Displaying 1 - 27 of 27
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Anichini, M., De Heer Kloots, M., & Ravignani, A. (2020). Interactive rhythms in the wild, in the brain, and in silico. Canadian Journal of Experimental Psychology, 74(3), 170-175. doi:10.1037/cep0000224.
Abstract
There are some historical divisions in methods, rationales, and purposes between
studies on comparative cognition and behavioural ecology. In turn, the interaction between
these two branches and studies from mathematics, computation and neuroscience is not
usual. In this short piece, we attempt to build bridges among these disciplines. We present a
series of interconnected vignettes meant to illustrate how a more interdisciplinary approach
looks like when successful, and its advantages. Concretely, we focus on a recent topic,
namely animal rhythms in interaction, studied under different approaches. We showcase 5
research efforts, which we believe successfully link 5 particular Scientific areas of rhythm
research conceptualized as: Social neuroscience, Detailed rhythmic quantification,
Ontogeny, Computational approaches and Spontaneous interactions. Our suggestions will
hopefully spur a ‘Comparative rhythms in interaction’ field, which can integrate and
capitalize on knowledge from zoology, comparative psychology, neuroscience, and
computation. -
De Boer, B., Thompson, B., Ravignani, A., & Boeckx, C. (2020). Analysis of mutation and fixation for language. 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. 56-58). Nijmegen: The Evolution of Language Conferences. -
De Boer, B., Thompson, B., Ravignani, A., & Boeckx, C. (2020). Evolutionary dynamics do not motivate a single-mutant theory of human language. Scientific Reports, 10: 451. doi:10.1038/s41598-019-57235-8.
Abstract
One of the most controversial hypotheses in cognitive science is the Chomskyan evolutionary conjecture that language arose instantaneously in humans through a single mutation. Here we analyze the evolutionary dynamics implied by this hypothesis, which has never been formalized before. The hypothesis supposes the emergence and fixation of a single mutant (capable of the syntactic operation Merge) during a narrow historical window as a result of frequency-independent selection under a huge fitness advantage in a population of an effective size no larger than ~15 000 individuals. We examine this proposal by combining diffusion analysis and extreme value theory to derive a probabilistic formulation of its dynamics. We find that although a macro-mutation is much more likely to go to fixation if it occurs, it is much more unlikely a priori than multiple mutations with smaller fitness effects. The most likely scenario is therefore one where a medium number of mutations with medium fitness effects accumulate. This precise analysis of the probability of mutations occurring and going to fixation has not been done previously in the context of the evolution of language. Our results cast doubt on any suggestion that evolutionary reasoning provides an independent rationale for a single-mutant theory of language.Additional information
Supplementary material -
Garcia, M., & Ravignani, A. (2020). Acoustic allometry and vocal learning in mammals. Biology Letters, 16: 20200081. doi:10.1098/rsbl.2020.0081.
Abstract
Acoustic allometry is the study of how animal vocalisations reflect their body size. A key aim of this research is to identify outliers to acoustic allometry principles and pinpoint the evolutionary origins of such outliers. A parallel strand of research investigates species capable of vocal learning, the experience-driven ability to produce novel vocal signals through imitation or modification of existing vocalisations. Modification of vocalizations is a common feature found when studying both acoustic allometry and vocal learning. Yet, these two fields have only been investigated separately to date. Here, we review and connect acoustic allometry and vocal learning across mammalian clades, combining perspectives from bioacoustics, anatomy and evolutionary biology. Based on this, we hypothesize that, as a precursor to vocal learning, some species might have evolved the capacity for volitional vocal modulation via sexual selection for ‘dishonest’ signalling. We provide preliminary support for our hypothesis by showing significant associations between allometric deviation and vocal learning in a dataset of 164 mammals. Our work offers a testable framework for future empirical research linking allometric principles with the evolution of vocal learning. -
Garcia, M., Theunissen, F., Sèbe, F., Clavel, J., Ravignani, A., Marin-Cudraz, T., Fuchs, J., & Mathevon, N. (2020). Evolution of communication signals and information during species radiation. Nature Communications, 11: 4970. doi:10.1038/s41467-020-18772-3.
