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WIREs Cogn Sci
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Combinatoriality in the vocal systems of nonhuman animals

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A key challenge in the field of human language evolution is the identification of the selective conditions that gave rise to language's generative nature. Comparative data on nonhuman animals provides a powerful tool to investigate similarities and differences among nonhuman and human communication systems and to reveal convergent evolutionary mechanisms. In this article, we provide an overview of the current evidence for combinatorial structures found in the vocal system of diverse species. We show that considerable structural diversity exits across and within species in the forms of combinatorial structures used. Based on this we suggest that a fine‐grained classification and differentiation of combinatoriality is a useful approach permitting systematic comparisons across animals. Specifically, this will help to identify factors that might promote the emergence of combinatoriality and, crucially, whether differences in combinatorial mechanisms might be driven by variations in social and ecological conditions or cognitive capacities. This article is categorized under: Cognitive Biology > Evolutionary Roots of Cognition Linguistics > Evolution of Language
Graphical illustration of combinatorial structures in nonhuman vocal systems. Each combinatorial structure shows one representative species. Shapes distinguish sound elements, colors distinguish meaning (black indicates absence of functional‐ or context‐specific meaning). Phonocoding: combination of meaningless elements into sequence that lacks functional‐ or context‐specific meaning (e.g., whales, songbirds). Multi‐element calls: combination and reuse of meaningless elements to generate context‐specific/functionally meaningful calls (e.g., chestnut‐crowned babblers). Temporal structures: meaning‐differentiating temporal variation (e.g., number of element repetitions) within a string of repeated sounds (e.g., pied babblers, Mexican free‐tailed bats). Intermediate structures: combination of meaningful calls into sequence reflecting intermediate stages experienced by the caller (e.g., wedge‐capped capuchins, gorillas). Segmental concatenations: concatenation of invariable, stand‐alone segment (generally identity‐encoding) with variable, bound segment (generally motivation/behavior‐encoding) (e.g., banded mongooses, Diana monkeys). Meaning‐modifying structures: combination of individually meaningful (variable) call with (invariable) meaning‐modifying affix (e.g., Campbell's monkeys). Meaning‐derived call combinations: combination of meaningful calls into sequence with derived meaning (e.g., Japanese tits, pied babblers). Idiomatic structures: combination of meaningful calls into sequence with unrelated meaning (e.g., putty‐nosed monkeys). Stochastic structures: sequence whose meaning is encoded by proportional sound/call contribution at particular parts of the sequence (e.g., black‐fronted titi monkeys). Animal drawings were done by Zinaida Bogdanova
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Linguistics > Evolution of Language
Cognitive Biology > Evolutionary Roots of Cognition

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