Fourier analysis with a newly developed goodness-of-fit value, a generate-and-test approach where data was overlaid with varying artificial beats, and the analysis of inter-onset-intervals and calculations of a normalized Pairwise Variability Index (nPVI). We discuss the advantages and disadvantages of the methods and we also show suggestions on how to best visualize rhythm analysis results. Furthermore, we developed a decision tree that will enable researchers to select a suitable and comparable method on the basis of their data.Author summary: In the analysis of animal communication more and more interest is shown in rhythm of animal communication and what information this might convey. In this paper, we establish a workflow to analyze the temporal structure–namely the rhythm–of any particular animals’acoustic signal with methods that are applicable for a wide range of signals and results that are easily comparable and interpretable. This workflow will enhance the understanding of rhythmicality in animals’ acoustic signals as well as facilitate comparison between species. Methods we conducted ranged from simple distributional and visual analysis to higher mathematics such as Fourier analysis. All analyses rely on Inter-Onset-Intervals, the duration between the beginning of one element and the next. We used different datasets from two neotropical bat species as well as from the sperm whale. With this selection, we cover very short sequences with only few elements up to sequences of around 200 elements, multisyllabic and monosyllabic sequences and social communication as well as sounds used for orientation and foraging.