Abstract
Communicating species identity is a key component of many animal signals. However, whether selection for species recognition systematically increases signal diversity during clade radiation remains debated. Here we show that in woodpecker drumming, a rhythmic signal used during mating and territorial defense, the amount of species identity information encoded remained stable during woodpeckers’ radiation. Acoustic analyses and evolutionary reconstructions show interchange among six main drumming types despite strong phylogenetic contingencies, suggesting evolutionary tinkering of drumming structure within a constrained acoustic space. Playback experiments and quantification of species discriminability demonstrate sufficient signal differentiation to support species recognition in local communities. Finally, we only find character displacement in the rare cases where sympatric species are also closely related. Overall, our results illustrate how historical contingencies and ecological interactions can promote conservatism in signals during a clade radiation without impairing the effectiveness of information transfer relevant to inter-specific discrimination. -
Geambasu, A., Toron, L., Ravignani, A., & Levelt, C. C. (2020). Rhythmic recursion? Human sensitivity to a Lindenmayer grammar with self-similar structure in a musical task. Music & Science. doi:10.1177%2F2059204320946615.
Abstract
Processing of recursion has been proposed as the foundation of human linguistic ability. Yet this ability may be shared with other domains, such as the musical or rhythmic domain. Lindenmayer grammars (L-systems) have been proposed as a recursive grammar for use in artificial grammar experiments to test recursive processing abilities, and previous work had shown that participants are able to learn such a grammar using linguistic stimuli (syllables). In the present work, we used two experimental paradigms (a yes/no task and a two-alternative forced choice) to test whether adult participants are able to learn a recursive Lindenmayer grammar composed of drum sounds. After a brief exposure phase, we found that participants at the group level were sensitive to the exposure grammar and capable of distinguishing the grammatical and ungrammatical test strings above chance level in both tasks. While we found evidence of participants’ sensitivity to a very complex L-system grammar in a non-linguistic, potentially musical domain, the results were not robust. We discuss the discrepancy within our results and with the previous literature using L-systems in the linguistic domain. Furthermore, we propose directions for future music cognition research using L-system grammars. -
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. -
Heinrich, T., Ravignani, A., & Hanke, F. H. (2020). Visual timing abilities of a harbour seal (Phoca vitulina) and a South African fur seal (Arctocephalus pusillus pusillus) for sub- and supra-second time intervals. Animal Cognition, 23(5), 851-859. doi:10.1007/s10071-020-01390-3.
Abstract
Timing is an essential parameter influencing many behaviours. A previous study demonstrated a high sensitivity of a phocid, the harbour seal (Phoca vitulina), in discriminating time intervals. In the present study, we compared the harbour seal’s timing abilities with the timing abilities of an otariid, the South African fur seal (Arctocephalus pusillus pusillus). This comparison seemed essential as phocids and otariids differ in many respects and might, thus, also differ regarding their timing abilities. We determined time difference thresholds for sub- and suprasecond time intervals marked by a white circle on a black background displayed for a specific time interval on a monitor using a staircase method. Contrary to our expectation, the timing abilities of the fur seal and the harbour seal were comparable. Over a broad range of time intervals, 0.8–7 s in the fur seal and 0.8–30 s in the harbour seal, the difference thresholds followed Weber’s law. In this range, both animals could discriminate time intervals differing only by 12 % and 14 % on average. Timing might, thus be a fundamental cue for pinnipeds in general to be used in various contexts, thereby complementing information provided by classical sensory systems. Future studies will help to clarify if timing is indeed involved in foraging decisions or the estimation of travel speed or distance.Additional information
supplementary material -
Hoeksema, N., Villanueva, S., Mengede, J., Salazar-Casals, A., Rubio-García, A., Curcic-Blake, B., Vernes, S. C., & Ravignani, A. (2020). Neuroanatomy of the grey seal brain: Bringing pinnipeds into the neurobiological study of vocal learning. 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. 162-164). Nijmegen: The Evolution of Language Conferences. -
Jacoby, N., Margulis, E. H., Clayton, M., Hannon, E., Honing, H., Iversen, J., Klein, T. R., Mehr, S. A., Pearson, L., Peretz, I., Perlman, M., Polak, R., Ravignani, A., Savage, P. E., Steingo, G., Stevens, C. J., Trainor, L., Trehub, S., Veal, M., & Wald-Fuhrmann, M. (2020). Cross-cultural work in music cognition: Challenges, insights, and recommendations. Music Perception, 37(3), 185-195. doi:10.1525/mp.2020.37.3.185.
