Acoustic sequences in non‐human animals: a tutorial review and prospectus
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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, ‘A nalysing 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.
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Standardising methodologies will avoid confusion and enable meaningful comparisons across species, populations, sexes, developmental stages, and behavioural contexts.
“Standardising methodologies will avoid confusion and enable meaningful comparisons across species, populations, sexes, developmental stages, and behavioural contexts (Broughton 1963; Ruxton and Schaefer 2011; Kershenbaum et al. 2016).”
From Roars, Rumbles, and Resonance: A Systematic Review and Meta‐Analysis of Crocodylian Acoustic Signals (2026) · cites this paper as Kershenbaum et al. 2016
The Abstract (page unavailable) proposes a uniform approach to clarify terminology and facilitate comparative studies across taxa, supporting the claim’s general point. The Introduction likewise describes bridging inconsistent terminology across disciplines and taxa. The paper does not specifically say that standardisation will enable comparisons across populations, sexes, developmental stages, and behavioural contexts. (LLM, conf=0.82)
Experimental — accuracy not yet measuredDifferences in acoustic signalling syntax may allow species identification and potentially contain contextual, individual and/or behavioural information.
“Differences in acoustic signalling syntax (i.e., the sequence of signal elements) may allow species identification and potentially contain contextual, individual and/or behavioural information (Garrick et al. 1978; Kershenbaum et al. 2016).”
From Roars, Rumbles, and Resonance: A Systematic Review and Meta‐Analysis of Crocodylian Acoustic Signals (2026) · cites this paper as Kershenbaum et al. 2016
Section VI(3), “How do individual differences in acoustic sequences arise?”, says variation in syntax may reflect aggression and that phrase sequences can distinguish individual birds. Section IV(7) describes different call sequences carrying different meanings, supporting the possibility of contextual or behavioural information. Section VI(4) discusses a possible role for sequence syntax in speciation, but does not establish that syntactic differences allow species identification. (LLM, conf=0.82)
Experimental — accuracy not yet measuredConceptual work on sequence structure, mammalian source-filter theory, and signal embedding in social networks provides a theoretical foundation for interpreting meaning in animal vocalizations.
“Conceptual work on sequence structure (Kershenbaum et al. 2016), mammalian source–filter theory (Taylor and Reby 2010), and signal embedding in social networks (Seyfarth and Cheney 2017; Snijders and Naguib 2017) provide a theoretical foundation.”
From Listening forward: emerging roles of bioacoustics in ecology, evolution, and conservation (2026) · cites this paper as Kershenbaum et al. 2016
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- Authors
- Kershenbaum, A., Blumstein, D. T., Roch, M. A., Akçay, Ç., Backus, G., Bee, M. A., Bohn, K., Cao, Y., Carter, G., Cäsar, C., Coen, M., DeRuiter, S. L., Doyle, L., Edelman, S., Ferrer‐i‐Cancho, R., Freeberg, T. M., Garland, E. C., Gustison, M., Harley, H. E., Huetz, C., Hughes, M., Hyland Bruno, J., Ilany, A., Jin, D. Z., Johnson, M., Ju, C., Karnowski, J., Lohr, B., Manser, M. B., McCowan, B., Mercado, E., Narins, P. M., Piel, A., Rice, M., Salmi, R., Sasahara, K., Sayigh, L., Shiu, Y., Taylor, C., Vallejo, E. E., Waller, S., & Zamora‐Gutierrez, V.
- Affiliations
- National Institute for Mathematical and Biological Synthesis (US)
- University of California, Los Angeles (US)
- San Diego State University (US)
- Cornell University (US)
- North Carolina State University (US)
- University of Minnesota (US)
- Florida International University (US)
- The University of Texas at Dallas (US)
- University of Maryland, College Park (US)
- University of St Andrews (GB)
- University of Wisconsin–Madison (US)
- Search for Extraterrestrial Intelligence (US)
- Universitat Politècnica de Catalunya (ES)
- University of Tennessee System (US)
- National Oceanic and Atmospheric Administration (US)
- University of Michigan (US)
- New College of Florida (US)
- Université Paris-Sud (FR)
- College of Charleston (US)
- Department of Psychology Hunter College and the Graduate Center, The City University of New York 365 Fifth Avenue New York NY 10016 U.S.A.
