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      Compositionality in animals and humans

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          Abstract

          A key step in understanding the evolution of human language involves unravelling the origins of language’s syntactic structure. One approach seeks to reduce the core of syntax in humans to a single principle of recursive combination, merge, for which there is no evidence in other species. We argue for an alternative approach. We review evidence that beneath the staggering complexity of human syntax, there is an extensive layer of nonproductive, nonhierarchical syntax that can be fruitfully compared to animal call combinations. This is the essential groundwork that must be explored and integrated before we can elucidate, with sufficient precision, what exactly made it possible for human language to explode its syntactic capacity, transitioning from simple nonproductive combinations to the unrivalled complexity that we now have.

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          Formulaic Language and the Lexicon

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            Language evolution: semantic combinations in primate calls.

            Syntax sets human language apart from other natural communication systems, although its evolutionary origins are obscure. Here we show that free-ranging putty-nosed monkeys combine two vocalizations into different call sequences that are linked to specific external events, such as the presence of a predator and the imminent movement of the group. Our findings indicate that non-human primates can combine calls into higher-order sequences that have a particular meaning.
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              Campbell's monkeys concatenate vocalizations into context-specific call sequences.

              Primate vocal behavior is often considered irrelevant in modeling human language evolution, mainly because of the caller's limited vocal control and apparent lack of intentional signaling. Here, we present the results of a long-term study on Campbell's monkeys, which has revealed an unrivaled degree of vocal complexity. Adult males produced six different loud call types, which they combined into various sequences in highly context-specific ways. We found stereotyped sequences that were strongly associated with cohesion and travel, falling trees, neighboring groups, nonpredatory animals, unspecific predatory threat, and specific predator classes. Within the responses to predators, we found that crowned eagles triggered four and leopards three different sequences, depending on how the caller learned about their presence. Callers followed a number of principles when concatenating sequences, such as nonrandom transition probabilities of call types, addition of specific calls into an existing sequence to form a different one, or recombination of two sequences to form a third one. We conclude that these primates have overcome some of the constraints of limited vocal control by combinatorial organization. As the different sequences were so tightly linked to specific external events, the Campbell's monkey call system may be the most complex example of 'proto-syntax' in animal communication known to date.
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                Author and article information

                Journal
                PLoS Biol
                PLoS Biol
                plos
                plosbiol
                PLoS Biology
                Public Library of Science (San Francisco, CA USA )
                1544-9173
                1545-7885
                15 August 2018
                August 2018
                15 August 2018
                : 16
                : 8
                : e2006425
                Affiliations
                [1 ] Department of Comparative Linguistics, University of Zurich, Zurich, Switzerland
                [2 ] Department of Psychology, University of Warwick, Coventry, United Kingdom
                [3 ] Psycholinguistics Laboratory, Department of Comparative Linguistics, University of Zurich, Zurich, Switzerland
                [4 ] Institute of Biology, University of Neuchatel, Neuchatel, Switzerland
                [5 ] School of Psychology and Neuroscience, University of St Andrews, St Andrews, United Kingdom
                Author notes

                The authors have declared that no competing interests exist.

                ‡ These authors share first authorship on this work.

                Author information
                http://orcid.org/0000-0003-1504-1801
                Article
                pbio.2006425
                10.1371/journal.pbio.2006425
                6093600
                30110319
                dd3e0cd7-d0bb-493b-9333-37829c4e53e4
                © 2018 Townsend et al

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                Page count
                Figures: 1, Tables: 0, Pages: 7
                Funding
                University of Zurich Research Priority Program (grant number URPP, Evolution in Action; URPP U-702-06). Received by SWT and BB. Swiss National Science Foundation (grant number SWT: PP003_163860; SE: PP003_163860; P1ZHP3_151648; KZ grant: 31003A_166458). Received by SWT, SE and KZ. European Research Council under the European Union’s 7th Framework Programme (grant number FP7/2007-2013/ ERC grant agreement no (615988)). Received by SS. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
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