Abstract
Many foundational questions in the psychology of music require cross-cultural approaches, yet the vast majority of work in the field to date has been conducted with Western participants and Western music. For cross-cultural research to thrive, it will require collaboration between people from different disciplinary backgrounds, as well as strategies for overcoming differences in assumptions, methods, and terminology. This position paper surveys the current state of the field and offers a number of concrete recommendations focused on issues involving ethics, empirical methods, and definitions of “music” and “culture.” -
Ravignani, A., & Kotz, S. (2020). Breathing, voice and synchronized movement. Proceedings of the National Academy of Sciences of the United States of America, 117(38), 23223-23224. doi:10.1073/pnas.2011402117.
Additional information
Pouw_etal_reply.pdf -
Ravignani, A., Barbieri, C., Flaherty, M., Jadoul, Y., Lattenkamp, E. Z., Little, H., Martins, M., Mudd, K., & Verhoef, T. (
Eds. ). (2020). The Evolution of Language: Proceedings of the 13th International Conference (Evolang13). Nijmegen: The Evolution of Language Conferences. doi:10.17617/2.3190925.Additional information
Link to pdf on EvoLang Website -
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. -
Wilson, B., Spierings, M., Ravignani, A., Mueller, J. L., Mintz, T. H., Wijnen, F., Van der Kant, A., Smith, K., & Rey, A. (2020). Non‐adjacent dependency learning in humans and other animals. Topics in Cognitive Science, 12(3), 843-858. doi:10.1111/tops.12381.
Abstract
Learning and processing natural language requires the ability to track syntactic relationships between words and phrases in a sentence, which are often separated by intervening material. These nonadjacent dependencies can be studied using artificial grammar learning paradigms and structured sequence processing tasks. These approaches have been used to demonstrate that human adults, infants and some nonhuman animals are able to detect and learn dependencies between nonadjacent elements within a sequence. However, learning nonadjacent dependencies appears to be more cognitively demanding than detecting dependencies between adjacent elements, and only occurs in certain circumstances. In this review, we discuss different types of nonadjacent dependencies in language and in artificial grammar learning experiments, and how these differences might impact learning. We summarize different types of perceptual cues that facilitate learning, by highlighting the relationship between dependent elements bringing them closer together either physically, attentionally, or perceptually. Finally, we review artificial grammar learning experiments in human adults, infants, and nonhuman animals, and discuss how similarities and differences observed across these groups can provide insights into how language is learned across development and how these language‐related abilities might have evolved. -
Ravignani, A., & Thompson, B. (2017). A note on ‘Noam Chomsky – What kind of creatures are we? Language in Society, 46(3), 446-447. doi:10.1017/S0047404517000288.
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Ravignani, A., Honing, H., & Kotz, S. A. (2017). Editorial: The evolution of rhythm cognition: Timing in music and speech. Frontiers in Human Neuroscience, 11: 303. doi:10.3389/fnhum.2017.00303.