- Pennsylvania State University (US)
- Marquette University (US)
- City University of New York (US)
- University of California San Diego (US)
- University of Maryland, Baltimore County (US)
- University of Zurich (CH)
- University of California, Davis (US)
- University at Buffalo, State University of New York (US)
- Division of Biological Anthropology University of Cambridge Pembroke Street Cambridge CB2 3QG U.K.
- California State University, San Marcos (US)
- University of Georgia (US)
- Nagoya University (JP)
- Woods Hole Oceanographic Institution (US)
- Tecnológico de Monterrey (MX)
- Montana State University (US)
- University of Cambridge (GB)
- Corresponding author
- Kershenbaum, Arik
- ORCID
- Kershenbaum, Arik
- Blumstein, Daniel T.
- Roch, Marie A.
- Akçay, Çağlar
- Backus, Gregory
- Bee, Mark A.
- Bohn, Kirsten
- Carter, Gerald
- Cäsar, Cristiane
- DeRuiter, Stacy L.
- Edelman, Shimon
- Ferrer‐i‐Cancho, Ramon
- Freeberg, Todd M.
- Garland, Ellen C.
- Gustison, Morgan
- Huetz, Chloé
- Hughes, Melissa
- Ilany, Amiyaal
- Jin, Dezhe Z.
- Johnson, Michael
- Manser, Marta B.
- McCowan, Brenda
- Mercado, Eduardo
- Narins, Peter M.
- Salmi, Roberta
- Sasahara, Kazutoshi
- Sayigh, Laela
- Shiu, Yu
- Taylor, Charles
- Vallejo, Edgar E.
- Waller, Sara
- Zamora‐Gutierrez, Veronica
- Venue
- Biological Reviews
- Publisher
- Wiley
- Published
- 2014-11-26
- Volume
- 91
- Issue
- 1
- Pages
- 13-52
- ISSN
- 0006-3231 (ISSN-L)
- 1464-7931
- 1469-185X
- PMC
- PMC4444413
- OpenAlex
- W1514601846
- Cited by
- 337
- References
- 424
- Citations by year
- 2026: 27 · 2025: 33 · 2024: 31 · 2023: 34 · 2022: 46 · 2021: 36 · 2020: 36 · 2019: 16 · 2018: 27 · 2017: 26 · 2016: 18 · 2015: 6 · 2014: 1
- FWCI
- 2.73
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Biological reviews/Biological reviews of the Cambridge Philosophical Society (publishedVersion)
- PubMed (publishedVersion)
- Liverpool John Moores University (acceptedVersion)
- Europe PMC (PubMed Central) (acceptedVersion)
- PubMed Central (acceptedVersion)
- LA Referencia (Red Federada de Repositorios Institucionales de Publicaciones Científicas) (acceptedVersion, other-oa)
- UPCommons institutional repository (Universitat Politècnica de Catalunya) (acceptedVersion, other-oa)
- Apollo (University of Cambridge) (acceptedVersion)
- Deep Blue (University of Michigan) (submittedVersion)
- e-Publications@Marquette (Marquette University) (submittedVersion)
- HAL (submittedVersion)
- ePrints Soton (University of Southampton) (submittedVersion)
- eScholarship (California Digital Library) (submittedVersion)
- RECERCAT (Consorci de Serveis Universitaris de Catalunya) (submittedVersion, public-domain)
- St Andrews Research Repository (St Andrews Research Repository) (submittedVersion)
- Tokyo Tech Research Repository (Tokyo Institute of Technology) (submittedVersion)
- Funders
- NSF (DEB-1119660, IOS-0842759)
- NIH (R01DC009582)
- ONR (N0001411IP20086)
- NOPP (ONR/BOEM) (N00014-11-1-0697)
- U.S. Office of Naval Research
- Spanish Ministry of Science and Innovation (BASMATI (TIN2011-27479-C04-03))
- National Research Council Postdoctoral Fellowship
- CONACYT, Mexico (I010/214/2012)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Developmental Biology › Animal Vocal Communication and Behavior
- Physical Sciences › Environmental Science › Ecology › Marine animal studies overview
- Social Sciences › Cultural Studies › Language and cultural evolution
- Keywords
- animal vocal communication, acoustic unit, sequence structure, bird song, evolution of communication, sociality, interdisciplinary research framework
- MeSH terms
- Acoustics, Animals, Markov Chains, Models, Biological, Perception, Vocalization, Animal
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