Koen de Reus


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  • Henry, M. J., Cook, P. F., de Reus, K., Nityananda, V., Rouse, A. A., & Kotz, S. A. (2021). An ecological approach to measuring synchronization abilities across the animal kingdom. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 376: 20200336. doi:10.1098/rstb.2020.0336.


    In this perspective paper, we focus on the study of synchronization abilities across the animal kingdom. We propose an ecological approach to studying nonhuman animal synchronization that begins from observations about when, how and why an animal might synchronize spontaneously with natural environmental rhythms. We discuss what we consider to be the most important, but thus far largely understudied, temporal, physical, perceptual and motivational constraints that must be taken into account when designing experiments to test synchronization in nonhuman animals. First and foremost, different species are likely to be sensitive to and therefore capable of synchronizing at different timescales. We also argue that it is fruitful to consider the latent flexibility of animal synchronization. Finally, we discuss the importance of an animal's motivational state for showcasing synchronization abilities. We demonstrate that the likelihood that an animal can successfully synchronize with an environmental rhythm is context-dependent and suggest that the list of species capable of synchronization is likely to grow when tested with ecologically honest, species-tuned experiments.
  • de Reus, K., Soma, M., Anichini, M., Gamba, M., de Heer Kloots, M., Lense, M., Bruno, J. H., Trainor, L., & Ravignani, A. (2021). Rhythm in dyadic interactions. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 376: 20200337. doi:10.1098/rstb.2020.0337.


    This review paper discusses rhythmic dyadic interactions in social and sexual contexts. We report rhythmic interactions during communication within dyads, as found in humans, non-human primates, non-primate mammals, birds, anurans and insects. Based on the patterns observed, we infer adaptive explanations for the observed rhythm interactions and identify knowledge gaps. Across species, the social environment during ontogeny is a key factor in shaping adult signal repertoires and timing mechanisms used to regulate interactions. The degree of temporal coordination is influenced by the dynamic and strength of the dyadic interaction. Most studies of temporal structure in interactive signals mainly focus on one modality (acoustic and visual); we suggest more work should be performed on multimodal signals. Multidisciplinary approaches combining cognitive science, ethology and ecology should shed more light on the exact timing mechanisms involved. Taken together, rhythmic signalling behaviours are widespread and critical in regulating social interactions across taxa.
  • De Heer Kloots, M., Carlson, D., Garcia, M., Kotz, S., Lowry, A., Poli-Nardi, L., de Reus, K., Rubio-García, A., Sroka, M., Varola, M., & Ravignani, A. (2020). Rhythmic perception, production and interactivity in harbour and grey seals. In A. Ravignani, C. Barbieri, M. Flaherty, Y. Jadoul, E. Lattenkamp, H. Little, M. Martins, K. Mudd, & T. Verhoef (Eds.), The Evolution of Language: Proceedings of the 13th International Conference (Evolang13) (pp. 59-62). Nijmegen: The Evolution of Language Conferences.
  • de Reus, K., Carlson, D., Jadoul, Y., Lowry, A., Gross, S., Garcia, M., Salazar Casals, A., Rubio-García, A., Haas, C. E., De Boer, B., & Ravignani, A. (2020). Relationships between vocal ontogeny and vocal tract anatomy in harbour seals (Phoca vitulina). In A. Ravignani, C. Barbieri, M. Flaherty, Y. Jadoul, E. Lattenkamp, H. Little, M. Martins, K. Mudd, & T. Verhoef (Eds.), The Evolution of Language: Proceedings of the 13th International Conference (Evolang13) (pp. 63-66). Nijmegen: The Evolution of Language Conferences.
  • Ravignani, A., & de Reus, K. (2019). Modelling animal interactive rhythms in communication. Evolutionary Bioinformatics, 15, 1-14. doi:10.1177/1176934318823558.


    Time is one crucial dimension conveying information in animal communication. Evolution has shaped animals’ nervous systems to produce signals with temporal properties fitting their socio-ecological niches. Many quantitative models of mechanisms underlying rhythmic behaviour exist, spanning insects, crustaceans, birds, amphibians, and mammals. However, these computational and mathematical models are often presented in isolation. Here, we provide an overview of the main mathematical models employed in the study of animal rhythmic communication among conspecifics. After presenting basic definitions and mathematical formalisms, we discuss each individual model. These computational models are then compared using simulated data to uncover similarities and key differences in the underlying mechanisms found across species. Our review of the empirical literature is admittedly limited. We stress the need of using comparative computer simulations – both before and after animal experiments – to better understand animal timing in interaction. We hope this article will serve as a potential first step towards a common computational framework to describe temporal interactions in animals, including humans.

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  • Ravignani, A., Kello, C. T., de Reus, K., Kotz, S. A., Dalla Bella, S., Mendez-Arostegui, M., Rapado-Tamarit, B., Rubio-Garcia, A., & de Boer, B. (2019). Ontogeny of vocal rhythms in harbor seal pups: An exploratory study. Current Zoology, 65(1), 107-120. doi:10.1093/cz/zoy055.


    Puppyhood is a very active social and vocal period in a harbor seal's life Phoca vitulina. An important feature of vocalizations is their temporal and rhythmic structure, and understanding vocal timing and rhythms in harbor seals is critical to a cross-species hypothesis in evolutionary neuroscience that links vocal learning, rhythm perception, and synchronization. This study utilized analytical techniques that may best capture rhythmic structure in pup vocalizations with the goal of examining whether (1) harbor seal pups show rhythmic structure in their calls and (2) rhythms evolve over time. Calls of 3 wild-born seal pups were recorded daily over the course of 1-3 weeks; 3 temporal features were analyzed using 3 complementary techniques. We identified temporal and rhythmic structure in pup calls across different time windows. The calls of harbor seal pups exhibit some degree of temporal and rhythmic organization, which evolves over puppyhood and resembles that of other species' interactive communication. We suggest next steps for investigating call structure in harbor seal pups and propose comparative hypotheses to test in other pinniped species.
  • Ravignani, A., Garcia, M., Gross, S., de Reus, K., Hoeksema, N., Rubio-Garcia, A., & de Boer, B. (2018). Pinnipeds have something to say about speech and rhythm. In C. Cuskley, M. Flaherty, H. Little, L. McCrohon, A. Ravignani, & T. Verhoef (Eds.), Proceedings of the 12th International Conference on the Evolution of Language (EVOLANG XII) (pp. 399-401). Toruń, Poland: NCU Press. doi:10.12775/3991-1.095.

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