Abstract
This editorial serves a number of purposes. First, it aims at summarizing and discussing 33 accepted contributions to the special issue “The evolution of rhythm cognition: Timing in music and speech.” The major focus of the issue is the cognitive neuroscience of rhythm, intended as a neurobehavioral trait undergoing an evolutionary process. Second, this editorial provides the interested reader with a guide to navigate the interdisciplinary contributions to this special issue. For this purpose, we have compiled Table 1, where methods, topics, and study species are summarized and related across contributions. Third, we also briefly highlight research relevant to the evolution of rhythm that has appeared in other journals while this special issue was compiled. Altogether, this editorial constitutes a summary of rhythm research in music and speech spanning two years, from mid-2015 until mid-2017 -
Ravignani, A., & Sonnweber, R. (2017). Chimpanzees process structural isomorphisms across sensory modalities. Cognition, 161, 74-79. doi:10.1016/j.cognition.2017.01.005.
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Ravignani, A., Gross, S., Garcia, M., Rubio-Garcia, A., & De Boer, B. (2017). How small could a pup sound? The physical bases of signaling body size in harbor seals. Current Zoology, 63(4), 457-465. doi:10.1093/cz/zox026.
Abstract
Vocal communication is a crucial aspect of animal behavior. The mechanism which most mammals use to vocalize relies on three anatomical components. First, air overpressure is generated inside the lower vocal tract. Second, as the airstream goes through the glottis, sound is produced via vocal fold vibration. Third, this sound is further filtered by the geometry and length of the upper vocal tract. Evidence from mammalian anatomy and bioacoustics suggests that some of these three components may covary with an animal’s body size. The framework provided by acoustic allometry suggests that, because vocal tract length (VTL) is more strongly constrained by the growth of the body than vocal fold length (VFL), VTL generates more reliable acoustic cues to an animal’s size. This hypothesis is often tested acoustically but rarely anatomically, especially in pinnipeds. Here, we test the anatomical bases of the acoustic allometry hypothesis in harbor seal pups Phoca vitulina. We dissected and measured vocal tract, vocal folds, and other anatomical features of 15 harbor seals post-mortem. We found that, while VTL correlates with body size, VFL does not. This suggests that, while body growth puts anatomical constraints on how vocalizations are filtered by harbor seals’ vocal tract, no such constraints appear to exist on vocal folds, at least during puppyhood. It is particularly interesting to find anatomical constraints on harbor seals’ vocal tracts, the same anatomical region partially enabling pups to produce individually distinctive vocalizations. -
Ravignani, A., & Norton, P. (2017). Measuring rhythmic complexity: A primer to quantify and compare temporal structure in speech, movement, and animal vocalizations. Journal of Language Evolution, 2(1), 4-19. doi:10.1093/jole/lzx002.
Abstract
Research on the evolution of human speech and phonology benefits from the comparative approach: structural, spectral, and temporal features can be extracted and compared across species in an attempt to reconstruct the evolutionary history of human speech. Here we focus on analytical tools to measure and compare temporal structure in human speech and animal vocalizations. We introduce the reader to a range of statistical methods usable, on the one hand, to quantify rhythmic complexity in single vocalizations, and on the other hand, to compare rhythmic structure between multiple vocalizations. These methods include: time series analysis, distributional measures, variability metrics, Fourier transform, auto- and cross-correlation, phase portraits, and circular statistics. Using computer-generated data, we apply a range of techniques, walking the reader through the necessary software and its functions. We describe which techniques are most appropriate to test particular hypotheses on rhythmic structure, and provide possible interpretations of the tests. These techniques can be equally well applied to find rhythmic structure in gesture, movement, and any other behavior developing over time, when the research focus lies on its temporal structure. This introduction to quantitative techniques for rhythm and timing analysis will hopefully spur additional comparative research, and will produce comparable results across all disciplines working on the evolution of speech, ultimately advancing the field.Additional information
lzx002_Supp.docx -
Ravignani, A. (2017). Interdisciplinary debate: Agree on definitions of synchrony [Correspondence]. Nature, 545, 158. doi:10.1038/545158c.
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Ravignani, A., & Madison, G. (2017). The paradox of isochrony in the evolution of human rhythm. Frontiers in Psychology, 8: 1820. doi:10.3389/fpsyg.2017.01820.
Abstract
Isochrony is crucial to the rhythm of human music. Some neural, behavioral and anatomical traits underlying rhythm perception and production are shared with a broad range of species. These may either have a common evolutionary origin, or have evolved into similar traits under different evolutionary pressures. Other traits underlying rhythm are rare across species, only found in humans and few other animals. Isochrony, or stable periodicity, is common to most human music, but isochronous behaviors are also found in many species. It appears paradoxical that humans are particularly good at producing and perceiving isochronous patterns, although this ability does not conceivably confer any evolutionary advantage to modern humans. This article will attempt to solve this conundrum. To this end, we define the concept of isochrony from the present functional perspective of physiology, cognitive neuroscience, signal processing, and interactive behavior, and review available evidence on isochrony in the signals of humans and other animals. We then attempt to resolve the paradox of isochrony by expanding an evolutionary hypothesis about the function that isochronous behavior may have had in early hominids. Finally, we propose avenues for empirical research to examine this hypothesis and to understand the evolutionary origin of isochrony in general. -
Ravignani, A. (2017). Visualizing and interpreting rhythmic patterns using phase space plots. Music Perception, 34(5), 557-568. doi:10.1525/MP.2017.34.5.557.
Abstract
STRUCTURE IN MUSICAL RHYTHM CAN BE MEASURED using a number of analytical techniques. While some techniques—like circular statistics or grammar induction—rely on strong top-down assumptions, assumption-free techniques can only provide limited insights on higher-order rhythmic structure. I suggest that research in music perception and performance can benefit from systematically adopting phase space plots, a visualization technique originally developed in mathematical physics that overcomes the aforementioned limitations. By jointly plotting adjacent interonset intervals (IOI), the motivic rhythmic structure of musical phrases, if present, is visualized geometrically without making any a priori assumptions concerning isochrony, beat induction, or metrical hierarchies. I provide visual examples and describe how particular features of rhythmic patterns correspond to geometrical shapes in phase space plots. I argue that research on music perception and systematic musicology stands to benefit from this descriptive tool, particularly in comparative analyses of rhythm production. Phase space plots can be employed as an initial assumption-free diagnostic to find higher order structures (i.e., beyond distributional regularities) before proceeding to more specific, theory-driven analyses. -
Filippi, P., Jadoul, Y., Ravignani, A., Thompson, B., & de Boer, B. (2016). Seeking Temporal Predictability in Speech: Comparing Statistical Approaches on 18 World Languages. Frontiers in Human Neuroscience, 10: 586. doi:10.3389/fnhum.2016.00586.
Abstract
Temporal regularities in speech, such as interdependencies in the timing of speech events, are thought to scaffold early acquisition of the building blocks in speech. By providing on-line clues to the location and duration of upcoming syllables, temporal structure may aid segmentation and clustering of continuous speech into separable units. This hypothesis tacitly assumes that learners exploit predictability in the temporal structure of speech. Existing measures of speech timing tend to focus on first-order regularities among adjacent units, and are overly sensitive to idiosyncrasies in the data they describe. Here, we compare several statistical methods on a sample of 18 languages, testing whether syllable occurrence is predictable over time. Rather than looking for differences between languages, we aim to find across languages (using clearly defined acoustic, rather than orthographic, measures), temporal predictability in the speech signal which could be exploited by a language learner. First, we analyse distributional regularities using two novel techniques: a Bayesian ideal learner analysis, and a simple distributional measure. Second, we model higher-order temporal structure—regularities arising in an ordered series of syllable timings—testing the hypothesis that non-adjacent temporal structures may explain the gap between subjectively-perceived temporal regularities, and the absence of universally-accepted lower-order objective measures. Together, our analyses provide limited evidence for predictability at different time scales, though higher-order predictability is difficult to reliably infer. We conclude that temporal predictability in speech may well arise from a combination of individually weak perceptual cues at multiple structural levels, but is challenging to pinpoint.Additional information
https://www.frontiersin.org/article/10.3389/fnhum.2016.00586/full#supplementary… -
Geambaşu, A., Ravignani, A., & Levelt, C. C. (2016). Preliminary experiments on human sensitivity to rhythmic structure in a grammar with recursive self-similarity. Frontiers in Neuroscience, 10: 281. doi:10.3389/fnins.2016.00281.
Abstract
We present the first rhythm detection experiment using a Lindenmayer grammar, a self-similar recursive grammar shown previously to be learnable by adults using speech stimuli. Results show that learners were unable to correctly accept or reject grammatical and ungrammatical strings at the group level, although five (of 40) participants were able to do so with detailed instructions before the exposure phase. -
Ravignani, A., Delgado, T., & Kirby, S. (2016). Musical evolution in the lab exhibits rhythmic universals. Nature Human Behaviour, 1: 0007. doi:10.1038/s41562-016-0007.
Abstract
Music exhibits some cross-cultural similarities, despite its variety across the world. Evidence from a broad range of human cultures suggests the existence of musical universals1, here defined as strong regularities emerging across cultures above chance. In particular, humans demonstrate a general proclivity for rhythm2, although little is known about why music is particularly rhythmic and why the same structural regularities are present in rhythms around the world. We empirically investigate the mechanisms underlying musical universals for rhythm, showing how music can evolve culturally from randomness. Human participants were asked to imitate sets of randomly generated drumming sequences and their imitation attempts became the training set for the next participants in independent transmission chains. By perceiving and imitating drumming sequences from each other, participants turned initially random sequences into rhythmically structured patterns. Drumming patterns developed into rhythms that are more structured, easier to learn, distinctive for each experimental cultural tradition and characterized by all six statistical universals found among world music1; the patterns appear to be adapted to human learning, memory and cognition. We conclude that musical rhythm partially arises from the influence of human cognitive and biological biases on the process of cultural evolution. -
Ravignani, A., & Cook, P. F. (2016). The evolutionary biology of dance without frills. Current Biology, 26(19), R878-R879. doi:10.1016/j.cub.2016.07.076.
Abstract
Recently psychologists have taken up the question of whether dance is reliant on unique human adaptations, or whether it is rooted in neural and cognitive mechanisms shared with other species 1, 2. In its full cultural complexity, human dance clearly has no direct analog in animal behavior. Most definitions of dance include the consistent production of movement sequences timed to an external rhythm. While not sufficient for dance, modes of auditory-motor timing, such as synchronization and entrainment, are experimentally tractable constructs that may be analyzed and compared between species. In an effort to assess the evolutionary precursors to entrainment and social features of human dance, Laland and colleagues [2] have suggested that dance may be an incidental byproduct of adaptations supporting vocal or motor imitation — referred to here as the ‘imitation and sequencing’ hypothesis. In support of this hypothesis, Laland and colleagues rely on four convergent lines of evidence drawn from behavioral and neurobiological research on dance behavior in humans and rhythmic behavior in other animals. Here, we propose a less cognitive, more parsimonious account for the evolution of dance. Our ‘timing and interaction’ hypothesis suggests that dance is scaffolded off of broadly conserved timing mechanisms allowing both cooperative and antagonistic social coordination.Additional information
Experimental Procedures and Two Tables -
Ravignani, A., Fitch, W. T., Hanke, F. D., Heinrich, T., Hurgitsch, B., Kotz, S. A., Scharff, C., Stoeger, A. S., & de Boer, B. (2016). What pinnipeds have to say about human speech, music, and the evolution of rhythm. Frontiers in Neuroscience, 10: 274. doi:10.3389/fnins.2016.00274.
Abstract
Research on the evolution of human speech and music benefits from hypotheses and data generated in a number of disciplines. The purpose of this article is to illustrate the high relevance of pinniped research for the study of speech, musical rhythm, and their origins, bridging and complementing current research on primates and birds. We briefly discuss speech, vocal learning, and rhythm from an evolutionary and comparative perspective. We review the current state of the art on pinniped communication and behavior relevant to the evolution of human speech and music, showing interesting parallels to hypotheses on rhythmic behavior in early hominids. We suggest future research directions in terms of species to test and empirical data needed.
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